WESTERN SANTA BARBARA CHANNEL 

- FINAL REPORT I 

Volume 5 of 14 

The Annelida Part 2 - Polyehaeta: Ph yllodoada (Syllidae and Scale-Bearing 

Fami ties), ~hphinamida, and Eunicida 
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I OCS Study

i MMS 95-0016

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| TAXONOMIC ATLAS OF THE BENTHIC FAUNA OF

THE SANTA MARIA BASIN AND

i WESTERN SANTA BARBARA CHANNEL

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'i FINAL REPORT Volume 5 of 14

I The Annelida Part 2 m Polychaeta: Phyllodocida (Syllidae and Scale-Bearing

Families), Amphinomida, and Eunicida

i Edited by:

i JamBdgitte es A. Hilbig Blake

Paul H. Scott

I Submitted by:

Science Applications International Corporation

I San 10260Diego, CampCalifornia us Point Drive

92121

For:

I U.S. Department of the Interior

Minerals Management Service

i Pacific OCS Region 770 Paseo Camarillo

Camarillo, California 93010

i Under Contract No. 14-35-0001-30484

i Minerals Pa Management Service cific OCS Region November 1995

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DISCLAIMER

This report has been reviewedby the PacificOuter ContinentalShelfRegion,MineralsMangement i

Service, U.S. Department of the Interior and approved for publication. The opinions, findings,

conc necessarily lusionsreflect or recothemmviewendof ations the eMinerals xpressedManagement in this reportSearervicthose e. Mention of the ofauthors, trade names and dodoes not i

not consitiute endorsement or recommendation for use. This report has not been edited for

conformity with Minerals Management Service editorial standards, i

TAXONOMIC DISCLAIMER i

This report is not deemed nor intended to be a valid publicationfor the naming of new taxa as

stipulated in the International Code of Zoological Nomenclature, Article 8b. i

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i PROJECT ORGANIZATION

I PROGRAM MANAGER

I DR. Aa,mRmVL. L/ssym_ Science Applications International Corporation

10260 Campus Point Drive

I San Diego, California 92121

I DEPUTY PROGRAM MANAGER

i DR. JAMES A. BLAKE ENSR Consulting and Engineering

89 Water Street

I Woods Hole, Massachusetts 02543

I PROJECT TECHNICAL OFFICER

I MR.FRAr_ J. MANACO,

U.S. Department of the Interior

i Minerals Management Service, Pacific OCS Region

770 Paseo Camarillo

I Camarillo, California 93010

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CONTRIBIYrING AUTHORS

I DR. JAMES A. BLAKE

DR. BRmrrrE HILBIG

Ms. LESLIEH. HARRIS

DR. JERRY D. KtmEYOV

i MR. R. EUGENE RuFF

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' I iii
ACCESS NUMBER 30484.5 

TECHNICAL SUMMARY 

STUDY TITLE: Taxonomic Atlas of the Santa Maria Basin and Western Santa Barbara Channel. 

REPORT TITLE: Taxonomic Atlas of the Santa Maria Basin and Western Santa Barbara Channel. Volume 

5. The Annelida Part 2 - Polychaeta: Phyllodicida (Syllidae and Scale-Bearing Families), Amphinomida, 

and Eunicida. 

CONTRACT NUMBER: MMS Contract No. 14-35-0001 -30484. 

SPONSORING OCS REGION: Pacific. 

APPLICABLE PLANNING AREA: Southern California. 

FISCAL YEARS OF PROJECT FUNDING: 1990; 1991 ; 1992; 1993; 1994; 1995. 

COMPLETION DATE OF REPORT November 1995.Taxonomic Atlas of the Benthic Fauna of the Santa 

Maria Basin and Western Santa Barbara Channel. Volume 5. The Annelida Part 2 - Polychaeta: Phyllodicida 

(Syllidae and Scale- Bearing Families), Amphinomida, and Eunicida. 

COSTS: FY 1989, $229,013; FY 1990, $1 50,000; FY 1991 $1 50,000; FY 1 993 $1 50,000; FY 1995 $45,371. 

CUMULATIVE PROJECT COST $724,384 

PROJECT MANAGER: Andrew L. Lissner. 

AFFILIATION: Science Applications International Corporation. 

ADDRESS: 10260 Campus Point Drive, San Diego, California 92121. 

PRINCIPAL INVESTIGATORS': James A. Blake, Andrew L. Lissner. 

KEY WORDS: Marine invertebrates, Annelida, Syllidae, Aphroditidae, Polynoidae, Acoetidae, Pholoidae, 

Sigalionidae, Amphinomidae, Euphrosinidae, Onuphidae, Eunicidae, Lumbrineridae, Oenonidae, Dowilleidae, 

California, Santa Maria Basin, Santa Barbara Channel, continental shelf. 

BACKGROUND: The Taxonomic Atlas of the Santa Maria Basin and western Santa Barbara Channel is an 

extension of the benthic reconnaissance (Phase I) and monitoring programs (Phase II) that were conducted 

by the MMS since 1983. The organisms that were collected as part of those programs provide the material 

on which the Atlas is developed. In order to fully document the fauna collected by those programs, a series 

of 14 volumes will be prepared that provide keys, descriptions, and illustrations of the benthic fauna of the 

hard and soft substrate environments. A team of 40 experts on the fauna has been assembled to carry out 

this work and their contributions are distributed among the 14 volumes. 

OBJECTIVES: The objectives of Volume 5 are to continue coverage of the Polychaeta that was initiated in 

Volume 4. This volume concludes treatment of the so-called "errant" families. The following 13 families are 

treated in this volume: Syllidae, Aphroditidae, Polynoidae, Acoetidae, Pholoidae, Sigalionidae, 

Amphinomidae, Euphrosinidae, Onuphidae, Eunicidae, Lurnbrineridae, Oenonidae, Dorvilleidae. 

DESCRIPTION: Volume 5 concludes treatment of the so-called "erranl' families and includes the Syllidae, 

5 scale-bearing families (Aphroditidae, Polynoidae, Acoetidae, Pholoidae, and Sigalionidae), Ampt~inomidae, 

Euphrosinidae, and the Eunicida (Eunicidae, Onuphidae, Lumbrineridae, Oenonidae, and Dorvilleidae). 

The 13 chapters are organized into sections that include the morphology, taxonomic history, biology, keys to 

species, and descriptions of genera and species. Each species is fully illustrated with relevant characteristics 
ACCESS NUMBER 30484.5 

labeled and identified. 

SIGNIFICANT CONCLUSIONS: The keys, detailed descriptions, and illustrations to the 13 families treated 

in this volume represent a major contribution to polychaete systematics for the eastern Pacific. Many 

poorly known species are newly defined with new illustrations. The chapter on Syllidae defines a total of 34 

species in the subfamilies Exogoninae (10, 6 new), Eusyllinae (12, 5 new), and Syllinae (12, 1 new, 3 

provisional). Among the Polynoidae 22 species in 13 genera are included, with one new genus and species 

described. New species of Pholoe, Onuphis, Parophryrotrocha, Ninoe, and Drilonereis are described. 

STUDY RESULTS: A total of 11 5 species in 57 genera are treated. One genus in the Polynoidae is new to 

science. Eighteen species are new to science as follows: Syllidae (12), Polynoidae (I), Pholoidae (I), 

Onuphidae (I), Lumbrineridae (I), Oenonidae (I), and Dorvilleidae (1). Three additional species are 

provisional new taxa, to be named in a later publication. The chapter on the Syllidae represents a major 

treatment of eastern Pacific genera and species of the subfamities Exogoninae, Eusyllinae, and Syllinae. 

The species descriptions, especially of the new and provisional taxa are a major contribution. The 22 

described species of Polynoidae include several poorly known taxa that are illustrated in detail for the first 

time. One new genus and species is described. Previous California records of Lepidonotus sguamatus are 

referred to L. spicuius. Keys are provided to all species from the study area and in some cases, to species 

from adjacent areas. The treatment of other scale-bearing families is mostly limited to the material available 

from the MMS collections. For the Aphroditidae only juveniles are available from the MMS collections 

precluding any extensive review of California species. Sigalionidae and Acoetidae include previously 

known species, with updated descriptions and keys provided. For the Pholoidae, a new species of Pholoe 

is described. The amphinomid, Chloeia pinnata was found to have unusual bifurcate notosetae in posterior 

segments. These setae and other features of this common species are carefully illustrated; taxonomic 

implications of these findings are discussed. In the Euphrosinidae, 2 species were identified and described. 

Among the euniciform families, new species of Onuphis, Ninoe, Drilonereis, and Parophryotrocha are 

described. In the Oenonidae, Arabella pecfinata and A. protomutans are newly reported for California. 

Keys are provided for all species treated. 

STUDY PRODUCT: Blake, J.A. Hilbig, B., and P.H. Scott (Editors). 1995. Taxonomic Atlas of the Benthic 

Fauna of the Santa Maria Basin and Westem Santa Barbara Channel. Volume 5. The Annelida Part 2- 

Polychaeta: Phyllodocida (Syllidae and Scale-Bearing families), Amphinornidae, and Eunicida. A final 

report prepared by Science Applications International Corporation, for the U.S. Department of the Interior, 

Minerals Management Service, Pacific OCS Region, Camarilla, CA. OCS Study MMS 95-01 6. Contract 

NO. 14-35-0001 -30484. 

'P.lls affiliation may be different than that listed for Project Manager(s). 

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ACCESS NUMBER 30484.5

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Southern California PlanningArea Indicatingthe TaxonomicAtlas Study Region. n

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DOCUMENTATION 1. _rr NO. MMS 95-0016 :2" 3. Reci_s ,_¢==,=t_No.

I PAGE 4. Title eitd Subtitle L Report Date

Taxonomic Atlas of the Behthic Fauna of the Santa Maria Basin and Western Santa November 1995

Barbara Channel. Volume 5. The Annelida Part 2 m Polychaeta: Phyllodicida (Syllidae 6.

I 7.andAuthor(s) Scale-Bearin 9 Families), Amphinomida, and Eunicida. 8. Performini OrlPnJzation Rept, No.

James A. Blake, Brigitte Hilbig and Paul H. Scott (editors)

I 9. Peffm'minl Organization Name end Address 10. Projectrresk/Work Unit No.

Science AppJications international Corporation z:. co..,_c) o, G,I,t(_) No.

10260 Campus Point Drive, San Diego, California 92121

i (c) 14-35-0001-30484 (G)

|2. Sponsoring Oqlmnl:lr=Ltlon Name end Address 'J3. Type of Report & P_.riod Cov_re6

i MPacific ineralOCS ManaRegion, gement Service, U. S. Department of the Interior, Final 5 of 14 volumes, 770 Paseo Camarillo 9/89 to 11/95

Camaritlo, California 93010 14.

I ....

IS. Sul_,el'nlntary Notls

Contributing authors: James Blake, Brigitte Hilbig, Jerry Kudenov, Leslie Harris, R. Eugene Ruff

i |8. Al_l=rlCt {LIm_.: 200 words)

Volume 5 is the second annelid volume, and continues coverage of the Polychaeta that began with Volume 4. This

I volume concludes treatment of the so-called "errant" families. The following 13 families are treated in this volume: Syllidae, Aphroditidae, Porynoidae, Acoetiodae, Pholoidae, Sigalionidae, Amphinomidae, Euphrosinidae, Onuphidae,

Eunicidae, Lumbrineridae, Oenonidae, and Dorvilleidae. A total of 115 species in 57 genera are treated. One genus

in the Polynoidae is new to science. Eighteen species are new to science as follows: Syllidae (12), Polynoidae (1),

I Pholoidae (1), Onuphidae (1), Lumbrineridae (1), Oenonidae (1), and Dorvilleidae (1). Three additional species of Syllidae are provisional new taxa, to be named later. The chapter on the Syllidae represents a major treatment of

eastern Pacific genera and species of the subfamilies Exogoninae, Eusyllinae, and Syllinae. The species descriptions,

especially of the new and provisional taxa are a major contribution. The 22 described species of Polynoidae include

I several Previouspoorly California known taxa that are illustrated in detail for the first time. One new genus and species is described. records of Lepidonotus squamatus are referred to L. spicufus. Keys are provided to all species

from the study area and in some cases, to species from adjacent areas. The treatment of other scale-bearing

families is mostly limited to the material available from the MMS collections. For the Aphroditidae, only juveniles are

I available include previously from the MMS knowncollections species, with precluding updatedanydescriptions extensive review and keys. of California For the Pholoidae, species. Sigalionidae a new speciesandofAcoetidae Phofoe is

described. The amphinomid, Chfoeia pinnate was found to have unusual bifurcate notosetae in posterior segments.

These setae and other features of this common species are carefully illustrated; taxonomic implications of these

I families, findings anewre dspecies iscussed.of Onuphis, In the Euph Ninoe, rosiDdfonereis nidae, two, spandeciParoph es wereryotrocha identifieare d anddescribed. describedIn. theAmOeonngonida thee,eunArabelfa iciform

pectinata and A. protomutans are newly reported for California. Keys are provided for all species treated.

I Mr. tl_¢um_w_. Amblysls s. Ossch_:ters

Marine invertebrates, Annelida Polychaeta, Syllidae, Aphroditidae, Polynoidae, Acoetiodae, Pholoidae, Sigalionidae,

i Amphinomidae, Euphrosinidae, Onuphidae, Eunicidae, Lumbrineridae, Oenonidae, and Dorvilleidae, taxonomy,

California, Santa Maria Basin, Santa Barbara Channel, continental shelf.

b- Idantlk_slOpon-Ended Terms

I

Taxonomic Atlas, keys to species, faunal guide, marine ecology, Annelida, Polychaeta, marine benthos.

i ¢. COSATI FIsM/Gmup

|L Availabtllty W.&tement 19. Security Class {This Repo4't) 21. No. af PaBes

I Unlimited Unclassified 372

20. Security Class (This Pile} 22, Price

See ANSI..-Z39.18) See fnstruct/ons on Reverse OPTIONAL FORM 272 (4-71)

i (Fon_ulrly I_rl s.-35)

Deportment of Com_eccl
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Table of Contents n

1. FAMILYSYLLIDAEGRUBE, 1850 (by Jerry D. Kudenov and Leslie H. Harris) m

Introduction ...................................................................................................................................... 1 •

Morphology ..................................................................................................................................... 1 II

Systematic History ........................................................................................................................... 2

Principal Diagnostic Traits .............................................................................................................. 2 •

Subfamily Autolytinae ........................................................................................................ 3 II

Subfamily Exogoninae ........................................................................................................ 3

Subfamily Eusyllinae .......................................................................................................... 3 II

Subfamily Syllinae .............................................................................................................. 3 l

Biology ............................................................................................................................................. 5

Key to the Subfamilies and Genera of the Syllidae .......................................................................... 5 II

List of Syltid Species ....................................................................................................................... 7 |

Subfamily Exogoninae ........................................................................................................ 7

Subfamily Eusyllinae .......................................................................................................... 8 ann

Subfamily Syllinae .............................................................................................................. 8

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Description of Species ..................................................................................................................... 8

Key to Species of Brania From the Santa Mafia Basin ....................................... 10 i

Brania brevipharyngea Banse, 1972 ...................................................... 10 |

Brania cah'forniensis Kudenov and Harris, new species .................... 12

Key to the Subgenera of Exogone ........................................................................ 14 Ii

Key to Known Species of the Subgenus Exogone (Exogone) from the Northeast l

Pacific .................................................................................................................. 15

Exogone (Exogone) lourei Berkeley and Berkeley, 1938 ....................... 15 ill

gxogone (Exogone) dwisula Kudenov and Harris, new species ........ 17 n

Key to the Known Species of the Subgenus Exogone (Parexogone) from the Pacific

Coast .................................................................................................................... 20

Exogone (Parexogone) molesta (Banse, 1972) ..................................... 20 I[

Exogone (Parexogone) acutipalpa Kudenov and Harris, new species..

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Exogone (Parexogone) breviseta Kudenov and Harris, new species. 24 II

Key to Selected Species of the Subgenus Exogone (Sylline) ............................... 27

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Key to Species of Sphaerosyllis from the Northeast Pacific ................................ 28

Sphaerosyllis californiensis Hartman, 1966 .......................................... 28 n

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SphaerosyUis bilineata Kudenov and Harris, new species ................. 31 !11

Sphaeresyllis ranunculus Kudenov and Harris, new species ............ 33

Subfamily Eusyllinae ........................................................................................................ 35 I

Key to the Species of Dioplosyllis from California ............................................. 36

Dioplosyllis lagunae (Hartman, 1961), new combination .................... 36

Dioplosyllis tridentata Kudenov and Harris, new species .................. 38 l

Key to Species of Eusyllis from California ......................................................... 40

Eusyllis blomstrandi Malmgren, 1867 ................................................... 41

Eusyllis habei Imajima, 1966 ................................................................. 43 •

Eusyllis longicirrata Imajima, I966 ...................................................... 45

Key to Known Species of Odontosyllis from the Northeast Pacific .................... 47

Odontosyllisphosphorea Moore, 1909 .................................................. 47 n
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I Odontosyllisfragilis Kudenov and Harris, new species ..................... 49

Opisthodonta mitcheUi Kudenov and Harris, new species ................. 51

i Key to the Species of Pionosyllis from California ............................................... 54 PionosylIis magnifica Moore, t 906 ....................................................... 54

PionosyUis articulata Kudenov and Harris, new species ................... 55

i Key to Known Species of Syllides from the Northeast Pacific ............................ 58 SyIlides reishi Dorsey, 1978 ................................................................... 59

Syllides mikeli Kudenov and Harris, new species ............................... 61

Subfamily Syllinae ............................................................................................................ 63

I Key to the MMS Species of Ehlersia from the Northeast Pacific ....................... 64

Ehlersia heterochaeta (Moore, 1909) .................................................... 64

EhIersia hyperioni (Dorsey and Phillips, 1987) ..................................... 66

I Eurysyllis spicum Kudenov and Harris, new species ......................... 68

Geminosyllis ohma (Imajirna and Hartman, 1964) ................................ 71

Syllis spongiphila Verrill, 1885 .............................................................. 73

I to the of from California 75

Key Species Trypanosyllis (Trypanosyllis)

Trypanosyllis (Trypanosyllis) coeliaca nipponica Imajima and Hartman,

1964 ........................................................................................................ 76

i Trypanosyllis (Trypanosyllis) A 77

sp.

Trypanosyllis (Trypanosyllis) sp. B ....................................................... 78

Trypanosyllis (Trypanedenta) sp.A ........................................................ 80

I Key to Species of TyposylIis from California 82 Typosyllis bella Chamberlin, 1919 ......................................................... 85

Typosyllis hyalina (Grube, 1863) ........................................................... 87

I ALiterature cknowledgements Cited 89 .............................................................................................................................. 89

I 2. FAMILYAPIIRODITIDAEMALMGREN,1867 CoyJames A. Blake)

Introduction.................................................................................................................................... 99

I Morphology 99 Biology ........................................................................................................................................... 99

Taxonomic History ...................................................................................................................... 101

I Key to the Genera of Aphroditidae from California ..................................................................... 101 Descriptions of Species ................................................................................................................ 102

Aphroditaparva Moore, 1905 .............................................................. 102

I Literature Cited ............................................................................................................................ 103

3. FAMILYPOLYNOIDAEMALMGREN,1867 CoyR. Eugene Ruff)

I Introduction .................................................................................................................................. 105

Morphology ................................................................................................................................. 105

I Taxonomic History 107 Distribution and Biological Notes ................................................................................................ 108

Key to the Species of Polynoidae ................................................................................................. 109

I Description of Species ................................................................................................................. 112

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Arctonoefragilis (Baird, 1863) ............................................................ 113 m

Arctonoe pulchra (Johnson, 1897) ....................................................... 115

Arctonoe vittata (Grube, 1855) ............................................................ 116 m

Bylgides macrolepidus (Moore, 1905) ................................................. 119 l

Eucranta anoculata (Moore, 1910) ..................................................... 121

Eunoe senta (Moore, 1902) ................................................................. 123 1

Gaudichaudius iphionelloides (Johnson, 1901) ................................... 126 m

Harmothoe fragilis Moore, 1910 ......................................................... 128

Harmothoe hirsuta Johnson, 1897 ....................................................... 130 I

Harmothoe imbricata (Linnaeus, 1767) ............................................... 132 l

Harmothoe multisetosa (Moore, 1902) ............................................... 134

Hesperonoe laevis Hartman, t961 ....................................................... 136 I

Lepidasthenia berkeleyae Pettibone, 1948 ........................................... 138 l

Lepidasthenia longicirrata Berkeley, 1923 .......................................... 140

Lepidonotus spiculus (Treadwell, 1906) new combination ................ 142 m

Malmgreniella baschi Pettibone, 1993 ................................................ 145 |

MaImgreniella macginitiei Pettibone, 1993 ........................................ 147

Malmgreniella nigralba (Berkeley, 1923) ........................................... 149 m

Malmgreniella scriptoria (Moore, t910) ............................................ 151 m

Subadyte mexicana Fauchald, 1972 .................................................... 153

m

Tenoniapriops (Hartman, 1961) ......................................................... 156 m

Ysideria hastata Ruff, new species ..................................................... 158 m

Acknowledgements ...................................................................................................................... 161

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LiteratureCited ............................................................................................................................ 162 l

4. FAMILYACOETIDAEKINBERG, 1865 (by James A. Blake) m

Introduction .................................................................................................................................. 167 mmm

Morphology ................................................................................................................................. 167

Biology ......................................................................................................................................... 168 m

Taxonomic History ...................................................................................................................... 168 m

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Key to the Genera and Species From California .......................................................................... 169

Descriptions of Species ................................................................................................................ 169 Illl

Acoetes pacifica (Treadwell, 1914) ...................................................... 170 m

Literature Cited ............................................................................................................................ 173 m

5. FAM_Y PHOLOIDAEKINBERG, 1858 (by James A. Blake) m

Introduction .................................................................................................................................. 175 m

Morphology ................................................................................................................................. 175 1

Taxonomic History ...................................................................................................................... 175

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Biology ......................................................................................................................................... 176 m

Key to the Genera and Species of Pholoidae ................................................................................ 178 1

Descriptions of Species ................................................................................................................ 178 I

Pholoe glabra Hartman, 1961 .............................................................. 179 1

Pholoe courtneyae Blake, new species ............................................... 180 m

Pholoides asperus (Johnson, 1897) ...................................................... 185

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Literature Cited ............................................................................................................................ 186

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I 6. FAMILYSIGALIONIDAEKINBERG, 1856 (by James A. Blake)

I Introduction Morphology .................................................................................................................................. ................................................................................................................................. 189 189

Taxonomic History ...................................................................................................................... 190

I Biology Key to the Genera and Species from California ........................................................................... 190 190

Descriptions of Species ................................................................................................................ 192

I Sthenelaisfusca Sthenelais berlceIeyi JohnsoPettibone, n, 1897 1971 ............................................................ .................................................... 192 194

Sthenelais verruculosa Johnson, 1897 ................................................. 196

i Sthenelais tertiaglabra Moore, 1910 ................................................... 198 Sthenelanella uniformis Moore, 1910 .................................................. 200

Sigalion spinosus (I-lartman, 1939) ...................................................... 202

i Literature Cited ............................................................................................................................ 204

7. FAMILYA2_ItINOIVlIDAELA__VaC, K, 1818 (by Jerry D. Kudenov)

I Introduction .................................................................................................................................. 207

Morphology ................................................................................................................................. 207

i Taxonomic Biology ......................................................................................................................................... History ...................................................................................................................... 208 209

Description of Species ................................................................................................................. 209

i Chloeia pinnata Moore, 1911 .............................................................. 209 Literature Cited ............................................................................................................................ 213

i 8. FAMa1LYEt_HROSImDAE WmL_, 1851 (by Jerry D. Kudenov)

Introduction .................................................................................................................................. 217

i Morph Taxonomic ologyHistory ................................................................................................................................. 217 ...................................................................................................................... 218

Biology ......................................................................................................................................... 219

i Key to the Species of Euphrosinidae ............................................................................................ 219 Description of Species ................................................................................................................. 220

: Euphrosine arctia Johnson, 1897 ......................................................... 220

i m Euphrosine bicirrata Moore, 1905 ....................................................... 223

[ l Literature Cited ............................................................................................................................ 227

9. FAMILYONUPtlIDAEK.INBERG, 1865 (by Brigitte Hilbig) !

Introduction .................................................................................................................................. 229

i Morphology ................................................................................................................................. 229 Taxonomic History ...................................................................................................................... 231

Distribution and Biological Notes ................................................................................................ 231

Key to the Ormphidae .................................................................................................................. 233

I Description Species ................................................................................................................. 235

of

Kinbergonuphis vexilIaria (Moore, 1911) ............................................ 235

i Mooreonuphis exigua (Shisko, 1981) .................................................. 238
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Mooreonuphis nebulosa (Moore, 1911) ............................................... 240 I

Mooreonuphis segmentispadix (Shisko, 1981) .................................... 242

Nothria occidentalis Fauchald, 1968 ................................................... 244 i

Onuphis affinis Hilbig, new species ................................................... 246 II

Onuphis elegans (Johnson, 1901) ........................................................ 249

Onuphis geophiliformis (Moore, 1903) ................................................ 251 •

Onuphis iridescens (Johnson, 1901) .................................................... 253 II

Onuphis sp. "intermediates". ................................................................ 255

Paradiopatra parva (Moore, 1911) ...................................................... 257 a

Rhamphobrachium longisetosum Berkeley and Berkeley, 1938 .......... 259 II

Literature Cited ............................................................................................................................ 261

10. FAMILYEtrNICmAE SAVmNV, 1818 (by Brigitte Hilbig) I

Introduction .................................................................................................................................. 263 am

Morphology |................................................................................................................................. 263

Taxonomic History ...................................................................................................................... 264

Distribution and Biological Notes ................................................................................................ 264 I!

Key to the Eunicidae .................................................................................................................... 265 |

Description of Species ................................................................................................................. 266

Eunice americana Hartman, 1944 ....................................................... 267 •

Eunice multicylindri Shisko, 1981 ....................................................... 269 ii

Eunice multipectinata Moore, 1911 ..................................................... 269

Eunice vittatopsis Fauchald, 1970 ....................................................... 273 am

Marphysa conferta Moore, 1911 .......................................................... 275 |

Literature Cited ............................................................................................................................ 277

11. F_tn.v LtWmRINERmAE MAI_GR_, 1867 (by Brigitte Hilbig) I

Introduction .................................................................................................................................. 279 I

Morphology ................................................................................................................................. 279

Taxonomic History ...................................................................................................................... 280

Distribution and Biological Notes ................................................................................................ 280 Im

Key to the Lumbrineridae ............................................................................................................ 281 !1

Description of Species ................................................................................................................. 284

Eranno bicirrata (Treadwell, 1929), new combination ...................... 284 Im

Eranno lagunae (Fauchald, 1970) ....................................................... 286 II

Lumbrineris californiensis Hartman, 1944 .......................................... 288

Lumbrineris cruzensis Hartman, 1944 ................................................. 290 m

Lumbrineris index (Moore, 1911) ........................................................ 292 •

mm

Lumbrineris inflata Moore, 1911 ......................................................... 294

Lumbrinerisjaponica (Marenzeller, 1879) .......................................... 296

Lumbrineris latreilli Audouin and Milne-Edwards, 1834 .................... 298 •

Lumbrineris limicola Hartman, 1944 ................................................... 300 I

Ninoe gemmea Moore, 1911 ................................................................ 302 /

Ninoe palmata Moore, 1903 ................................................................ 304 B

Ninoe tridentata HJlbig, new species .................................................. 306 I

Ninoe sp. C ........................................................................................... 306

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I Ninoe sp. D ........................................................................................... 308

Scoletoma tetraura (Schmarda, 1861) ................................................. 309

i Literature Cited ............................................................................................................................ 311

12. FAMILYOENONIDAEKINBERG, 1865 (by Brigitte Hilbig)

I Introduction .................................................................................................................................. 315

Morphology ................................................................................................................................. 315

n Taxonomi Distributionc His andtoryBiological ...................................................................................................................... Notes ................................................................................................ 316 317

Key to the Oenonidae ................................................................................................................... 317

n Description of Species ................................................................................................................. Arabella 319 (Arabella) iricolor (Montagu, 1804) ..................................... 320

Arabella (ArabeUa)pectinata Fauchald, 1970 .................................... 322

i Arabella Arabella (Arabella) protomutans Orensanz, 1990 ............................... 324 (Arabella)semimaculata (Moore, 1911) ............................... 326

Drilonereisfalcata Moore, 1911 .......................................................... 328

i Drilonereis longa Webster, 1879 ......................................................... 330 Drilonereis mexicana Fauchald, 1970 ................................................. 332

Drilonereis nuda Moore, 1909 ............................................................. 334

i Drilonereis spectabilis Hilbig, new species ........................................ 336 LiteratureCited ............................................................................................................................ 338

13. FAMILYDORVILLEIDAECHAMBERHN,1919 (by Brigitte Hilbig)

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Introduction .................................................................................................................................. 341

i Morphology ................................................................................................................................. 341 Taxonomic History ...................................................................................................................... 342

Distribution and Biological Notes ................................................................................................ 343

Key to the Dorvilleidae ................................................................................................................ 344

I Description of Species ................................................................................................................. 349

Dorvillea (Schistomeringos) annuIata (Moore, 1906) ......................... 350

Do rviUea (Schistomeringos) longicornis (Ehlers, 1901) ..................... 352

Parophryotrocha brevicapitis Hilbig, new species .............................

355

Parougia batia (Jumars, 1974), new combination .............................. 357

Pettiboneia brevipalpa Hilbig and Ruff, 1990 ..................................... 359

I Literature Cited 361

APPE_tx A ................................................................................................................................. 365

I APPENDIXB ................................................................................................................................. 367

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List of Figures i

Figure 1.1. Brania brevipharyngea: A, anterior end, dorsal view, setae schematic; B, parapodium 16, l

posterior view; C, inferiormost compound falciger; D, superior dorsal simple seta; E, acicula. I

(A-E, redrawn from Banse, 1972) .................................................................................... 11 i

I

Figure 1.2. Brania californiensis Kudenov and Harris, new species : A, anterior end, dorsal view; B,

parapodium 6; C, superior compound falciger, setiger 6; D, median compound falciger, setiger i

6; E, dorsal simple seta, setiger 6; F, acicula, setiger 6. (A-F, originals by JDK) ............ 12 I

Figure t.3. Exogone (Exogone) lourei: A, anterior end, dorsal view; B, dorsal simple seta from midbody

region; C, superior spiniger from midbody region; D, compound spiniger, setiger 2; E, inferior i

compound falciger, with detail of blade, setiger 2;F, ventral simple seta. (A-E after Uebelacker, II

1984) ................................................................................................................................. 17

Figure 1.4. Exogone (Exogone) dwisula Kudenov and Harris, new species: A, anterior segments, dorsal H

view; B, right parapodium, setiger 2, ventral view; C, right parapodium, setiger 9, posterior

view; D, compound falciger with can opener-like blades, setiger 1; E, superior compound •

falciger with can opener-like blades, setiger 2; F, compound falciger, setiger 5; G, dorsal |

simple seta, setiger 1;H-I, dorsal simple setae: H, setiger 6, ventral view; I, setiger 17, lateral

view; J, acicula, setiger 1; K, acicula, setiger 17. (A-K, originals by JDK) ..................... 18 i

Figure 1.5. Exogone (Parexogone) molesta: A, anterior end, dorsal view slightly from the right side; B,

parapodium 12; C, falciger with long blade from setiger 4; D, falciger with medium long

blade from setiger 16; E, falciger with short blade from setiger 4; F-G, dorsal simple seta I

from two positions from setiger 8; H, same, far posterior segment; I, ventral simple seta from

posterior setiger; J, acicula from setiger 16; K, parapodium 1, dorsal view; L, ventral simple

seta, far posterior setiger. (A-H, redrawn from Banse, 1972; I-L, originals by JDK) ...... 21 H

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Figure 1.6. Exogone (Parexogone) acutipalpa Kudenov and Harris, new species: A, anterior end, dorsal

view; B, same, dorsal view; C, right parapodium 1, dorsal view; D, left parapodium 16,

posterior view; E-H, compound falcigers from setiger 2: E, superior seta with long blade; F, Ii

same, with shorter blade; G, lower intermediate seta; H, inferior seta; I-L, compound falcigers

from setiger 12: I, superior seta with long blade; J, intermediate seta; K, low intermediate

seta; L, inferior seta; M-P, compound falcigers from setiger 23: M, superior seta; N, upper II

intermediate seta; O, lower intermediate seta; P,inferior seta; Q-R, dorsal simple setae: Q,

i

setiger 1;R, setiger 23; S, ventral simple seta from setiger 35; T-W, aciculae: T, setiger 1;U,

setiger 12; V, setiger 9; W, setiger 30. (A-W, originals by JDK) ...................................... 23 1

Figure 1.7. Exogone (Parexogone) breviseta Kudenov and Harris, new species: A, anterior end, dorsal

view; B, pharynx and setiger 1, ventral view; C, anterior end, dorsolateral view; D, left 1

parapodium 1, dorsal view; E, left parapodium 12, posterior view; E right parapodium 25, l

anterior view; G-I, superior compound falcigers from setiger 2; J, same from setiger 3; K-M,

compound falcigers from setiger 4: K, superior seta; L-M, upper intermediate setae; N-P, •

compound falcigers from setiger 25: N, superior seta; O, intermediate seta; P,inferior seta; I

Q, superior seta from midbody segment; R, same from far posterior segment; S-T, dorsal

simple setae: S, from setiger 2; T,from midbody; U, from far posterior segment; V-Y, aciculae: •

V, from setiger 1; W, from midbody segment; X, from setiger 25; Y, from far posterior l

segment. (A-Y, originals by JDK) ..................................................................................... 25

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I Figure 1.8. Sphaerosyllis catiforniensis: A, anterior segments, dorsal view (Goleta 6/HI); B, prostomium,

dorsal view; C, left parapodium, setiger 7, anterior view; D, left parapodium, setiger 30,

i anterior compoundview; E, fight parapodium, setiger 22, posterior view (LBGS B8EII); F-G, superior falcigers, setiger 2; H, inferior comtx)und falciger, setiger 2; I-K, compound falcigers

from setiger 31: I, superior seta; J, intermediate seta; K, inferior seta; L-N, dorsal simple

i setae: L, setiger 28; M, setiger 32; N, setiger 33; O, ventral simple seta, setiger 31; P-Q, aciculae: P,setiger 2; Q, setiger 31; R, Posterior end, dorsal view. (A-R, originals by JDK.)

............................................................................................ .............................................. 29

I Figure 1.9. Sphaerosyllis B, bilineata Kudenov and Harris, new species: A, anterior segments, dorsal view; prostomium and setigers I-2, lateral view; C, ventrum, median setiger; D, superior

compound falciger, setiger 8. (A-D, originals by J. Dorsey) ............................................. 32

I Figure 1.10. Sphaerosyllis ranunculus Kudenov and Harris, new species: A, anterior end, dorsal view; B,

prostomium, dorsal view; C, pmstomium, ovigerous female, dorsal view; D, rightparapodium,

i setiger 7, anterior view; E, fight parapodium, setiger 16, anterior view; F-G, compound falcigers from setiger 4: F, superior seta; G, inferior seta; H-I, compound falcigers from

setiger 5: H, inferior seta; I, superior seta; J, compound falciger, setiger 20; K, compound

falciger, setiger 22; L, dorsal simple seta, setiger 18; M, ventral simple seta, setiger 23; N,

I acicula, setiger 15; O, acicula, setiger 19. (A-O, originals by JDK) ................................ 34

Figure 1.11. Dioplosyllis lagunae: A, anterior end, dorsal view; B, prostomium and anterior segments,

I dneuroaciculae, orsal view; C, pharynx; D, setiger 7, anterior view; E, compound falcigers, sefiger 7; F, setiger 7. (A-E redrawn from Mueller and Fauchald, 1976) .................... 37

Figure 1.12. Dioplosyllis tridentata Kudenov and Harris, new species: A, anterior segments, dorsal view;

I B, prostomium and setiger 1, lateral view; C, left parapodium, median segment, posterior

view; D-E, superior compound falcigers; F,inferior compound falciger; G, acicula. (A-C, E,

F, redrawn from originals by J. Dorsey; D, original by JDK) .......................................... 39

I Figure 1.13. Eusyllis bloomstrandi: A, anterior end, dorsal view; B, parapodium 1;C, parapodium 15; D,

parapedium 17 from epitokous specimen; E, inferior compound falciger, parapodium 1;E

I inferio body segment; r compoundH, aciculae, falciger, middle parapodiumbody segment; 1. (A-H, modified G, superiorandcoredrawn mpound fromfalciger, Hartmann- middle

Schrrder, 1971) ................................................................................................................ 42

I Figure 1.'14. Eusyllis habei: A, anterior end, dorsal view; B, pharynx, dissected; C, parapodium 18; D,

superior compound falciger, middle segment; E, inferior compound falciger, middle segment;

F, superior compound falciger, posterior segment; G, inferior compound fatciger, posterior

I segment; H, dorsal simple seta, posterior segment; I, ventral simple seta, same posterior segment as H; J, acicula, median parapodium; K, anterior end, ventral view. (A-K, redrawn

from Imajima, 1966c) ....................................................................................................... 44

I Figure 1.15. Eusyllis longicirrata: A, anterior end, dorsal view, distal parts of antennae, tentacular and

dorsal cirri omitted; B, pharynx dissected; C, parapodium 11;D, superior compound falciger,

i pampodium 11; E, inferior compound falciger, parapodium 11; F, aciculae, pampodium 11. (A-F, redrawn from Imajima, 1966c) ................................................................................ 46

Figure 1.16. Odontosyllisphosphorea: A, anterior end, dorsal view; B, teeth and lateral chitinized regions

I of pharynx, frontal view; C, anterior parapodium; D, medial biramous parapodium from epitokous specimen; E, compound falciger, setiger 10; F, same, setiger 70; G, same, from

inferior fascicle; H, dorsal simple seta, posterior segment; I, ventral simple seta, posterior

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segment; J, acicula, setiger 1; K, same, posterior segment; L, same, setiger 70. (A, modified •

from Blake, 1975; B, E-F, H-L, redrawn from Banse, 1972; C-D, G, redrawn from Moore,

1909) ................................................................................................................................. 48

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Figure 1.17. Odontosyllisfragilis Kudenov and Harris, new species: A, anterior segments, dorsal view;

B, left parapodium, setiger 11, setae omitted, anterior view; C, left parapodium, setiger 11,

including dorsal cirrus and setae, anterior view; D, left parapodium, setiger 36, anterior II

view; E, pharynx, dissected, dorsolateral view; F,superior compound falciger, setiger 11;G, I

inferior compound falciger, setiger 11; H, paired aciculae, setiger 11; I, paired aciculae

embedded in acicular lobe, setiger 11.(A-I, originals by JDK) ........................................ 50 I

Figure 1.18. Opisthodonta mitchelli Kudenov and Harris, new species: A, Anterior end, dorsal view; B,

right parapodium, setiger 4, anterior view; C, superior compound falciger, setiger 4; D, •

inferior compound falciger, setiger 4; E, shaft tip of compound falciger, posteroventral view; II

F, shaft tip of compound falciger, ventral view; G, shaft tip of compound falciger, dorsal

view; H, Acicula. (A-G, redrawn from originals by J. Dorsey) ........................................ 52 •

Figure 1.19. Pionosyllis magnifica: A, anterior end, dorsal view; B, parapodium; C, compound falciger.

(A-C, redrawn from Moore, 1906) ................................................................................... 55

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Figure 1.20. Pionosyllis articulata Kudenov and Harris, new species: A, anterior end, dorsal view; B, I

pharynx and setiger 1, ventral view; C, superior compound falciger, setiger 6; D, inferior

compound falciger, setiger 6; E, superior compound falciger, with detail of tip, setiger 40; F, •

inferior compound falciger, setiger 40; G, dorsal simple seta, setiger 40; H, ventral simple |

seta, setiger 40. (A-H, redrawn from originals by J. Dorsey) ........................................... 56

Figure 1.21. Syllides reishi: A, anterior end, dorsal view; B, prostomium and everted pharynx, dorsal B

view; C, parapodium 16, posterior view; D, dorsal simple seta, parapodium 10; E, compound

falciger, parapodium 10; F, shaft end of compound falciger, ventral view; G, acicula,

parapodium 10. (A-G, redrawn from J. Dorsey, 1978) ..................................................... 60 I

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Figure 1.22. SyIlides mikeli Kudenov and Harris, new species: A, anterior end, dorsal view; B, prostomium

and pharynx everted, dorsal view; C, pharynx, dorsal view; D, left parapodium, setiger 5, II

dorsal view; E, left parapodium, setiger 7, anterior view; F, right parapodium, setiger 9, E

anterior view; G, right parapodium, setiger 19, anterior view; H-J, compound falcigers from

setiger 1; H-I, superior setae; J, inferior seta; K-M, compound falcigers from setiger 3: K,

superior seta; L, intermediate seta; M, inferior seta; N-O, inferior compound falcigers, setiger II

6; P-Q, compound falciger from setiger 20; P, superior seta; Q, inferior seta; R-T, dorsal

simple setae: R, setiger 1; S, setiger 16;T, setiger 20 with distal hood, dorsal view; U, ventral

simple seta, setiger, far posterior segment; V-W, aciculae: V, setiger 2; W, setiger 19. (A-W, E

originals by JDK) .............................................................................................................. 62

Figure 1.23. Ehlersia heterochaeta: A, anterior end, male epitoke, dorsal view; B, anterior end, atoke, •

dorsal view; C, anterior end, atoke, dorsal view; D, inferior compound falciger, anterior II

segment; E, inferior compound falciger, posterior segment; E superior compound falciger,

anterior segment, with detail of tip; G, superior compound falciger, posterior setiger; H, •

ventral simple seta, posterior setiger. (A-H, after Dorsey and Phillips, 1987) .................. 65 |

Figure 1.24. Ehlersia hyperioni: A, anterior end, holotype, dorsal view; B, anterior end, holotype, ventral

view; C, anterior end, lateral view; D, left parapodium, setiger 9, posterior view; E, right I

parapodium, setiger 78, posterior view; F, inferior compound falciger, anterior setiger; G, m

superior compound falciger, posterior setiger; H, inferior compound falciger, posterior setiger;

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I I, superior compound falciger, posterior setiger, with detail of tip not to scale; J, dorsal

simple seta, posterior setiger; K, ventral simple seta, posterior setiger; L, paired aciculae,

I anteri modified or setiger; M, acicula, posterior setiger; N, pygidium, holotype, dorsal view. (A-N, from Dorsey and Phillips, 1987) ........................................................................ 67

Figure 1.25. EurysylIis spicum Kudenov and Harris, new species: A, anterior segments, dorsal view

(SBMNH); B, prostomium, dorsal view (SBMNH); C, anterior segments, dorsal view (Sta. BRC-13); D, anterior segments, ventral view (Sta. BRC-13); E, right parapodium, setige r

I 1, posterior view; F, right parapodium, setiger 25, posterior view; G, right pampodium,

I setiger 50, posterior view; H, superior compound falciger, setiger 11; I, inferior compound falciger, setiger 11; J-K, superior compound falcigers, setiger 25; L-M, superior compound

falcigers, setiger 50; N, inferior compound falciger, setiger 50; O, acicula, setiger 11; P,

I acicula, setiger 25; Q, acicula, setiger 50. (A-Q, originals by JDK) ................................ 69

Figure 1.26. GeminosylIis ohma: A, anterior end with pharynx everted, dorsal view; B, anterior end,

I detail of pharyngeal armature, dorsolateral view; C, parapodium 18; D, bifid simple seta with smooth cutting edge from anterior parapodium; E, bifid simple seta with minute serrations

along cutting edge from same anterior parapodium as D; F, ventral simple seta, posterior

parapodium; G, acicula, posterior parapodium. (A-G, redrawn from Imajima, 1966d.).. 72

I Figure 1.27. SyUis spongiphila: A, anterior end, dorsal view; B, parapodium 21 with long dorsal cirrus;

C, composite falciger, setiger 1; D, pseudocomposite falciger, setiger 1; E, same, median

i parapodium; parapodium 22. F, ventral simple seta, posterior parapodium; G, aciculae; H, short dorsal cirrus, (A-H, redrawn from Imajima, 1966d.) ................................................... 74

Figure 1.28. Trypanosyllis (Trypanosyllis) coeliaca nipponica: A, anterior end, dorsal view; B, median

I parapodium; C, superior compound bidentate falciger, setiger 6; D, median compound falciger,

setiger 6; E, inferior compound unidentate falciger; F, acicula, distal end. (A-F, redrawn

Imajima and Hartman, 1964) ............................................................................................ 76

I Figure 1.29. Trypanosyllis (Trypanosyllis) sp. A: A, prostomium and setiger 1, dorsal view, (USNM); B,

right parapodium, setiger 13, anterior view; C, same, posterior view; D, superior compound

I falciger, compoundsetiger falciger, 13;setiger E, intermediate 13; G, inferior cmpound cmpoundfalciger, falciger, setiger setiger13;13; F, H,lowacicula, intermediate setigr

13; I, Posterior end, dorsal view. (A-I, originals by JDK) ................................................ 78

I Figure 1.30. Trypanosyllis (Trypanosyllis) B: A, prostomium and setigers 1-2, dorsal view (USNM); sp.

B, prostomium, dorsal view (SBMNH); C, everted pharynx showing trepan, dorsal view

(SBMNH); D, everted pharynx showing trepan, ventral view; E, median segment, anterior

I view; F, superior compound falciger, median setiger; G, intermediate compound falciger, median setiger; H, inferior compound falciger, median setiger; I, ventral simple seta, setiger

25; J, inferior acicula, median setiger; K, superior acicula, median setiger. (A-K, originals by

I JDK) 79

Figure 1.31. Trypanosyllis (Trypanedenta) sp. A: left parapodium from far posterior segment, posterior

i view; B-C, compound falcigers from sefiger 15; D-F, compound falcigers from far posterior segment: D, superior seta; E, intermediate seta; F,inferior seta; G, aciculae from far posterior

segment. (A-G, originals by JDK.) ................................................................................... 81

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Figure 1.32. Typosyllis alternata: A, anterior segments, dorsal view; B, prostomium, dorsal view; C, left I

parapodium, median segment, anterior view; D, left parapodium, median segment, dorsal

cirri omitted, anterior view; E, right parapodium, setiger 14, anterior view; F, superior i

compound falciger, median segment; G-H, J, compound falcigers from setiger 14: G, superior i

seta; H, intermediate seta; J, inferior seta; I, inferior compound falciger, median segment; KL, dorsal simple setae: K, median segment; L, setiger 14; M, ventral simple seta, setiger 14; •

N-O, aciculae: N, median segment; O, setiger 14; P, pygidium, dorsal view. (A-P, originals II

by JDK) ............................................................................................................................ 84

Figure 1.33. Typosyllis bella: A, prostomium, dorsal view; B, median segments with paired transverse i

bars of pigment per segment, dorsal view; C, fight parapodium, setiger 2, anterior view; D, I

superior compound falciger, setiger 2; E, inferior compound falciger, setiger 2; E, paired

aciculae, emerging through acicular lobe, superior acicula on right. (A-E, originals by JDK) H

.......................................................................................................................................... 6 B

Figure 1.34. TyposylIis hyalina: A, anterior end, dorsal view; B, median parapodium with short dorsal m

cirrus, posterior view; C, median parapodium with long dorsal cirrus, posterior view; D, I

superior compound falciger, parapodium 10; E, inferior compound falciger, parapodium 10;

F, compound falciger, median parapodium; G, compound falciger, posterior parapodium; H, i

posterior parapodium; I, dorsal simple seta, posterior parapodium; J, ventral simple seta i

from same parapodium as I; K, aciculae from median parapodium. (A-K, redrawn from

Imajima, 1966e) ................................................................................................................ 88 •

Figure 2.1. Aphroditidae: A,Aphrodita, entire animal, dorsal view; B,Laetmonice, entire animal, dorsal

view; C, harpoon seta; D, Aphrodita, anterior end, dorsal view, felt removed; E, parapodium

of cirrigerous segment, anterior view. (A-B, after Mclntosh; C, after Hartman; D, after i

Fordham; E, after Pettibone) .......................................................................................... 100 II

Figure 2.2. Aphrodita parva: A, anterior end, dorsal view; B, three types of neurosetae with spurs; C, HI

fight

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parapodium from segment 10; D, bearded neuroseta; E, notoseta. (after Moore, 1905).

103

B

Figux_e3.I. Polynoid morphology: A, anterior end of typical harmothoid polynoid, dorsal view; B, ventral i

view of same; C, prostomia of representatives of three polynoid subfamilies showing insertion

pattern of lateral antennae ............................................................................................... 106

i

Figure 3.2. " Arctonoefragilis: A, anterior end, dorsal view; B, median right elytron; C, median cirrigerous i

parapodium, anterior view; D, notoseta; E, neurosetae .................................................. 114

Figure 3.3. Arctonoe pulchra: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous i

parapodium, anterior view; D, notoseta; E, neurosetae .................................................. 116 !1

Figure 3.4. Arctonoe vittata: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous i

parapodium, anterior view; D, notoseta; E, superior neuroseta; F, inferior neuroseta .... I 18 H

Figure 3.5. Bylgides macrolepidus: A, anterior end, dorsal view; B, median left elytron with detail of

microtubercles and marginal papillae; C, median cirrigerous parapodium, posterior view; D, BE

notoseta; E, median neuroseta with detail of aristate tip ................................................. 120 II

Figure 3.6. Eucranta anoculata: A, anterior end, dorsal view; B, median fight elytron; C, median i

cirrigerous parapodium, anterior view; D, tip of notoseta; E, tip of superior neuroseta; F,tip i

of inferior neuroseta (D-F from Moore, 1910) ............................................................... 122

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I Figure 3.7. Eunoe senta: A, anterior end, dorsal view; B, median right elytron; C, median cirrigerous

parapodium, posterior view; D, superior notoseta; E, median notoseta; E distal portion of

i median neuroseta ............................................................................................................ 124 Figure 3.8. Gaudichaudius iphionelloides: A, anterior end, dorsal view; B, median right elytron; C,

median cirrigerous parapodium, anterior view; D, superior notoseta; E, inferior notoseta; F,

I distal portion of median neuroseta .................................................................................. 127

Figure 3.9. Harmothoe fragiIis: A, anterior end, dorsal view; B, median left elytron with detail of

i microtubercles notoseta; E, distal andportion marginalof papillae; C, median cirrigerous parapodium, anterior view; D, superior neuroseta; F, tip of median neuroseta ................. 129

Figure 3.10. Harmothoe hirsuta: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous

I parapodium, anterior view; D, notoseta; E, tip of median neuroseta .............................. 131

Figure 3.11. Harmothoe imbricata: A, anterior end, dorsal view; B, median right elytron with detail of

i microtubercles and marginal papillae; C, median cirrigerous parapodium, anterior view; D, distal portion of median notoseta; E, distal portion of median neuroseta ........................ 133

Figure 3.12. Harmothoe multisetosa: A, anterior end, dorsal view; B, median right elytron; C, median

I cirrigerous inferior neuroseta parapodium, ............................................................................................................ posterior view; D, notoseta; E, tip of median neuroseta; F,tip135of

i Figure 3.13. Hesperonoe laevis: A, anterior end, dorsal view; B, median cirrigerous parapodium, anterior view; C, superior notoseta; D, inferior notoseta; E, distal portion of superior neuroseta; F,

distal portion of inferior neuroseta .................................................................................. 137

I Figure 3.14. Lepidasthenia berkeleyae: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous parapodium, anterior view; D, superior neuroseta; E, distal portion of median

neuroseta ......................................................................................................................... 139

I Figure 3.15. Lepidasthenia longicirrata: A, anterior end, dorsal view; B, anterior cirrigerous parapodium,

posterior view; C, superior neuroseta; D, distal portion of median neuroseta; E, distal portion

of inferior neuroseta ........................................................................................................ 141

I Figure 3.16. Lepidonotus spiculus, new combination: A, anterior end, dorsal view; B, first right elytron;

C, median left elytron; D, detail of elytral surface from anterior elytron (top group), median

I elytron, notoseta;andG, posterior inferior notoseta; elytron; E, H, median distal half cirrigerous of medianparapodium neuroseta,.................................. anterior view; F, superior 143

Figure 3.17. MaImgrenieUa baschi: A, anterior end, dorsal view; B, median right elytron; C, median

I cirrigerous parapodium, posterior view; D, superior neuroseta; E, portion

distal of median

neuroseta ......................................................................................................................... 146

I Figure 3.18. Mal cirrigerous mgreniellaparapodium, macginitieianterior : A, anterior view,end, withdorsal detail view; of neuropodial B, median left supraacicular elytron; C,lobe; medianD,

distal portion of median neuroseta with detail of tip ....................................................... 148

I Figure 3.19. MaImgrenielIa nigralba: A, anterior end, dorsal view; B, posterior right elytron with detail

of pigmented reticular cell; C, median cirrigerous parapodiurn, anterior view; D, notoseta; E,

distal portion of superior neuroseta; E distal portion of median bifid neuroseta; G, distal

I portion of inferior knob-tipped neuroseta 150

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Figure 3.20. MalmgrenieUa scriptoria: A, anterior end, dorsal view; B, median right elytron with detail of I

surface; C, median cirrigerous parapodium, anterior view, with detail of neuropodial

supraacicular lobe; D, notoseta; E, distal portion of median neuroseta with detail of tip..... •

........................................................................................................................................ 152 l

Figure 3.21. Subadyte mexicana: A, anterior end, dorsal view; B, median right elytron with detail of l

marginal papillae; C, median cirrigerous parapodium, anterior view; D, notoseta; E, superior

neuroseta; E median neuroseta; G, inferior neuroseta .................................................... 154 i

Figure 3.22. Tenonia priops: A, anterior end, dorsal view, with pharynx everted; B, median left elytron; i

C, median cirrigerous parapodium, anterior view; D, notoseta; E, distal portion of inferior II

neuroseta ......................................................................................................................... 157

Figure 3.23. Ysideria hastata, new species, holotype: A, anterior end, dorsal view; B, posterior end, dorsal i

view, Ieft posteriormost parapodium underdeveloped ..................................................... 159 m

Figure 3.24. Ysideria hastata, new species, holotype: A, right eighth elytron from segment 15; B, left I

cirrigerous parapodium from segment 18, anterior view; C, inferior and superior notosetae, II

segment 18; D, median neuroseta from 2nd segment; E, superior neuroseta, segment 18; F,

median neuroseta; G, inferior neuroseta ......................................................................... 160 •

Figure 4.1. Acoetespacifica: Anterior end, dorsal view, left parapodium of segment 2 omitted; B, elytron

from segment 25; C, elytragerous parapodium of segment 2, anterior view; D, cirrigerous

parapodium of segment 3, anterior view; E, elytragerous parapodium of segment 25, anterior i

view, acicula and spinning gland dotted; F, notoseta from segment 2; G, neuroseta from i

segment 2; H, upper neuroseta from segment 3; I, middle neuroseta from segment 3; J, lower

neuroseta from segment 3; K, upper neurosetae from segment 9 (types a and b indicated).

(after Pettibone, 1989) .................................................................................................... 171

Figure 4.2. Polyodontespanamensis: A, anterior end, dorsal view; B, elylragerous parapodium of segment •

2, anterior view; C, notoseta from segment 2; D, middle and lower neurosetae from segment |

2; E, cirrigerous parapodium of segment 3, posterior view; F-H, upper (F), middle (G), and

lower (H) neursetae from segment 3; I, elytragerous parapodium of segment 9, anterior view, m

aciculae and spinning gland dotted; J-K, upper (J, types a and b indicated) and lower (K) |

neurosetae from segment 9. (after Pettibone, 1989) .............................................................

172

Figure 5.1. Pholoe glabra: A, anterior end, dorsal view; B, middle parapodium, anterior view; C, first •

elytron; D, middle elytron, with insets showing details of marginal papillae; E, superior

notopodial seta; F, neuropodial compound falciger. (all except insets after Hartman, 1961) •

........................................................................................................................................ 180

Figure 5.2. Pholoe courtneyae (Paratypes, Sta. 3-15, CASIZ 103569): A, anterior end, dorsal view, m

with only second elytron included for clarity; B, first and second elytra from right side of •

m

another paratype, with insets showing details of elytral papillae [not to scale]; C, elytron

from a posterior segment; D, parapodium from setiger 10 in posterior view, with insets of iim

different parapodial papillae [not to scale]; E, short geniculate serrated notoseta; F, long, thin

serrated notoseta; G, neuropodial falciger from middle of series .................................... 182

Figure 5.3. Pholoides aspera: A, entire animal, dorsal view; B, anterior end, dorsal view; C, dorsolateral •

view of prostomium and left tentaculophore, acicula dotted; D, parapodium from segment 2, II

anterior view; E, middle parapodium with dorsal tubercle, anterior view; F, middle elytral

beating parapodium, posterior view; G, elytron; H, neuropodiai compound falciger. (A, from •

Pettibone, 1953; B, from Johnston, 1897; C-H, after Pettibone, 1992) .......................... 184 l

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I Figure 6.1. Sthenelaisfusca: A, anterior end, dorsal view; B, anterior parapodium, posterior view; C,

etytron with insets showing tubercles and papillae; D, simple neuroseta; E, superior compound

i neuroseta; F,stout neuropodial falciger. (A, after Blake, 1975; B-C, after Pettibone,_197 lb; D-F, after Hartman, 1939) .............................................................................................. 193

Figure 6.2. Sthenelais berkeleyi: A, anterior parapodium; B, elytron with insets showing tubercles and

I papillae; C-E, compound neurosetae from middle group as insets. (all after Pettibone, 197lb) ........................................................................................................................................ 195

i Figure 6.3. Sthenelais and tubercles; verrucul C, anterior osa: A,parapodium anterior end,indorsal posterior view;view; B, elytron D, middle with insets parapodium showingin papillae anterior

view; E, simple neurosetae; F, compound neuroseta with 9 articulations. (all after Hartman,

i 1939) ............................................................................................................................... I97 Figure 6.4. Sthenelais tertiaglabra: A, anterior end, dorsal view; B, elytron with details of tubercles; C,

compound neuroseta from segment 2; D, simple spinose neuroseta; E, slender, unidentate

I neuropodial neuropodial falciger; falciger. F, (A-Dstoutafter bidentate Moore,neuropodial 1910; rest original) falciger; .......................................... G-H, anteroventral compound 199

Figure 6.5. SthenelaneUa uniformis: A, anterior end, dorsal view; B, anterior end, ventral view; C, ftrst

I elytron with inset showing marginal papillae; D, fifth elytron; E, third parapodium, anterior

view; F, middle parapodinm, anterior view; G, neurosetae from segment 2; H, neuropodial

falcigers from middle parapodium. (A-C, E-G, after Pettibone, 1969; D, H, after Hartman,

I t939) ............................................................................................................................... 201

Figure 6.6. Sigalion spinosus: A, prostomium, dorsal view; B, elytron with inset showing detail of marginal

I papillae; multiarticulated C, anterior neuroseta. parapodium; (all after D, middle Hartman, parapodium; 1939) ..................................................... E, simple neuroseta; F-G, compound 203

Figure 7.1. Chloeia pinnata: A, anterior end, dorsal view; B, same, anterodorsal view; C, same,

I view; D, worm, view; E, setiger 10, view; F,

anteroventral entire ventral anterodorsal left

setiger 12, posterior view; G, posterior end, dorsal view. Setae omitted for clarity. All scales

= 1 mm. Scale 1, D; 2, E; 3, F; 4, A-C; 5, G................................................................. 210

I Figure 7.2. ChIoeia pinnata: A, notoseta, from superior of fascicle, with A' showing detail of tip; B,

same, with B' showing detail of surface; C-E, bifurcate harpoon notosetae; F-K, spurred

I nfascicle; otoseta I-J, frominferior innerofsetal fascicle; field:K,F,superior center ooff fasci fascicle; cle; L-N, G, superi spinose or onotosetae, f fascicle;fromH, anteri outermost or of

whorl: L, superior of fascicle; M, anterior of fascicle; N, inferior of fascicle; O, notoacicula;

I P-S, neuroseta; spurredR, neurosetae from outer field of fascicle: P, superior neuroseta; Q, anterior inferior neuroseta; S, posterior neuroseta; T-W, spurred neurosetal capillaries

from inner setal field: T, superior neuroseta; U, anterior neuroseta; V-W, inferior neurosetae;

i X-A X, B, surface details of neurosetae: X, Y,AA, spurred neurosetae from outer field of fascicle; superior neuroseta; Y, posterior neuroseta; AA, anterior neuroseta; Z, AB, spurred

neurosetal capillaries from inner field of fascicle: Z, longest capillary from center of fascicle;

i AB, shorter capillary from inferior of fascicle; AC, neuroacicula. Scales: 1 = .001 ram, X- AB; 2 = .001 mm, B'; 3 = .01 mm, A-W, AC and .005 ram, A' .................................... 212

Figure 8.1. Euphrosine arct/a: A, anterior end, dorsal view; B, prostomium, dorsal view; C, anterior

I end, anteroventral view; D, left setiger 13, posterior view; E, Branchiae Nos. 1-5, depicting dorsal and lateral cirri, posterior view; F, pygidium, posteroventral view. All scales = 1mm.

Scale 1, A, C, D, F; 2, E; 3, B........................................................................................ 221

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i xxi
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Figure 8.2. Euphrosine arctia: A, superior notoseta, tier 1; B,inferior notoseta, tier 1; C, superior notoseta, I

tier 3; D, inferior notoseta, tier 3; E, middle notoseta, detail of shaft, tier 3; F,long superior

ringent notoseta, tier 2; G, short superior ringent notoseta, tier 2; H, short inferior ringent i

notoseta, tier 2; I, ringent notoseta, detail of shaft, tier 2; J, long superior neuroseta; K, long l

neuroseta from center of fascicle; L, long inferior neuroseta; M, short inferior neuroseta; N,

subinferiormost neuroseta; O, inferiormost neuroseta; P, short anterior neuroseta; Q, long •

anterior neuroseta; R, long posterior neuroseta; S, short posterior neuroseta; T, superior I

neuroseta, detail of shaft; U, anterior acicula; V, superior acicula. Scales: 1= .001 mm, E, I,

T; 2 = .005 mm, A-D, F-H, J-S, U-V. ............................................................................. 222 •

Figure 8.3. Euphrosine bicirrata: A, anterior end, dorsal view; B, prostomium, dorsal view; C, anterior

end, anteroventral view; D, right segment 9, posterior view; E, left setiger 12, posterior view;

F, pygidinm, posteroventrat view. All scales = 1 ram. Scale 1, A; 2, D-E; 3, C-F; 4, B ...... i

........................................................................................................................................ 224

Figure 8.4. Euphrosine bicirrata: A, superior notoseta, tier 1; B. inferior notoseta, tier 1; C, superior I

notoseta, tier 4; D, inferior notoseta, tier 4; E, superior notoseta, detail of shaft, tier 4; F, |

inferior notoseta, detail of shaft, tier 4; G, superiormost notoseta, tier 2; H, superior notoseta,

tier 2; I, inferior notoseta, tier 2; J, superior ringent notoseta, tier 3; K, inferior ringent i

notoseta, tier 3; L, largest superiormost neuroseta; M, superior neuroseta; N, long neuroseta l

from center of fascicle; O, same, from point below center of fascicle; P, subinferiormost

neuroseta; Q, inferiormost neuroseta; R, short anterior neuroseta; S,long anterior neuroseta; i

T, long posterior neuroseta; U, short posterior neuroseta; V, superior neuroseta, detail of H

shaft; W, anterior acicula. Scales: 1 = .001 ram, E, F, V; 2= .005 ram, A-D, G-U, W... 225

Figure 9.1. Morphological features of onuphids: prostomial and peristomial appendages and jaws. A, i

anterior end, dorsal view; B, C, schematic dorsal and ventral views of prostomium; D,

mandibles; E, maxillae. (all from Paxton, 1986a) .......................................................... 230

Figure 9.2. Morphological features of onuphids: parapodia and setae. A, fn-st parapodium, Nothria; B, i

i

anterior parapodium, Onuphis; C, middle parapodium, Diopatra; D, E, scoop-shaped and

flat pectinate setae; F, compound spiniger (Mooreonuphis); G, subacicular hook; H, I, m

pseudocompound hooks; J, simple hooded hook ("large median hook"); K, pseudocompound

hook (Paradiopatra). (all from Paxton, 1986a) ............................................................. 232

n

Figure 9.3. Kinbergonuphis vexillaria: A, anterior end, dorsal view; B, maxillae; C, parapodium 2, •

anterior view; D, middle parapodium, anterior view; E, posterior parapodium, anterior view;

F, pseudocompound seta; G, simple set,a;H, pectinate seta, setiger 2; I, same, setiger 10; J,

subacicular hook ............................................................................................................. 237 i

Figure 9.4. Mooreonuphis exigua: A, anterior end, dorsal view (after Shisko, 1981); B, anterior

parapodium, anterior view; C, simple set,a; D, slender pseudocompound hook; E, stout

pseudocompound hook; F, compound spiniger ................................................................ 239 i

Figure 9.5. Mooreonuphis nebulosa: A, anterior end, dorsal view; B, posterior end, dorsal view; C,

maxillae; D, mandibles; E, free end of acicula; F, anterior parapodium, anterior view; G, •

middle parapodium, anterior view; H, slender pseudocompound hook; I, stout pseudocompound l

hook; J, large hook; K, limbate seta; L, pectinate seta; M, subacicular hooks. (C after Moore,

1911; F, G after Gathof, 1984) ....................................................................................... 241 i

i

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i
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I Figure 9.6. Mooreonuphis segmentispadix: A, anterior end, dorsal view; B, maxillae; C, parapodium 1,

anterior view; D, prebranchial parapodium, anterior view; E, parapodium 12, anterior view;

i E, G,far posterior parapodium, anterior view; F,slender and stout (left) pseudocompound hook; compound spiniger; H, subacicular hook; I, upper simple seta; J, lower simple seta; K,

pectinate seta; L, free tip of acicula ................................................................................ 243

I Figure 9.7. Nothria occidentalis: A, anterior end, dorsolateral view; B, parapodium 1, anterior view; C, parapodium 2, anterior view; D, middle parapodium, anterior view; E-F, simple and

pseudocompound hook, setiger 1;G-H, uni- and bidentate pseudocompound hooks, setiger

I 2; I, nearly compound hook, setiger 3; J, pectinate seta; K-M, anterior, middle, and posterior simple setae ..................................................................................................................... 245

i Figure 9.8. Onuphis affinis Hilbig new species: A, anterior end, dorsal view; B, maxillae; C, mandibles; D-E, parapodium 1 and 4, anterior view; F-G, middle and posterior parapodium, anterior

view; H, pseudocompound hook; I, subacicular hook; J, simple seta; K, pectinate seta. (all

from non-type specimens) ............................................................................................... 248 II

Figure 9.9. Onuphis elegans: A, anterior end, dorsal view; B, middle parapodium with wide gill; C,

pseudocompound hooks, setiger 2; D, subacicular hook. (all from Blake, 1975a) ......... 250

I Figure 9.10. Onuphis geophiliformis: A, anterior end, dorsal view; B, maxillae; C, first parapodium,

anterior view; D, fourth parapodium, anterior view; E, middle parapodium, anterior view; F,

pseudocompound hooks; G, subacicular hook; H, pectinate seta; I, posterior end, dorsal

I view. (A-H after Maekawa and Hayashi, 1989) ............................................................ 252

Figure 9.11. Onuphis iridescens: A, anterior end, dorsal view; dashes: pigment pattern; B, maxillae; C,

I mandible; H, stout andD-E, slenparapodia der pseudocompound 2 and 15, anterior setae;view; I, subacicular F, middlehook; parapodium, J, simpleanterior seta ........... view;254GFigure 9.12. Onuphis sp. "intermediates", specimen indentified and dissected by Hobson: A, anterior end,

I view; B, parapodium or 3, view; C-E pseudocompound same,

dorsal 2 anterior hooks from

dosal to ventral; G, subacicular hook; H, pectinate seta ................................................. 256

I Figure 9.13. Paradiopatra anterior views parva (after: Fauchald, A, anterior1968); end, dorsal E, pseudocompound view; B, maxillae; seta (after C-D, Fauchald, parapodia 1968); 1 and F-7,

G, subacicular hooks, anterior and far posterior setiger; H-I, pectinate setae, anterior and far

I posterior setiger; J, posterior end, dorsal view ................................................................ 258 Figure 9.14. Rhamphobrachium longisetosum: A, anterior end, dorsal view; B, maxillae; C, mandibles;

D, parapodium 4, anterior view; E, far posterior parapodium, anterior view; E recurved

I hook; G-I, pseudocompound setae; J-K, pectinate setae; L, subacicular hook. (A, F after Berkeley and Berkeley, 1938; B, C after Hartman, 1944; D, G-I after Paxton, 1986b). 260

I Figure 10.1. Eunice region, americana anterior view; : A, Danterior , compound end, dorsal falciger; view; E, B, pectinate maxillae;seta;C,F, parapodium subacicularfromhook; branchial G, tips

of aciculae. (D-G after Fauchald, 1992) ......................................................................... 268

I Figure 10.2. Eunice multicylindri: A, anterior end, lateral view; B-C, parapodia from branchial and postbranchial region, anterior vierw; D, compound falciger; E, pectinate seta; F,subacicular

hook. (A, D-F after Fauchald, 1992) .............................................................................. 270

I Figure 10.3. Eunice multipectinata: A, anterior end, dorsal view, proboscis partly everted; B, middle

parapodium, anterior view (after Fauchald, 1992); C, pectinate seta; D, compound falciger;

i E, subacicular hook; F, tips of aciculae; G, posterior end, dorsal view.......................... 272
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Figure 10.4. Eunice vittatopsis: A, anterior end, dorsal view; B-D, anterior, middle, and posterior parapodia, m

anterior view; E, detail of limbate seta; F, pectinate seta; G, anterior (left) and posterior

compound falciger; H, subacicular hook. (B-D, G, H after Fauchald, 1992)................ .274 n

Figure 10.5. Marphysa conferta: A, anterior end, dorsal view; B, middle parapodium, anterior view; C,

pectinate seta; D, compound falciger; E, subacicular hook............................................ 276

m

Figure 11.1. Eranno bicirrata: A, anterior end, dorsal view, peristomium slightly distorted because of m

everted proboscis; B, posterior end, dorsal view; C, prostomium and partly extended proboscis,

lateral view; D, maxillae, dorsal view, left MIII- MV omitted, right MIII - MV slightly m

displaced for better visibility; E, mandibles; F-H, anterior, median, and posterior parapodium, l

anterior view; I, hooded hook, setiger 15; J, same, setiger 120 (from Fauchald, 1970). 285

Figure 11.2. Eranno lagunae: A, anterior end, dorsal view; B, maxillae, dorsal view, in situ; C-F, anterior, n

median, posterior, and far posterior parapodium, anterior view; G, hooded hook, anterior

m

setiger, with detail of crest; H, same, posterior setiger .................................................... 287

m

Figure 11.3. Lumbrineris californiensis: A, anterior end, dorsal view; B, some posterior segments, dorsal m

view; c, maxillae, dorsal view, in situ; D, mandibles (after Hartman, 1968); E, anterior

parapodium, posterior view; F,same, anterior view; G, posterior parapodium, anterior view; •

H, compound hooded hooks, same fascicle, setiger 7; I, simple hooded hook (F, G, I after |

Fauchald, 1970) .............................................................................................................. 289

Figure 11.4. Lumbrineris cruzensis: A, anterior end, dorsal view (after Day, 1973); B, maxillae, dorsal n

view, in situ, right MIII and MIV after Hartman, 1944; C, mandibles; D-F, anterior, median,

and posterior parapodium, anterior view; G, simple hooded hooks, setiger 20; H, compound

hooded hooks, setiger 10 (C-F after Hartman, 1944) ..................................................... 291 m

m

Figure 11.5. Lumbrineris index: A, anterior end, dorsal view; B, posterior end, dorsal view; C, maxillae,

dorsal view, in sit-u;D, mandibles, adult specimen; E, same,juvenile specimen; F-H, anterior, •

posterior, and far posterior parapodium, anterior view; I, compound hooded hook, with detail m

of crest; J, simple hooded hook (G, H after Moore, 1911) ............................................. 293

Figure 11.6. Lumbrineris inflata: A, anterior end, dorsal view; B, posterior end, dorsal view; C, MII, m

MIII, and MIV, dorsal view; D, MI and maxillary carriers, dorsal view; E, mandibles; F-H,

first, median, and posterior parapodium, anterior view; I, compound hooded hook, setiger 6;

J, simple hooded hook (A, B, F-H after Imajima and Higuchi, 1975; C-E, I, J from Imajima m

and Higuchi, 1975) ......................................................................................................... 295

Figure 11.7. Lumbrinerisjaponica: A, anterior end, dorsal view; B, posterior end, dorsal view; C, MII, m

MRI, and MIV, dorsal view; D, MI and maxillary carriers, dorsal view; E, mandibles; F, first m

parapodium, anterior view; G, posterior parapodium, anterior view; H, limbate capillary

m

seta, setiger 5; I, compound hooded hook, setiger 1; J, simple hooded hook, setiger 60 (A, B, m

F, G after Imajima and Higuchi, 1975; C-E, H-J from Imajima and Higuchi, 1975) ..... 297 m

Figure 11.8. Lumbrineris latreilli: A, anterior end, dorsal view; B, MI and maxillary carriers, dorsal

view; C, MII, MIII, and MIV, dorsal view; D, mandibles; E-G, first, 10th, and posterior •

parapodium, anterior view; H, limbate capillary seta, setiger 1; I, compound hooded hook,

setiger 10; J, simple hooded hook, posterior setiger (all from Imajima and Higuchi, 1975).

299 m

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xxiv m
!

I Figure 11.9. Lumbrineris limicola: A, anterior end, dorsal view; B, maxillae, dorsal view, in situ; C,

mandibles; D-F, anterior, median, and posterior parapodium, anterior view; G, simple hooded

i hook; H, compound hooded hook, with detail of crest (C-F after Hartman, 1944; G from Fauchald, 1970) .............................................................................................................. 301

Figure 11.10. Ninoe gemmea: A, anterior end, dorsal view; B, same, ventral view; C, maxillae, dorsal

I view, in situ; D, mandibles; E-I, first through fifth parapodium, anterior view; J, 25th parapodium, anterior view; K, hooded hook, setiger 25; L, same, setiger 100; M, limbate

capillary seta, setiger 25 (A, B, E-I from holotype; D, K-M after Moore, 1911) ........... 303

I Figure 11.11. Ninoe palmata: A, anterior end, dorsal view; B, MI and maxillary carriers, dorsal view; C,

MII, MIll, and MIV, dorsal view; D, mandibles; E-G, first, 16th, and 75th parapodium,

anterior view; H, limbate capillary seta, setiger 75; I, hooded hook, anterior setiger; J, same,

I setiger 75 (all from Imajima and Higuchi, 1975) ........................................................... 305

Figure 11.12. Ninoe tridentata Hilbig, new species: A, anterior end, dorsal view; B, maxillae, dorsal view,

I inlastsitu; andC-G, last gilI-tmaring first, second,parapodium, third, lOth,anterior and 25thview; parapodium, J, limbate anterior capillaryview; seta, anterior H-l, second-to- setiger;

K, same, posterior setiger; L, hooded hook, anterior setiger; M, same, posterior setiger. (A,

I C-l: paratype USNM 170426; B, J-M: holotype) ........................................................... 307

Figure 11.13. Scoletoma tetraura: A, anterior end, dorsal view; B, maxillae, dorsal view; in situ; C,

mandibles; D, 10th parapodium, anterior view; E, posterior parapodium, anterior view; F,

I hook, setiger, crest; G, same, posterior setiger (C-G

hooded anterior with detail of after

Orensanz, 1973) .............................................................................................................. 310

I Figure 12.1. Arabella Orensanz,(A.1974); ) iricolorC,: A, mandibles anterior end, (afterdorsal Pettibone, view (from1963);BlaD-E, ke, 1975); anteriorB, maxillae and posterior (after

parapodium, anterior view (after Fauchald, 1970); F,smooth limbate seta (after Pettibone,

I 1963); G, serrated limbate seta ....................................................................................... 321

Figure 12.2. Arabella (A.) pectinata: A, anterior end, dorsal view; B, middle setigers, dorsal view; C,

mandibles; D, maxillae; E-E anterior and posterior parapodium, anterior view; G, finely

I serrated seta, anterior parapodium; H-J, middle and lower seta from posterior

upper,

parapodium. (B, E, F after Fauchald, 1970) ................................................................. 323

I Figure 12.3. Arab middleellaand(A.posterior ) protomutans parapodium, : A, anteanterior rior end,view; dorsalF, vie serrated w; B,supra-aeicular maxillae; C, msetae, andibles; posterior D-E,

parapodium; G, ventralmost setae, posterior parapodium. (all after Orensanz, 1990 except

for left seta in F and right seta in G)............................................................................... 325 Figure 12.4. Arabella (A.) semimaculata: A, anterior end, dorsal view; B, maxillae; C, mandibles (after

Moore, 1911); D-E, anterior and posterior parapodium, anterior view (after Fauchald, 1970);

I F,fmely serrated seta; G-H, coarsely serrated seta, lateral and frontal view (detail) ..... 327

Figure 12.5. Drilonereisfalcata: A, anterior end, dorsal view; B, middle setigers, dorsal view; C, anterior

i end, lateral view; D, mandibles; E, maxillae; F, middle parapodium, anterior view; G-H, seta in lateral and frontal view; I, spine. (D-F after Orensanz, 1974) .................................... 329

Figure 12.6. Drilonereis longa: A, anterior end, dorsal view; B, maxillae and mandibles; C-D, anterior

I and afterposte Pettibone, rior parapo 1963)di

.....................................................................................................

um, anterior view; E, limbate seta. (A-D after Uebelacker, 1984;331E

I
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Figure 12.7. Drilonereis mexicana: A, parapodium, anterior view; B, maxillae; C, seta (all from Fauchald, m

1970) ............................................................................................................................... 333

Figure 12.8. Drilonereis nuda: A, anterior end, dorsal view; B, maxillae; C, posterior parapodium, anterior n

view; D, supraacicular seta; E, spine; E subacicular seta .............................................. 335 U

Figure 12.9. Drilonereis spectabilis Hilbig, new species: A, anterior end, dorsal view; B, posterior segments, •

dorsal view; C, mandibles; D, maxillae; E, posterior parapodium, anterior view; F-G, seta, I

lateral and frontal view; H, spine. (all from holotype) .................................................... 337

Figure 13.1. Dorvilleidae: A, capillary supraacicular setae; B, cultriform supraacicutar seta i

(Ophryotrocha); C, long-tined furcate seta; D, geniculate and short-tined furcate setae; E, i

bidentate subacicular setae; F, spinigerous subacicular seta (Exallopus); G, falcigerous

subacicular seta (Ophryotrocha); H, modified supraacicular setae (Exallopus); I-J, modified n

subacicular setae, heterogomph and homogomph (Exallopus); K, cultriform ventralmost seta I

(Ophryotrocha); L, subbiramous parapodium; L, uniramous parapodium; N, habitus of

interstitial species (A renotrocha)(from various sources) ................................................ 345 n

w

Figure 13.2. Dorvilleidae: A-E, maxillae; A, with maxillary carriers, two pairs of basal plates, and and

four rows of free denticles; B, with forceps and seven pairs of free denticles in four rows •

(Ophryotrocha); C, with two rows of denticles; D, reduced to smooth, soft plates U

(Pseudophryotrocha); E, modified, with two rows of denticles (Anchidorvillea); F-J,

mandibles; F, Dorvillea; G, Ophryotrocha; H, Anchidorvillea; I, modified, L-shaped •

(Ophryotrocha); J, rodlike (Pseudophryotrocha)(from various sources) ....................... 347 |

Figure 13.3. Dorvillea (Schistomeringos) annu/ata: A, anterior end, dorsal view; B, maxillae; C, mandibles;

D, middle parapodium, ventral setal lobe extended; E, subacicular setae, shortest and longest n

blade of fascicle; F, furcate supraacicular seta; G, capillary supraacicular seta (after Fauchald,

1970) ............................................................................................................................... 351

n

Figure 13.4. Dorvillea (Schistomeringos) longicornis: A, anterior end, dorsal view (eyes faded); B, superior i

base plate, detail of proximal teeth; C-G, superior free denticles from proximal to distal: C,

very close to base plate, D-E, middle of superior row, F-G, ftrst and fifth denticle; H-J, distal i

inferior free denticles; K, parapodium, anterior view, ventral setal lobe extended; L, same,

ventral setal lobe retracted; M, furcate supraacicular seta; N, subacicular seta. (D, E, G after

i

Fauchald, 1970) .............................................................................................................. 353

Figure 13.5. Parophryotrocha brevicapitis Hilbig, new species: A, anterior end and last setigers of fragment,

dorsal view; B, same, ventral view; C, parapodium; D, spines; E, ventralmost capillary seta.

(all from holotype) .......................................................................................................... 356 B

Figure 13.6. Parougia batia: A, anterior end, dorsal view; B, maxillae; C, right mandible (after Jumars,

1974); D, parapodium, anterior view; E-F, two upper, short-bladed and one lower, longbladed subacicular seta; G, supraacicutar seta ............................................................... 358 ii

Figure 13.7. Pem'boneia brevipalpa: A, anterior end, dorsal view; B, anterior parapodium; C-D, capillary

and furcate supraacicular setae; E, subacicular seta; F, maxillae; G, mandibles. (from Hilbig •

and Ruff, 1990) .............................................................................................................. 360 l

!

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xxvi i
!

I Acknowledgments

I This Study was funded by the Pacific Outer Continental Shelf Region of the U.S. Department of the

Interior, Minerals Management Service, Washington, D.C., under Contract No. 14-35-0001-30484. Ms.

I Andrea Moe of the Santa BarbaraMuseum of NaturalHistory provided production assistance for this volume. The editors would like to thank Drs. KristianFauchald, Kirk Fitzhugh, Mary E. Petersen, Marian H.

Pettibone, andDavid Russell for providing critical reviews of individual chapters. The senior editor especially

I thanks Dr. Petersen for providing guidance on the pholoids that greatly improved the technical content of Chapter 5.

List of Acronyms

i

I AMNH American Museum of Natural History, New York, New York, USA.

I AMW Australia Museum, Sydney, Australia.

ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania.

I BLM Bureau of Land Management.

BMNH The Natural History Museum, London, United Kingdom

I BRA Refers transecttoA/aB.station designation from the MMS Phase I Reconnaissance: Benthic Rocky,

BRC Refers to a station designation from the MMS Phase I Reconnaissance: Benthic Rocky,

I transect C/D.

CASIZ California Academy of Sciences, Department of Invertebrate Zoology, San Francisco,

I California, USA.

LACM NaturalHistory Museum of Los Angeles County, Los Angeles, California. Includes the type

collections formerly lodged atAllan Hancock Foundation (AHF).

I MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA.

MMS Minerals Management Service.

I SCAMIT Southern California Association of Invertebrate Taxonomists.

SBMNH Santa Barbara Museum of Natural History, Santa Barbara, California, USA.

I USNM United States National Museum. A historical designation for the National Museum of Natural

History (NMNH), Smithsonian Institution, Washington, D.C., USA.

I Zoologisches Museum, Hamburg, Germany

ZMH

I

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i xxvii
IIIIIIIIIIIIIIIIIII
I

I

| 1. FAMILYSYLLIDAEGRUBE, 1850

by

I Jerry D. Kudenov I and Leslie H. Harris 2

I

Introduction

I

The SyIlidae represent one oftbe most diverse and systematically challenging families of Polychaeta,

I with tend toabobeutsmall, 70 genusually era andless overthan60010species mm long(Hartman, and 1 mm1959, wide,1965; andFauchald, are especially1977welt ; Pettibone, represented1982as). epibiot Syllidsa

on hard surfaces or on metazoans in intertidal, subtidal and shelf habitats. Other members of this family are

I alsofewabugenera ndantinordeeper widely waters, distributed suchinashal Sphaerosyllis low soft-bottoandm Braniella environmenints.slopeTheandSyllrise idaehabitats. is represented by only

Morphology

I The body is usually small, short and slender or threadlike, nearly cylindrical to dorsoventrally flattened

in cross section, and tapering anteriorly and posteriorly. The integument is generally smooth, but papillae of

various shapes and distributions are sometimes present.

I The prostomium tends to be wider than long. Eyes number two to three pairs; the posterior two pairs

are arranged trapezoidally, and the anterior pair tends to be eyespots associated with the anteriormost region

of prostomium. Antennae generally number three, including unpaired median and paired lateral antennae,

I may strongly articulated, or smooth; they are inExogonella, organs may

which be wrinkled absent Nuchal be

present as epaulettes or lobes extending from the posterior prostomial margin, or as continuous transverse or

paired postectal lobes. Palps are paired, usually large, roughly triangular in shape, range from being completely

I fused to free from one another, and project anteroventrally. However, the of palps Autolytus are greatly

reduced, not visible dorsally, and project ventrally from the prostomium.

i The pharynx is an eversible tube with a cuticular lining, and generally with a dorsomedian tooth that

tends to be marginal or subdistal when the pharynx is everted, although it may be present in the middle or

posterior half of the pharynx. Additionally, a distal circlet or arc of marginal pharyngeal teeth may accompany

the dorsomedian tooth, all of which may become so well developed that together they appear to form a crown

I or trepan (as Trypanosyllis). Sometimes the pharynx is described as being unarmed, i.e., pharyngeal teeth

in

are absent, as in Syllides. A distal circlet of 10 soft papillae is usually also present, although these may be

fewer in number or absent in some taxa. The proventriculus is conspicuous, with transversely striated muscles

I arranged rows, connects pharynx to posterior

in and the anterior the intestine.

_Department of Biological Sciences, University of Alaska Anchorage, 3211 Providence Drive, Anchorage, Alaska 99508

:Invertebrate Zoology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles,

I California 90007

I 1
!

1

The tentacular segment is achaetous, and generally forms a complete ring, although sometimes it is m

dorsally reduced or variously developed into an occipital flap that extends anteriorly over the prostomium, l

covering one or both pairs of posterior eyes. Ventrally the tentacular segment forms the lips of the mouth.

Tentacular cirri generally number two pairs, although only one pair may be present in Exogone; Exogonella

lacks cirri altogether. 1

Parapodia are uniramous and similar all along the body in nonreproductive individuals; they may be

biramous in reproductive individuals. Setae are generally compound falcigers (as in Typosyllis), or sometimes I1

compound spinigers (as in some species of Exogone and Pionosyllis), or are entirely simple (as inHaplosyllis). I

In some taxa, the articulation between the blade and the shaft is fused. A dorsal and ventral simple seta may

also be present in each fascicle for a varying number of segments. Aciculae are present in all parapodia, 1R

varying both in number and shape, i

Dorsal cirri vary in shape and length, sometimes alternating between long and short, and generally

present on all parapodia, although they may be absent from an anterior segment in some taxa. Dorsal cirri I1

may be articulated or moniliform (as inSyllis, TyposyIIis or Trypanosyllis); smooth (as inExogone, Odontosyllis l

or Pionosyllis); or irregularly wrinkled (as in Eusyllis or Streptosyllis). Ventral cirri are mostly digitiform,

and usually do not extend beyond the parapodial lobes; they are entirely absent from the subfamily Autolytinae. I

The pygidium has a pair of anal cirri, and sometimes also a ventromedian cirrus. The cirri are I!

usually slender, ranging from articulated to smooth.

!

Systematic History

I

Grube (1850) was the first to recognize syllids as a separate family, followed by Williams (1851 ) who

also considered the Syllidae as a separate family. Other important early works on syllids include Quatrefages •

(1865), Malaquin (1893), Gravier (1900), Haswell (1920a, b; 1921), Fauvel (1923, 1934a, b) and Rioja 1

(1925). Rioja (1925) divided the Syllidae into four subfamilies: Autolytinae, Exogoninae, Eusyllinae and

Syllinae, each of which aredefined and accompanied by a key below, and have been recognized by subsequent •

investigators (Fauchald, 1977; Pettibone, 1982). |

Recent literature of importance includes the studies by Berkeley and Berkeley (1938, 1945), Rioja

(1941, 1943), Uschakov (1955), Gidholm (1962, 1965, 1967a,b), Pettibone (1954, 1963, 1982), Imajima []

(1966a-e, 1967), Day (1967), Hartman (1961, 1968), Hartmann-Schrtder (1971), Fauchald (1977), Uebelacker I!

(1984), Russell (1987, t989a-b, 199I) and San Ma_n (1990, 1991a,b). The excellent series of papers by

Imajima (1966a-e, 1967) are undoubtedly the most important works that have consistently clarified important []

traits for the entire family. In this context, Imajima largely stabilized syllid systematics. II

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I

2 !
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I Principal Diagnostic Traits

I Uebelacker The(1984) four subfamilie are characterized s definedasbyfollows: Rioja (1925), and recognized by Imajima (1966a), Day (1967) and

I Subfamily Autolytinae

I one another, Diagdirected nosis. Body ventrally. threadlike, Nuchal4-2organs 0 mm present long, 0.5-1 eithermmas wconspicuous ide. Palps partiall epaulettes y toorcolobes. mpletelyTentacular fused to

cirri numbering 2 pairs. Antennae, tentacular cirri and dorsal cirri smooth. Ventral cirri absent. Pharynx

I long, coiled or sinuous, with a distal circlet of teeth (=trepan).

i Subfamily Exogoninae

Diagnosis. Body rather compact, less than 8 mm long, 0.8 mmwide. Palps fused dorsally for at least

I one-half pairs; mayto beall absent. their length. Antennae, Nuchaltentacular organs usuall cirriyandincondorsal spicuocirusri. smooth. Tentacular Ventral cirri ucirri suallypresent. numberiPharynx ng 1-2

usually straight, with a middorsal tooth or unarmed.

I Subfamily Eusyllinae

I Diagnosis. Body usually at least 10 mm long. Palps only fused basally, free from one another

distally. Nuchal organs large and conspicuous. Tentacular cirri numbering 1-2 pairs. Dorsal cirri smooth to

wrinkled. Ventral cirri present. Pharynx usually straight, with a middorsal tooth with or without a marginal

I series of teeth, or unarmed.

Subfamily Syllinae

i Diagnosis. Body usually at least 10 mm long, sometimes over 50 ram. Palps usually free basally.

Nuchal organs usually small and inconspicuous. Tentacular cirri numbering 2 pairs. Dorsal cirri strongly

articulated or moniliform. Ventral cirri present. Pharynx straight, with a middorsaI tooth, with or without

distal circlet of teeth (=trepan). !

tt is relatively easy to distinguish autolytins and exogonins from one another and from each of the

I obecause ther twokeysubfcharacters amilies. Unf somet ortuinamestelyo,verlap. it is ofteFor n diffic example, ult to idsome entifyeusyllins both eusymay llinshave and sydistinctly llins to aarticul subfamil atedy

dorsal cirri or basally fused palps and some syllins may have smooth dorsal cirri or basally separated palps

I (Dphysi ay,cal1967; factors Uebsuch elacker, astemperature 1984). Fusi(Piltz, on ofp1980), alps insuggesting the Eusyllinae that thisandparticular Syllinae ma character y be partl is not y cinformative. ontrolled by

Although distinctions between smooth to wrinkled versus articulated dorsal cirri of eusyUins and syllins are

I not alwayAs reliable, this trait is still the most practical one to use when placing these taxa into subfamilies.

synopsis of the majority of syllid genera is presented by Fauchald (1977). Important generic traits

include: body shape, epidermal papillae, prostomial antennae, occipital flap, tentacular cirri, nuchal organs,

I nature of dorsal and ventral cirri, nature and distribution of setae, aciculae, and nature and armature of the
!

n

pharynx. Perhaps knowledge of the pharynx is the most critical of these traits. The pharynx may be unarmed, •

lacking any sort of dentition, or may have an internal valve. An armed pharynx can show a variety of

dentition patterns: (a) a single middorsal tooth with a smooth or minutely dentate anterior rim or margin; (b) •

a continuous distal circlet of teeth (trepan) which may have an enlarged median tooth; (c) a discontinuous I

ventral arc or an incomplete circlet of teeth. The position of the middorsal tooth is also important because it

can range from distal, subdistal, or proximal in the pharynx. Pharyngeal dentition is often the only way to •

separate such genera as Eusyllis and Pionosyllis. l

Important species traits include: pigmentation patterns; shape and distribution of such structures as

antennae and eyes, palps, nuchal organs, pharynx, proventriculus, peristomial cirri, occipital flap, parapodia, n

dorsal and ventral cirri, setae and aciculae. Among the more important traits noted above are the setae, I

aciculae, and pharynx. Both setae and aciculae must be examined and illustrated from superior and inferior

regions of fascicles of anterior, middle and posterior parapodia using a compound microscope. It is also •

important to scan the entire specimen for the distribution and morphological changes in setae and aciculae l

along the body (Fauchald, 1977), especially in such genera asExogone, SylIides andStreptosyllis. Typically,

blades of compound falcigers tend to vary in length, decreasing both inferiorly within fascicles, and posteriorly R

along the body; aciculae generally decrease in both size and number per parapodium posteriorly along the I

body. Setae and aciculae must also be examined and illustrated using oil-immersion, particularly the ends of

shafts and blade tips of compound setae, and the tips of both aciculae and simple setae. The pharynx is •

critical to species identifications, and it is usually not protracted in preserved specimens. Sometimes it can be I

readily observed through the body wall of small translucent specimens. Otherwise the pharynx must be

removed if possible (Day, 1967; Fauchald, 1977; Uebelacker, 1984) and illustrated, i

One must take care when identifying or describing syllids. Often, it is necessary to examine individual

specimens one at a time. This does not mean that the taxon is so fraught with taxonomic difficulties that it

should remain the purview of a few specialists. However, it is clear that much of the present confusion i

surrounding syllid systematics is due to at least three obvious factors. First, many workers have described

new species, often in complicated genera, without having examined relevant type materials. Similarly, there

is a tendency for some workers to apply names of species described from such areas as Europe (especially by •

Fauvet, 1923) to species that have similar suites of characters from the western coast of North America

(Berkeley and Berkeley, 1948; Hartman, 1968). Such undue reliance on literature descriptions, instead of

types, has cumulatively destabilized species concepts. For example, earlier records ofExogone gemmifera •

represent at least one undescribed species (Banse and Hobson, t 974; this study). Some of these species may l

truly be "cosmopolitan" in distribution, but until the types are critically examined, there is simply no way of

knowing the number of "hidden" species involved. Secondly, welt-known species that may have limited •

dispersal abilities have been identified far from their type localities. Again, many such identifications are I

seldom compared to types, which adds greater uncertainty to syllid systematics and further complicates efforts

to identify species. A third factor is that a number of species are either too poorly known, original type m

specimens are no longer available, or Ieetotypes have not been redescribed. l

Prospects for stabilizing syllid systematics will be attainable only in the long term. Systematic practice

on and descriptions of syllids needs to be standardized. The best way to set this process in motion is for •

workers to compare their identifications to existing type material during revisionary studies of genera. Where

type material is not available, one must obtain and describe specimens from type localities and not rely solely

on published descriptions. Other systematic techniques should also be pursued in the study of syllids. For •

example, an intriguing karyological study (Curini-Galletti et al, 1991) summarized chromosomal data for l

syllids, and suggested that: (a) congeneric species usually have similar karyotypes; and (b) one likely mechanism

of karyological evolution involves chromosomal fission or fusion, while polyploidy is speculated to be another !

mechanism that may have led to high numbers of chromosomes in both the Eusyllinae and Syllinae. SyUids l

are an ecologically important group of infauna and epibiota, and efforts to render them more understandable

should be made by all interested potychaetologists, m

l

4 !
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'1 Biology

I often associated Syllids arewithusually sponges, small, hydroids, cryptic, corals, highly bryozoans, motile polychaetes ascidians,thatalgae dominate and seagrasses. hard substrata. SyllidsTheyof the are

subfamily Exogoninae are well represented in soft sediments where they inhabit superficial layers, while

i oless thersdiverse are commin deep on inwater, algal although mats and ccertain oral mbble. generaThesuchentire as SphaerosylIis family is best , Braniella representedandinSyllis shallowareseas, present. being

Most syllids are considered to be carnivorous, piercing the body walt of prey with their armored

I pharynges, Other taxa, and suchingesting as the Exogoninae body fluidsmaythrogenerally ugh the pbeumping selectiveactiondeposit of thefeeders highlyormus consume cular prdiatoms oventriculus. (Day,

1967; Fauchald and Jumars, 1979; Petfibone, 1982).

I Syllid reproductive biology is both interesting and complex (Heacox, 1980; Heacox andSchroeder, 1981a,b). Sexes are separate, but species can generally reproduce both asexually and sexually. Asexual

reproduction occurs through the formation of stolons which are posterior body segments of sexually derived

I adults (or parental atoke) that become transformed solely for the purpose of carrying gametes. Stolons are transient structures, with prostomia bearing appendages and 2 pairs of conspicuous eyes, long tentacular and

dorsal cirri, and biramous parapodia with notopodial fascicles of long natatory or swimming capillary setae.

I Each stolon can either be produced singly, in long serial chains, or in dense clusters, detaching periodically to

spawn (Dales, 1967; Schroeder and Hermans, 1975). Stolons of the Exogoninae, Eusyllinae and Syllinae

tend to be morphologically similar: male and female stolons are called a chaetosyllis stage; those of TrypanosyUis

I' and polybostrichus Eurysyllis are stages calledandahave tetraglong lene divergent stage. Stolons palps;ofemale f the Autstolons olytinaearearecalled dimorphi sacconereis c: malestages stolonswithareventral called

pouches or marsupia containing large yolky ova (Gidholm, 1962).

I Sexually reproducing syllids may spawn their gametes directly into seawater after they have become modified into swimming stages called epitokes. Epitokes generally have large eyes, and biramous parapodia

with notopodial fascicles of long natatory setae in certain segments. Epitokes eventually swarm to surface

I waters in response to various environmental cues where gametes are released en masse, producing free- swimming larvae. Another mode of sexual reproduction is brooding by females in which developing embryos

are attached directly in sacs to the dorsum, ventrum or parapodia. The embryos usually number two per

I segment, and develop directly into individuals with 4-6 segments that resemble small adults.

I Key to the Subfamilies and Genera of the Syllidae (After Imajima, 1966a; Fauchald, 1977)

I 1A. Ventral cirri absent; dorsal cirri smooth ......................................................................... Autolytinae 3

i lB. Ventral cirri present; dorsal cirri smooth to articulated .................................................................... 2

2A. Body small, usually less than 8 mm long; dorsal cirri smooth; palps fused for at least half their

lengths (Figs. 1.1A, 1.7B, 1.10A) ............................................................... Exogoninae ............... 4

2B. Body larger, usually more than 10 mm long; dorsal cirri smooth to articulated; palps not fused or

fused only basally (Figs. 1.15A, 1.19A) .......................................................................................... 3

i

3NospeciesoftheAutolytinaewereencounteredinthecollectionsexaminedaspart ofthisstudy.The Autolytinaeare,

I however,wellrepresentedin Californiawaters(Hartman,1968;L. Harris,personalobservation,1995).

! '
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3A. Palps fused basally (Figs. 1.19A, 1.20A); dorsal cirri smooth to wrinkled (Figs. 1.18A, 1.19A) ..... i

.................................................................................................................... Eusyllinae .................. 7

3B. Palps not fused basally (Figs. 1.27A, 1.28A); dorsal cirri articulated (Figs. 1.26C, 1.28A) ............ i

.................................................................................................................... Syllinae ................. 12 U

i

4A. Two pairs of tentacular cirri (Figs. 1.1A, 1.2A, 1.25B) ................................................................. 5 i

4B. One pair of tentacular cirri (Figs. 1.5A, 1.6A, 1.8A, 1.10A) ......................................................... 6

IN

5A. Dorsal cirri long and filiform (Figs. 1.1A, 1.2A) ................................................................... Brania n

5B. Dorsal cirri globular or spherical (Fig. 1.25E-G) ............................................................. Eurysyllis i

n

6A. Dorsal cirri papilliform or ovoid (Figs. 1.4C, 1.7D-F) ....................................................... Exogone

6B. Dorsal cirri pyriform or flask-shaped (Figs, 1.8C-E, 1.9A, t.10D-E) ........................ Sphaerosyllis m

7A. Eversible pharynx unarmed (Figs. 1.21B, 1.22B-C) ............................................................ SyUides n

7B. Eversible pharynx armed (Figs. 1.12A, 1.18A) ............................................................................... 8 n

8A. Pharynx with a single middorsal tooth (Figs. 1.20B, 1.26B, 1.30C,D, 1.32A) ............................... 9 n

8B. Pharynx with a distal circlet of smaller curved teeth (Figs. 1.1613,1.17E) .................. OdontosyUis

Im

9A. Middorsal tooth located in posterior region of eversible pharynx (Fig. 1.18A) ............ Opisthodonta i

9B. Middorsal tooth located in anterior region of eversible pharynx (Figs. 1.12A, 1.14B, 1.15B, 1.20B) im

..................................................................................................... **........................................... , .... 10 •

n

10A. Anterior margin or rim of eversible pharynx denticulated (Figs. 1.14B, 1.15B) ................... Eusyllis i

10B. Anterior margin or rim of eversible pharynx smooth ..................................................................... 11

11A. Parapodia long; palps 2 times longer than prostomium (Figs. 1.11C, 1.12A) ................ Dioplosyllis n

11B. Parapodia short and conical; palps maximally same length as prostomium (Fig. 1.20B) ..................

......................................................................................................................................... Pionosyllis n

12A. Pharynx with middorsal tooth (Figs. 1.23B, 1.25A, 1.26B, 1.32A) .............................................. 13

12B. Pharynx with middorsal tooth and trepan (Fig. 1.30C-D) ............................................................. 15 i

13A. Setae include compound falcigers (Fig. 1.27C), dorsal and ventral simple setae, and pseudocompound I

setae (Fig. 1.27D-E) .................................................................................................................. Syllis []

13B. Setae include compound falcigers, some with extremely long spiniger-like blades, and dorsal and

ventral simple setae ........................................................................................................................ 14 i

i

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I 14A. Compound falcigers and spiniger-like falcigers with extremely long bidentate blades at least 4 times

longer than inferiormost blades within fascicles of midbody segments (Figs. 1.23G, 1.24G,I) .........

I ............................................................................................................................................. Ehlersia

14B. Only compound falcigers present (Figs. 1.32F-I, 1.33D-E, 1.34D-J) ............................... Typosyllis

I 15A. Palps same length as prostomium (Fig. 1.26A); body cylindrical; setae simple (Fig. 1.26D-E) .......

...................................................................................................................................... Geminosyllis

I 15B. Palps shorter than prostomium (Figs. 1.29A, 1.30A-B); body flattened; setae compound (Figs. 29D- G, 30B-F) .................................................................................................................... Trypanosyllis

l List of Syllid Species

I In all, 16 genera and 34 species of syllids have been identified in MMS samples from the Santa Maria

Basin and western Santa Barbara Channel. One species is a new combination, and 14 other species in 11

I are new to science. The following species list is organized by subfamily. Type specimens are deposited

genera

in the National Museum of Natural History, Smithsonian Institution (NMN-H), Natural History Museum of

Los Angeles County (LACM-AHF), Santa Barbara Museum of Natural History (SBMNH) and the Australian

I Museum, Sydney (AM W); all non-type specimens of new taxa listed below are deposited at the LACM-AHF. As noted above, the Autolytinae are not represented in voucher collections, and were infrequently encountered

in MMS samples. Excellent papers on the Autolytinae by Gidliolm (1962, 1965, 1967a,b) and Imajima

I (1966b) should be consulted.

I Subfamily Exogoninae

i

II Brania brevipharyngea Banse, 1972

mm Brania californiensis Kudenov and Harris, new species

Exogone (Exogone) lourei Berkeley and Berkeley, 1938

I Exogone (Exogone) dwisula Kudenov and Harris, new species Exogone (Parexogone) molesta Banse, 1972

Exogone (Parexogone) acuffpalpa Kudenov and Harris, new species

I Exogone (Parexogone) breviseta Kudenov and Harris, new species Sphaerosyllis californiensis Hartman, 1966

Sphaerosyllis bilineata Kudenov and Harris, new species

I SphaerosyUis ranunculus Kudenov and Harris, new species

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Subfamily Eusyllinae I

I

Dioplosyllis lagunae (Hartman, 1961), new combination I

Dioplosyllis tridentata Kudenov and Harris, new species

Eusyllis blomstrandi Malmgren, 1867 •

Eusyllis habei Imajima, 1966c !

Eusyllis longocirrata Imajima, 1966c

Odontosyllis phosphorea Moore, 1909 l

Odontosyllisfragilis Kudenov and Harris, new species 1

Opisthodonta mitchelli Kudenov and Harris, new species

Pionosyllis magnifica Moore, 1906 l

Pionosyllis articulata Kudenov and Harris, new species I

Syllides reishi Dorsey, 1978

SyUides mikeli Kudenov and Harris, new species I

Subfamily Syllinae I

Ehlersia heterochaeta (Moore, 1909) I

Ehlersia hyperioni Dorsey and Phillips, 1987 HI

Eurysyilis spicum Kudenov and Harris, new species

HI

GeminosyUis ohma Imajima, 1966d m

Syllis spongiphila Verrill, 1885 1

TrypanosylIis (Trypanedenta) sp. A

m

Trypanosyllis (Trypanosyllis) coeliaca nipponica Imajima and Hartman, 1964 I

Trypanosyllis (Trypanosyllis) sp. A 1

Trypanosyllis (Trypanosyllis) sp. B

I1

Typosyllis alternata (Moore, 1908) n

Typosyllis bella (Chamberlin, 1919) 1

Typosyllis hyalina (Grube, 1863) 1B

I

Description of Species

ml

Subfamily Exogoninae

I

Genus Brania Quatrefages, 1866 I

Grubea Quatrefages, I865:35 (in part). I

Protogrubea Czemiavsky, 1881:414 (in part), i

Pseudobrania San Martin in Sard_i, 1984:7-13.

1W

Pseudobrania: San Mart/n, 1984a: 150-151. I

Grubeosyllis

Verrill, 1900, sensu San Mart/n, 1991b:716-717. I

Type species: Exogone pusilla Dujardin, 1851

I
m

m

m Diagnosis. Prostomium with 3 antennae. Palps well developed, fused at least basally, as long as

prostomium. Two pairs of tentacular cirri. Small nuchal organs present between prostomium and tentacular

m segment. Pharynx with single middorsal anterior tooth. Dorsal cirri longer than setal lobes. Ventral cirri as long as setal lobes. Embryos carried on dorsum of females. Aciculae acuminate with sharp filiform tips, or

distally notched or excavate, with blunt tip.

m Remarks. San Martin (1984a) proposed the Pseudobrania to include previously described

genus

Brania species that have (a) prostomial antennae, tentacular and dorsal cirri long and fusiform; (b) acuminate

acicula with distally sharp tips; and (c) palps completely united by a dorsal membrane, and not separated by

m a distal cleft or notch. This genus was prematurely described by Sard_i (1984) but because it was based upon San Martin's unpublished dissertation, the authorship San Martin in Sarda must be applied. However,

Pseudobrania is actually preoccupied in the Platyhelminthes, andSan Martin (199 lb:716-717) instead proposed

m using Mart the previously named genus, Grubeosyllis Verrill, 1900. According to the scheme proposed by San in, Brania sensu stricto differs from GrubeosyUis in having, (a) prostomial antennae, tentacular and

dorsal cirri clavate or truncate; (b) aciculae distally excavate with tooth or blunt tips; and (c) palps united

m basally bydorsal membrane, distally separated by a small notch or cleft. There are apparently other differences pertaining to morphology of the compound setae.

The necessity of San Martin's (1984a, 1991b) approach is highly questionable. Brania sensu lato

m includes 21 species (Fauchald, 1977), and is a The division of this tends to

relatively compact genus. genus

obscure similarities previously recognized among congeneric species. San Martin (1991b) also suggests a

closer relationship between members of his newly redefined GrubeosyUis and Parapionosyllis Fauvel, 1923,

m than between Grubeosyllis andBrania sensu stricto. This statement is difficult to comprehend. For example, Parapionosyllis differs from GrubeosylIis in having one, instead of two pairs of tentacular cirri, and in having

compound spinigers rather than compound falcigers. Finally, examinations of both B. brevipharyngea and a

m new Brania species described below from MMS materials suggests considerable overlap in the characters used by San Martin (1984a, 1991b) and Sard_i (I984). For example, the palps of the new MMS Brania

species are distally notched and separated from one another, the aciculae are distally blunt and excavate, and

m some of the dorsal cirri are fitiform, while others may be clavate. Similarly, the aciculae ofB. brevipharyngea are also excavate and distally blunt, although the palps appear to be fused by a dorsal membrane for their

entire length. Perhaps the criteria used for these generic distinctions rely too heavily on soft anatomical

I features that not only change allometfically over life cycles, but also are easily influenced by collecting efforts and preservation. The above features for the two MMS Brania species significantly overlap the two separate

sets of criteria defined by San Martin (1984a, 1991b) for both Grubeosyllis (=Pseudobrania) and Brania,

m call shedintconsiderable o serious quesdoubt tion the on the usefulness validityofofdivGrubeosyllis iding such .a relatively Therefore,comthepactraditional t genus (Brania generisensu c definition lato) andof

Brania is followed here, and GrubeosyUis is considered to be a junior synonym of Brania.

m Although Rioja (1941) described Brania heterocirra from Mexico, Westheide (1974) reported B.

heterocirra from the Galapagos, Japan (where Imajima (1966a) recorded it as B. clavata), and Washington

(where Banse (1972) in part identified it as B. brevipharyngea). Brania heterocirra is included in the key

m below since it seems likely to be encountered in the syllid fauna of southern California. There are several additional undescribed species of Brania in California (L. Harris, unpublished).

!

m

! 9
!

[]

Key to Species of Brania From the Santa Maria Basin •

1A. Dorsal cirri varying in length along body; pharynx and proventriculus in same number of segments; I

cutting surfaces of superior dorsal simple seta smooth to minutely serrated ....... Brania heterocirra

1B. Dorsal cirri subequal in length along body; pharynx and pmventriculus in different number of segments; 1

cutting surfaces of superior dorsal simple seta strongly serrated ..................................................... 2 II

Pharynx in 2 segments (Fig. 1.1A); proventficulus in 4 segments ................ Brania brevipharyngea !

2A.

n

2B. Pharynx in 4 segments (Fig. 1.2A); proventriculus in 2 segments (Fig. 1.2B) ..................................

........................................................................................................................ Brania californiensis I

Brania brevipharyngea Banse, 1972 IN

Figure 1.1

Brania brevipharyngea Banse, 1972:198-200, fig. 4.--Banse and Hobson, 1974:55. I

Pseudobrania brevipharyngea: San Martin, 1984a:172, 174.--Sard_, 1984:12.

Grubeosyllis brevipharyngea: San Martin, 1991b:716-717. 1

II

Material Examined. California: Santa Maria Basin, off Purisima Point, Sta. BRA-20 (1, SBMNH

142697). --Washington: False Bay, San Juan Island, holotype (USNM 40711), paratypes (5, USNM 40712). 1

I

Description. Total body length about 3 mm, 0.4 mm wide without parapodia, 0.6 mm with parapodia,

for 30-31 setigers (Banse, 1972); MMS specimen an anterior fragment 1.5 mm long, 0.4 mm wide, with about

20 setigers. Body of preserved specimens generally colorless except for slight pigmentation on tentacular n

segment. Setigers 7-15 ovigerous, each with 12 ova per segment in 1 specimen (Banse, 1972:199). U

Prostomium wider than long (Fig. 1.IA); 2 pairs of eyes, anterior pair largest, lenticulate, in trapezoidal mm

arrangement; additional pair of eyespots present on anterior prostomium medial to lateral antennae (Fig. II

1.1A). Antennae longer than prostomium, bottle-shaped; median antenna arising between 2 pairs of large

mira

eyes; lateral antennae arising from anterior part of prostomium (Fig. 1.1A). Palps about as long as prostornium, Ill

fused completely (Fig. 1.1A). Pharynx extending through setiger 2 (2.5 setigers in MMS specimen); middorsal

tooth subdistal, lacking distal circlet of soft papillae when everted. Proventriculus extending from setigers 3-

m

6, with 20-25 rows of muscle cells (17-18 in MMS specimen). Tentacular segment distinct dorsally (Fig.

1.1A), with 2 pairs of bottle-shaped tentacular cirri; dorsal pair longest, each about aslong as median pro stomial 1

antenna; ventral pair each about one-half length of dorsal ones (Fig. 1.1A). w

Pampodia of median segments truncate, with small, distally rounded presetal lobes (Fig. 1.1B). I

Compound falcigers distally bidentate (Fig. 1.IC), 7 to 9 per pampodium. Blades decreasing in length |

inferiorly within fascicles, superior blades twice length of inferior blades; all with serrated cutting edges.

Shafts of compound falcigers enlarged distally; superior distal surfaces serrated (Fig. 1.1C). Superior dorsal II

simple seta distally bidentate with serrated cutting surface (Fig. 1.1D), present in all setigers from setiger 1or 1

4. Ventral inferior simple seta present from setigers 20-21 to end of body. One acicula per parapodium,

distally blunt, beak-shaped (Fig. 1.1E). Dorsal cirri long, bottle-shaped, present on all segments (Fig. 1.1A,B). in

Ventral cirri digitiform, extending beyond parapodial lobes (Fig. 1.1B). !

n

Pygidium with paired anal cirri, each bottle-shaped, 2-3 times length of anterior dorsal cirri.

!

lO N
!

i

I

!

I c

I

I -- '

i

1

I

II

Figure 1.1. Brania brevipharyngea: A, anterior end, dorsal view, setae schematic; B, parapodium 16, posterior

view; C, inferiormost compound falciger; D, superiordorsal simple seta; E, acicula. (A-E, redrawn

I Banse, 1972)

from

!

Remarks. Brania brevipharyngea is characterized by the combination of a short pharynx about two

I segments long, and a relatively long proventricle about 4 segments in length (Banse, 1972:199-200). Westheide (1974) reassigned one paratype originally described by Banse as B. brevipharyngea to B. heterocirra Rioja,

1941.

I Distribution, British Columbia.

California; Washington;

i

II 11
!

Brania californiensis Kudenov and Harris, new species I

Figure 1.2 I

Material Examined. California: offPoint Arguello, Sta. BRA-6, hototype (USNM 170899).

Description. Holotype incomplete. Originally with 22 setigers; 16 setigers present at time of I_

description. Length 1.4 mm, width 0.12 mm at setiger 6, without parapodia. Specimen colorless except for II

golden pigment just below constriction on antennae and tentacular cirri. Specimen originally carrying embryos

dorsally on setiger 12 through 15 on the fight side and setigers I2 and 13 on the left side; embryos 70 pm in

length. []

Prostomium wider than long; eyes numbering 2 pairs, located posteriorly on the prostomium; anterior

pair largest, both sets lenticulate; additional pair of eyespots present on anterior prostomium medial to lateral 1

antennae (Fig. 1.2A). Antennae longer than prostomium, bottle-shaped; median antenna located just anterior M

to postefior pair of eyes (Fig. 1.2A). Palps barely longer than prostomium, fused completely, pointed distally.

Pharynx extending through setiger 4, bright reddish-brown; middorsal tooth subdistal; distal end appearing I

smooth (pharynx inverted in specimen) (Fig. 1.2A). Proventriculus extending from midpoint of setiger 4 to II

midpoint of setiger 6, with approximately 19 rows of muscle cells (Fig. 1.2A). Tentacular segment indistinct,

with 2 pairs of bottle-shaped tentacular cirri; dorsal pair longest, each about two-thirds as long as median 1

prostomial antenna; ventral pair each about two-thirds length of dorsal pair (Fig. 1.2A). II

!

A |||

.05 rim1

I

''

I

C D E F 1

!

Figure 1.2. Brania ealiJ'orniensis Kudenov and Harris, new species : A, anterior end, dorsal view; B,

parapodium 6; C, superior compound falciger, setiger 6; D, median compoundfalciger, setiger 6; •

E, dorsal simple seta, setiger 6; F, acicula, setiger 6. (A-F,originals by JDK)

12 I
!

I Parapodia of median segments truncate, presetal and postsetal lobes equal in length; slight prolongation

of superior tips around bases of superior simple setae (Fig. 1.2B). Compound falcigers distally bidentate

i (Fig. 1.2C,D), 1.2C,D), approximately 7 per parapodium. Blades decreasing in length inferiorly within fascicles (Fig. superior blades (Fig. 1.2C) twice length of inferior blades; all with serrated cutting edges (Fig.

1.2C,D). Superior blades decreasing gradually in length posteriorly. Shafts of compound falcigers enlarged

i distally; superior distal surfaces serrated (Fig. 1.2C,D). Superior dorsal simple seta distally bidentate with serrated cutting surface, present in all setigers beginning on setiger 1(Fig. 1.2E). Ventral inferior simple seta

present from setiger 18, continuing to at least setiger 22. One acicula per parapodium, distally blunt, beaki shaped (Fig. Dorsal1.2F). cirri tong, weakly bottle-shaped, present on all segments. Ventral cirri digitiform, subequal to

or shorter than parapodial lobes.

I Pygidium unknown.

Remarks. Brania californiensis andB. brevipharyngea Banse 1972, are similar in setal morphology,

setal distribution, and form of the dorsal cirri. Brania californiensis differs from B. brevipharyngea in

I having the pharynx present in four segments, instead of two, and the proventriculus in two segments instead of four. In addition, ventral cirri do not extend beyond the parapodial lobes ofB. californiensis, in contrast to

those ofB. brevipharyngea.

I Etymology. species name, californiensis, to state specimens were collected.

The refers the where the

Distribution. Southern California.

I Genus Exogone Orsted, 1845

I Type species: Exogone naidina Orsted, 1845

i Diagnosis. Prostomium with 3 antennae. Palps fused dorsally throughout most or all of length. One pair of tentacular cirri. Nuchal organs inconspicuous. Pharynx armed with single anterior middorsal tooth.

Tentacular cirri, dorsal and ventral cirri ovoid to papilliform. Dorsal and ventral cirri shorter than setal lobes.

Developing embryos attached ventrally to posterior margins of median segments of female (Imajima, 1966a;

I Fauchald, 1977; Uebelacker, 1984).

Remarks. San Martin (1991b) recognized three subgenera of the genus Exogone: (1) species of

I Exogone with unmodified (Parexogone) heterogomph Mesnil hinges and Caullery, and shaft1916, tips;have (2) species unmodified ofExogone compound (Exogone) "spinigers" Orsted,and1845, falcigers all haveall

modified compound "spinigers" and falcigers, all with modified heterogomph hinges and shaft tips; and (3)

i species of Exogone (Sylline) Clapar&te, 1864, all have compound setae strongly modified, with blades and shafts partially fused, or lacking blades. Unfortunately, San Martin (1991b) uses the term "spiniger" for any

compound seta with a long blade. Many of the compound "spinigers" noted by San Martin are actually

i compound falcigers with long, tapering, distally bidentate blades, and explains the use of quotation marks in his definition of the above subgenera. The term spiniger is or should be used only to denote a "seta that tapers

to a fine point" (Fauchald, 1977), and is used to describe the fine tapering blades of compound setae. San

Martin (1991a) presents a useful key to the known species of each subgenus of Exogone, and a key to these

subgenera is presented below.

Although the genus Exogone is well understood, several species from the Pacific coast of the United

i States Exogoneandgemmifera Canada havePagenstecher, been erroneousl 1862,y andidentified E. verugera as European (Clapar&ie, species.1868)Forfromexample, southernrecords and ofcentral both

California apparently cannot be verified, contrary to statements by Hartman (1968). Moreover, Westheide

It (1974) found that some materials previously identified as E. gemmifera both by Pettibone (1954) and Imajima

i ii

|
!

(1966a) represent a new species, E. naidinoides, later emended by Russell, 1991, which he also encountered i

in the Galapagos Islands and from Pt. Barrow, Alaska. Given these findings, the identity of two specimens

reported as E. gemmifera by Lee and Rho (1992) from the Yellow Sea of Korea needs to be verified. Similarly, I

Westheide (1974) found that previous records ofE. verugera from Japan (Imajima, 1966a) represent another m

species (E. occidentalis), also emended by Russell, 1991, that occurs in the Galapagos Islands; San Martfia

(1991 b) considers E. occidentalis to be the junior synonym of Exogone breviantennata Hartmann-Schr6der •

(1959); SCAMIT materials ofExogone sp. A from southern Califomia may be identical toE. breviantennata. i

Finally, Westheide (1974) discovered some materials identified and described as E. molesta by Banse (1972)

and E. multisetosa by Friedrich (1956) represented still another new species, Exogone longicornis. It is also •

likely that the specimens described by Banse and Hobson (1974) as E. gemmifera represent an undescribed Ii

species that is closely related, if not identical, to the MMS specimens identified as Exogone dwisula, new

species, below. It seems likely that some of the species described by Hartmann-Schr6der (1959) and Westheide am

(1974) will eventually be encountered in the syllid fauna of southern California: Exogone breviantennata, E. m

longicornis and E. naidinoides are included in the appropriate subfamily-level keys that follow.

i

Key to the Subgenera of Exogone

(after San Martin, 1991b) I

1A. Antennae closely set, often in middle of prostomium, between eyes (Fig. 1.3A); compound setae with n

highly modified blades, shafts and heterogomph joints (Figs. 1.3C-E, 1.4D-F); dorsal simple setae I[

similar in all segments (Fig. 1.3B), or differing in form between anterior and posterior segments (Fig.

i

1.4G-I) ............................................................................................................................................ 2 i

1B. Antennae widely spaced, with median antenna in middle of prostomium and lateral antennae in front 1

of eyes; compound setae with unmodified blades, shafts and heterogomph joints (Fig. 1.6F-P); dorsal

simple setae similar in all segments (Fig. 1.7S-U) ........................................ Exogone (Parexogone) Hi

2A. Compound setae with blades as long falcigers [=spinigers of San Martin] narrow, terminating in

minutely bidentate tips (Fig. 1.3C), and blades of short falcigers with subdistal tooth larger than I

distal tooth (Fig. 1.3E); shaft tips and heterogomph hinges especially of anterior setae strongly modified []

(Fig. 1.3E); posterior dorsal simple setae thicker, differing in form to anterior simple setae (Fig. 4GI) ........................................................................................................................ Exogone (Exogone) I

2B. Compound setae all strongly modified, with blades partially fused to shafts and bayonet-shaped

blades lacking; shaft tips distally truncate; dorsal simple setae generally similar throughout body,

distally truncate with serrated cutting surfaces ..................................................... Exogone (Sylline) U

Subgenus Exogone (Exogone) (3rsted, 1845 1

[]

Type species: Exogone naidina Orsted, 1845

Diagnosis. Antennae closely set together, usually arising between eyes; compound falcigers with I

modified blades, shaft tips and heterogomph joints; blades including long, narrow forms with minute, distally

bidentate tips, and shorter forms with subdistal teeth larger than distal teeth; dorsal simple setae of 2 forms, •

differing between anterior and posterior body segments. I

I

14 !
!

I Key to Known Species of the Subgenus Exogone (Exogone)

from the Northeast Pacific

n 1A. Dorsal cirri present on setiger 2 (Fig. 1.3A); shaft tips of compound spinigers of setiger 2 (sometimes

also setiger 1) massively enlarged compared to those of adjacent setigers, distally falciform (Fig.

i 1.3C-D) 2

1B. Dorsal cirri absent on setiger 2 (Fig. 1.4A); shaft tips of compound spinigers and falcigers of setiger

2 enlarged compared to those of medial and posterior segments, not distally falciform (Fig. 1.4E) ..

I .............................................................................................. * ......................................................... 3

I 2A. Anten times nlonger ae lonthan g, extending lateral antennae; to or beyproventriculus ond anterior prextending ostomial margin through(Fig. 4-5 segments, 1.3A), median withantenn 18-24arows 2-3

of muscle cells; superior dorsal simple setae f'u'stpresent from setiger 1 ..........................................

I ................................................................................................................ Exogone (Exogone) lourei 2B. Antennae short, not extending to anterior prostomial margin, median antenna about as long as lateral

antennae; proventriculus extending through 7-8 segments, with 25 or more rows of muscle cells;

I superior dorsal simple setae first present from setigers 10-14 ........... Exogone (Exogone) uniformis

n 3A. Blades of compound setae in setigers 1-3uniformly short, deeply bifid (Fig. 1.4D-E) .....................

.............................................................................................................. Exogone (Exogone) dwisula

3B. Blades of compound setae in setigers 1-3 may include short and awl- or needle-like spinigers, or

I falcigers of various lengths ............................................................................................................. 4

4A. Compound falcigers with blades tong and tapering all along body; antennae short, subequal in length,

much less than one-half length of prostomium; proventriculus extending through 2.5-3 segments with approximately 20 rows of muscle cells ...................................... Exogone (Exogone) breviantennata

4B. Compound spinigers with awl-like blades; antennae long, with median at least 4 times longer than

I lateral antennae, and usually longer than combined length of prostomium and palps; proventriculus

extending through 2 segments with 31 rows of muscle ceils; 2 aciculae in anterior parapodia .........

1 ....................................................................................................... Exogone (Exogone) longicornis

I

Exogone (Exogone) lourei Berkeley and Berkeley, 1938 I

I Figure 1.3

Exogone lourei Berkeley and Berkeley, 1938:44, figs. 6-12; 1948:79, fig. 117.--Rioja 1941: 703-4, pl. 3, fig.

I 10-13.--Berkeley, 1967:1055.--Pettibone, 1967:5. Hartman, 1968:425-426, figs. t-5.--Banse 1972:

200-202, figs. 5a-d.--Banse and Hobson 1968:16, fig. 4d-e; 1974:58, figs. 14h-j.--Perkins

1981:1092.--Uebelacker, 1984:30-39 to 30-41, fig. 30-34 a-f.--Russell, 1991:55-57, fig. 2.

n Exogone uniformis (in part, including pt. fig. 1,pl. 7, fig. 1);

Hartman 1961:73-74 6, Hartman 1968: 427-428

(in part, including figs. 1, 2).

Exogone dispar Pettibone, 1954:259 (in part).

I Exogone (Exogone) lourei: San Martin, 1991a: 728, 735. SphaerosylIis californiensis Hartman, 1966: 196-197, pl. 3 (in part).

!
!

Material Examined. California: Santa Maria Basin, Stas. BRA-16 (3); 65 (1); off Point Sal, Sta. []

R-8 (33: USNM 170915); off Point Sal, Stas. PJ-1 (2l); PJ-2 (75); PJ-3 (21); PJ-4 (2); PJ-5 (10); PJ-6 (50); |

PJ-7 (50); PJ-8 (10); PJ-9 (1); PJ-10 (65); PJ-11 (5); PH-F (?1, regenerating head); offPoint Arguello, Stas.

PH-I (3); PH-R (I); PH-U (2).--Western Santa Barbara Channel, BRC-1 (6, USNM 170914), (20, SBMNH 1

142698), (2l, regenerating head).--R/V VeIero IV Sta. 5028-57, 1 paratype (removed from paratype lot of R

SphaerosyUis caIiforniensis, LACM-AHF Poly 1707).--R/V Velero IV Sta. 5102-57 (37: holotype and

paratypes of E. uniformis, LACM-AHF Poly 170, 171,521). British Columbia: False Narrows, holotype !

(USNM 32895). |

Description. Up to 10 mm long, 0.2 mm wide, for 50 segments. Prostomium wider than long,

pentagonal; eyes numbering 2 pairs; anterior pair largest, lenticulate, widest apart; posterior pair not lenticulate; •

all arranged trapezoidally (Fig. 1.3A). Antennae arising together from anterior part of prostomium; median II

antenna fusiform, about as long as palps, 2-3 times length of paired lateral antennae; latter small, digitiform

(Fig. 1.3A). Palps long, completely fused (Fig. 1.3A). Nuchal organs small, paired, at postectal corners of II

prostomium. Pharynx usually extending to setiger 3 (Fig. 1.3A); anterior middorsal tooth subdistal, surrounded II

by distal circlet of 10 soft papillae when evened. Proventficulus usually extending through 4-5 segments,

with 18-24 rows of muscle cells (Fig. 1.3A). Tentacular segment fused to prostomium; tentacular cirri []

numbering 1 pair, each small, papilliform (Fig. 1.3A). I

Parapodia each with 5-9 setae per fascicle, including compound spinigers and falcigers plus simple

setae (Fig. 1.3B-F). One or 2 compound spinigers occupying superior positions of fascicles; blades long, •

narrow; shafts of those in setiger 2, sometimes also setiger 1, greatly enlarged, thick, falciform (Fig. 1.3C,D). II

Compound falcigers with blades of similar lengths, all distally bidentate with primary tooth terminal, minute

(Fig. 1.3E). Dorsal superior simple seta urtidentate, slender, distally curved, minutely serrated from setiger 1; •

becoming thicker, sharply bent in posterior segments (Fig. 1.3B). Ventral seta curved, distally bidentate with 1

bent tips in median and posterior segments (Fig. 1.3F). Dorsal and ventral cirri digitiform, present on all

setigerousPygidium segments.with pair of I long, cirriform anal cirri.

Remarks. Exogone lourei is characterized by long compound spinigers with greatly enlarged shafts

on setiger 2, that may also occur on setiger 1; a long median and short paired lateral antennae; the proventriculus i

extending through 4 to 5 segments with around 16 to 30 (usually 18to 24) rows of muscle cells. E. lourei was

I

originally described from Vancouver, and has subsequently been reported from various localities along the

Pacific coast of North America (Banse and Hobson, 1968; Hartman, 1968; Rioja, 194I), the Gulf of Mexico R

(Perkins, 1981; Uebelacker, 1984), Cuba and the Gulf of Mexico region (San Martin, 1991b), and Belize by U

Russell (1991) who confirmed both Perkins' and Uebelacker's records, and also emended the species description.

i

Various morphological characters were measured (by L. Harris)on 100 specimens ofExogone lourei; B

individuals ranging from 16 to 49 setigers. For example, the proventficulus extends through a range of 2-7

m

segments, is present in 4 to 5 segments in 70 specimens, and usually occurs from setigers 3-4 to 7-8 in i

individuals with non-everted pharynges. Rows of proventricular muscle cells range from 16-30, and usually 1

number 18-24 in 94 specimens. Superior dorsal simple setae begin anywhere in setigers 1-19, and usually are

found in setigers 1-9. Natatory setae are first present from setigers 13-15, and usually begin after setiger 15. I

Gametes are detectable in setigers 8-28, and usually seen in setigers 10-14. These and other data need to be 1

further analyzed for size-related relationships. As one would anticipate, specimens with fewer than 27 setigers 0il

have smaller pharynges (2-3 segments) and proventricules (3-4 segments), dorsal simple setae from setiger 1,

and proventricules with the same number of muscle rows as adults. B

i

Exogone lourei is similar only to E. uniformis Hartman, 1961 in that both have compound spinigers

with long blades and thick shafts in setiger 2. There tends to be considerable overlap in individual key i

characters between these species. Exogone lourei can be distinguished from E. uniformis by the combined 1

presence of a median antenna much longer than the paired lateral antennae, 18-24 rows of proventricular in

muscle cells and a proventficulus in 4-5 segments, instead of short, subequal antennae, with 25 or more rows I

of proventricular muscle cells, and a proventriculus in 7-8 segments. Most of the paratypes ofE. uniformis i

are actually specimens ofE. lourei.

I

16 I
!

!

, i !

,,, !!/

!

B C D E F

I

i Figure 1.3. Exogone (Exogone) lourei: A, anterior end, dorsal view; B, dorsal simple seta from midbody region;

C, superior spiniger from midbody region; D, compound spiniger, setiger 2; E, inferior compound

falciger, with detail of blade, setiger 2; F, ventral simple seta. (A-F, after Uebelacker, 1984)

I

Distribution. British Columbia; Washington; Oregon; California; ?Mexico; Gulf of Mexico: Texas,

i Louisiana, Mississippi, Alabama, Florida; Cuba; Spain.

i Exogone (Exogone) dwisula Kudenov and Harris, new species

Figure 1.4

I Exogone uniformis Hartman 1961:73-74 (in part); Hartman 1968:427-428 (in part) ?Exogone gemmifera: Banse and Hobson 1974:56.

Sphaerosyllis californiensis Hartman, 1966:196-197, pl. 3 (in part).

I Material Examined. California: Santa Maria Basin, BRA-6 rock (1); BRA-13 rock, 2 paratypes

(AM W. 22190); BRC-13 rock, 1 paratype (SBMNH 142658), (1); BRC-14 rock (4); BRA-16 rock, 4

i paratypes (LACM-AHF Poly 1662), (2); BRA-20 rock (3); BRA-25 rock, 3 paratypes (LACM-AHF Poly 1663) + (2).--Western Santa Barbara Channel, Stas. BRC-1 rock, 7 paratypes (USNM 170913); BRA-1

rock (3); BRA-2 rock (1).--La Jolla, off South Casas Reef, rocky intertidal (from Phyllospadix wash), 11

i

i 17
I

I

¢

A B D (E V |

l |

''L I

J

I K |

A D-K I

.05 mm .005 mm

I

Figure 1.4. Exogone (Exogone) dwisula Kudenov and Harris, new species: A, anteriorsegments, dorsal

view; B, fight parapodium,setiger 2, ventralview; C, fightparapodium,setiger9, posteriorview; i

D, compoundfalciger with canopener-likeblades, setiger 1; E, superiorcompoundfalciger with l

can opener-likeblades,setiger 2; F, compoundfalciger,sefiger 5; G, dorsalsimple set& sefiger 1;

g

H-I,dorsalsimple setae:H, setiger 6, ventralview; I, sefiger 17,lateralview;J, acicula,setiger 1;

K, acicula, setiger 17. (A-K, originals by JDK) •

1

June 1979, coll. D.B. Cadien, holotype (LACM-AHF Poly 1661).--R/V Velero IV Sta. 5028-57, 3 paratypes 1

(removed from paratype lot of SphaerosylIis californiensis; LACM-AHF Poly 1659).--R/V Velero IV Sta. ll

5102-57, 2 paratypes (removed from paratype lot of Exogone uniformis; LACM-AHF Poly 171).

m

Description. Holotype 3.9 mm long, 0.2 mm wide without parapodia at setiger 6, for 34 setigers. I

Body fragile, semicircular in cross section, arched dorsally. Natatory setae and gametes first present from

n

setigers 8-9. Brooding females with a single pair of embryos on ventrum of each segment from setigers 8-9. 1

Prostomium rectangular; 2 pairs of eyes, lenticulate, anterior and posterior eyes on each side connate, !

anterior pair largest; posterior pair sometimes covered by fold of tentacular segment (Fig. 1.4A). Antennae

long, club-shaped to fusiform; median antenna about twice length of prostomium, arising from a point in front I1

of anterior pair of eyes; lateral antennae one-half to three-fourths length of median antenna, arising slightly l

anterior to median antenna (Fig. 1.4A). All antennae arise at same level with median antenna when prostomium

contracted. Palps wider than long, completely fused, usually arched, projecting ventrally (Fig. 1.4A). Nuchal Im

organs inconspicuous, ciliated, arising from anterolateral margin of tentacular segment, continuing as bilobed 1

nuchal glands internally. Pharynx usually extending through setiger 2(3), with large, anterior middorsal

tooth, surrounded by distal circlet of 10 soft papillae when everted (Fig. 1.4A). Proventriculus present in 2.5 Ill

setigers, usually from setiger 3 through middle of setiger 5, with 14-16 rows of large muscle cells (Fig. 1.4A). II

Intestine with muscular valve-like structure at junction with proventriculus (Fig. 1.4A). Tentacular segment

m

longer than prostomium and shorter than segments following, with 1 pair of small, ovoid tentacular cirri I

anterolaterally (not visible as illustrated). I
!

!

I Parapodial lobes stout, conical (Fig. 1.4B,C). Three types of compound setae present: (a) falcigers with long, deeply bifid blades (resembling a metal can opener in side view) numbering 7-10 per fascicle in

setigers I-3, all blades smooth, subequal within each of these first 3 fascicles, decreasing in length from

I setigers 1 to 3, with shafts terminating in strongly serrated edges, sometimes obscured by blades (Fig. 1.4D,E); (b) spinigers with long blades usually numbering 1, occasionally 2, as superiormost seta in fascicles from

setiger 4 to end of body, increasing to maximal lengths in middle segments, decreasing posteriorly where

I blades are shortest (Fig. 1.4C); and (c) falcigers with short comblike bidentate blades numbering 3 in most fascicles from setiger 4, decreasing to 2 in far posterior fascicles, blades decreasing in length along body (Fig.

1.4F); cutting edges all coarsely dentate from setiger 4 becoming smooth in posterior segments, with shafts

I terminally tapered tipsserrated. with close-set Superiortransverse dorsal simple rows ofsetaspines present (Fig. from1.4G-I). setigerInferior 1 to endventral of body,simple terminating seta present in abruptly from

midbody setigers to end of body; bifid, with subdistal serrations. Aciculae usually 1 per parapodium in all

I setigers, occasionally 2 in anteriormost setigers; allterminating in distally enlarged blunt heads (Fig. 1.4J-K). Dorsal cirri digitiform, present on all setigers except setiger 2, arising from body wall just above

parapodial lobes (Fig. 1.4A-C). Ventral cirri digitiform, at least as large as dorsal cirri, not projecting beyond

I parapodial lobes (Fig. 1.4B,C).

Pygidium with 2 long, cirriform anal cirri, about as long as last 3 setigers.

i Remarks. Exogone dwisula is characterized by the presence of two pairs of lenticulate eyes, pharynx in two segments, proventriculus in 2.5 segments with 14-16 well-defmed rows of muscle cells, lacking dorsal

cirri on setiger 2; with three kinds of compound setae including (a) compound falcigers with smooth deeply

i bifid blades distally in setigers 1-3, (b) usually a single compound spiniger with an awl-shaped blade next to the dorsal superior simple seta from setiger 4 to end of body, and (c) several compound falcigers with short

distally bidentate blades with subdistal tooth much larger than distal tooth. The bifurcate nature of the awllike blades is difficult to discern, even under oil immersion at the optimum viewing angle. E. dwisula inhabits

I snug, branching tubes composed of fine silt.

Exogone dwisula can easily be misidentified as E. gemmifera (Pagenstecher, 1862). Although we

I have of somenotofexami her materials ned all ofconfirms the materials the ease idenwith tifiedwhich by Hartman these twoas Especies . gemmife canrabe, aconfused. preliminaryA primary re-examipoint nationof

confusion is the difference in proventricle length between European and eastern north Pacific specimens.

Banse and Hobson (1974:56) suggest that the Pacific forms ofE. gemmifera "...probably should be assigned

I to a new species."

Exogone dwisula andE. gemmifera are similar species having peculiar can opener-like blades on all

I falcigers setigers, whereas of the firstE.two gemmifera or three specimens setigers. ExfromogoneScandinavia dwisula alwa haveys them seemsintosetigers have these 1-2 setae (L. Harris, in thepers. fit three obs),

while other European and Israeli specimens have them in setigers 1-3 (Ben-Eliahu, 1977). Exogone dwisula

I differs 10 compound from E. gemmifera in having a proventriculus of 2.5 segments rather than 1.5-2 segments in length; 7- falcigers per fascicle in the first 2-3 setigers instead of 4 for specimens of similar sizes; and in

having the cutting surfaces of dorsal superior simple setae heavily serrated rather than smooth. Indications

I are that San Martfn (1991 b) has identified another species (Exogone sp. A) that may also be a member of this species complex. Two specimens ofE. dwisula were found in a paratype lot of E. uniformis Hartman.

Etymology. The term, dwisula, is the Indonesian name for a 2-pronged lance head, and here refers to

I the deeply bifid or can opener-like blades of compound falcigers. Distribution. Southern California; possibly also Washington and British Columbia.

!

I 19
I

!

Subgenus Exogone (Parexogone) Mesnil and Caullery, 1916

Type species: Paedophylax hebes Webster and Benedict, 1884 I

Diagnosis. Prostomial antennae not all arising between eyes; compound setae all with similar I

unmodified blades of varying lengths, shaft tips and heterogomph joints; dorsal simple setae of one form,

distally unidentate or bidentate.

I

Key to the Known Species of the Subgenus Exogone (Parexogone)

from the Pacific Coast i

1A. Transition between long and short blades of compound falcigers in all setigers gradual, with superior i

blades 1.5 to 3 times longer than inferior blades within fascicles (Fig. 1.5C-E, K) ......................... 2

1B. Transition between long and short blades of compound falcigers in setigers 1-4 abrupt, with superior i

blades 3 to 4 times longer than inferior blades within fascicles (Fig. 1.6E, H, 1.7K,P) ....................

........................................................................................................ Exogone (Parexogone) molesta

!

2A. Superior blades of compound falcigers 3 times longer than inferior blades (Fig. 1.6E, H); pharynx in

6-8 segments; all dorsal simple setae distally unidentate (Fig. 1.6Q, R) ............................................ n

.................................................................................................... Exogone (Parexogone) acutipalpa •

U

2B. Superior blades of compound falcigers 1.5 times longer than inferior blades (Fig. 1.7K, P); pharynx i

in 4 segments; all dorsal simple setae distally bidentate (Fig. 1.7S-U) ..............................................

I

...................................................................................................... Exogone (Parexogone) breviseta

I

Exogone (P,,rexogone) moles_ (Ba_e, 1972)

Figure 1.5 i

Exogone molesta Banse, 1972:203-205, fig. 6.--Banse and Hobson, 1968:15-16, fig. 4 a-c; 1974:58, fig. 1

14k,1. I

Exogone (Parexogone ) molesta : San Mart/n, 1991a:725.

I

Material Examined. California: Santa Maria Basin, off Point Estero, Sta. 3 (3); off Point Sal, Sta.

36 (1); off Purisima Point, Sta. 42(1); east of Point Conception, Sta. 85 (1); off Point Sal, Sta. R-8 (2, USNM •

170917); offPoint Sal, Stas. PJ-1 (22); PJ-2 (7); PJ-3 (4); PJ-4 (3); PJ-5 (2); PJ-6 (3); PJ-7 (64); PJ-8 (24); 1

PJ-9 (2); PJ-10 (21); PJ-11 (24); off Point Arguello, Sta. PH-R (1).--Western Santa Barbara Channel, Sta.

BRC-1 rock (2, USNM 170916). II

Description. Specimens about 3 mm long, 0.2 mm wide excluding parapodia, 0.3 mm wide including 1

parapodia, for up to 40 segments (Banse, 1972). Sexually mature individuals with both natatory setae and

gametes first present from setiger 11. i

|
i

I

' • Figure 1.5. Exogone (Parexogone) molesta: A, anterior end, dorsal view slightly from the right side; B,

: I parapodium 12; C, falciger with long blade from setiger 4; D, falciger with medium long blade

from sefiger 16; E, falciger with short blade from setiger 4; F-G, dorsal simple seta from two

i positions from setiger 8; H, same, far posterior segment; I, ventral simple seta from posterior

setiger, J, acicula from setiger 16; K, parapodium 1, dorsal view; L, ventral simple seta, far posterior

setiger. (A-H, redrawn from Banse, 1972; I-L, originals by JDK)

I Prostomium wider than long; eyes oblong, numbering 2 pairs, anterior pair largest, arranged

trapezoidally; eyespots absent (Fig. 1.5A). Antennae digitiform; median antenna longer than prostomium, 7-

I pair 10 times of length of lateral antennae; lateral antennae not extending beyond eyes; all arising between anterior eyes (Fig. 1.5A). Palps long, distally pointed, fused completely, poorly deffmed from prostomium,

depending on state of contraction (Fig. 1.5A). Pharynx extending to setigers 3-4, with distal anterior middorsal

I tlarge ooth (Fig. muscle1.5A) ceils. . Prove Intestine ntriculuswith extending valve-like fromstructure setigersat jun4-5ctionthroughwith4-4proventriculus. .5 segments, withTentacular about 20segment rows of

fused with prostomium; 1 pair of tentacular cirri, each about size of lateral antennae.

I
!

II

Parapodia short, distally rounded, with small superior presetal lobe (Fig. 1.5B). Setae consisting of •

up to 4 superior compound falcigers with long urtidentate blades each in setigers 1-15 (25) (Fig. 1.5C,K); and

compound falcigers with blades short, distally bidentate, with secondary teeth strongly reduced (Fig. 1.5D,E); 1

alltogether numbering about 12 per anterior parapodia, decreasing to 6 in median, and 4 inposterior parapodia. I

Blades all increasing in length from anterior to median body region, decreasing thereafter to end of body, and

decreasing in length inferiorly within fascicles; long superior blades of anterior fascicles 3-4 times length of •

inferior blades, 4-5 times in median segments, and 1.25 times length in posterior segments. Long superior l

blades of median segments twice the length of anterior blades, 4 times length of blades in posterior segments

with transition between long medial and short posterior blades sometimes abrupt; terminating in blunt, curved, n

unidentate tips in setiger 1 to around setiger 20, and either short and abruptly bidentate thereafter, or gradually I

decreasing in length and forming a slight subdistal tooth along setigers 20-25, and then becoming short and

conspicuously bidentate at around setiger 25. Short blades in first few anterior setigers fairly uniform in •

length. Cutting edges of long superior falcigerous blades f'mely serrated; shorter falcigerous blades much 1

more coarsely serrated, particularly inferior ones. Dorsal simple setae present in all setigers, distally pointed,

unidentate, usually with slightly serrated cutting surface (Fig. 1.5F-H). Ventral inferior simple seta distally n

pointed, curved, usually unidentate, with slightly serrated cutting surface, from around setiger 25 to end of I

body (Fig. 1.5I), sometimes bidentate (Fig. 1.5L). One acicula per parapodium, distally blunt, knob-shaped

(Fig. 1.5]).

Dorsal cirri digitiform, present on all setigers except setiger 2 (Fig. 1.5A). Ventral cirri digitiform, 1

i not extending beyond parapodial lobes (Fig. 1.5B).

Pygidium with paired, digitiform anal cirri, each about as long as last 1-3 setigers, n

n

Remarks. Although Banse (1972) originally described the superior compound setae as being

spinigerous, they lack tapering capillary tips so characteristic of spinigerous blades. In fact, the blade tips of

these setae are all distally blunt and curved, and a secondary tooth is evident in posterior fascicles. Additionally, l

all other compound falcigers are more appropriately described as being distally bidentate rather than either

unidentate or subbidentate.

Exogone molesta and E. sexoculata Hartmann-Schrrder (1979) are similar in having a long median 1

antenna and short lateral antennae, long pointed palps, superior compound falcigers with long blades three

n

times the length of inferior blades within fascicles, two distinctly different populations of blades having

abruptly different lengths within the same fascicle, and all blades coarsely serrated. Exogone molesta differs •

from E. sexoculata in having two pairs of eyes instead of eyes plus an additional pair of eyespots, all simple

!

setae distally unidentate rather than bidentate, and apparently lacks a distal nipple-shaped parapodia lobe

present in E. sexoculata. 1

Distribution. Southern California; Washington.

Exogone (Parexogone) acutipalpa Kudenov and Harris, new species I

Figure t.6 I

Materials examined. California: Santa Maria Basin, off Purisima Point, Sta. BRA- 16 rock (1); Sta. •

R-5, 2 paratypes, (USNM 170910); off Point Sal, Stas. PJ-1, 1 paratype (SBMNH 142656) + (1); PJ-2, 1 l

paratype (AM W. 22188); PJ-6 (1); PJ-7,holotype (USNM 170908), 1 paratype (LACM-AHF Poly 1681) +

(5); PJ-10, 2 paratypes (USNM 170909), 5 paratypes (LACM-AHF Poly 1682-1683). •

Description. Holotype 2.6 mm long, 0.2 mm wide without parapodia atsetiger 6; 0.25-0.3 mm wide

with parapodia at setiger 6, for up to 40 setigers.

I
I

I

I

I A B _,

I

I c

I

E-W

.01 nun

I i

. MNoQR,, S T _l I _t

I

! U W

I

i Figure 1.6. Ex viewogone ; B, sam(Parexogone) e, dorsal vieacutipalpa w; C, right Kud parapodiu env mand1, Hdorsal aris, viewnew; D, specie lefts: pamp A, odium anterior 16e,nd,posterior dorsal

view; E-H, compound falcigers from setiger 2: E, superior seta with long blade; F, same, with

I shorter blade; G, lower intermediate seta; H, inferior seta; I-L, compound falcigers from sefiger 12"

I, superior seta with long blade; J, intermediate seta; K, low intermediate seta; L, inferior seta; ME compound falcigers from setiger 23: M, superior seta; N, upper intermediate seta; O, lower

intermediate seta; P, inferior seta; Q-R, dorsal simple setae: Q, setiger 1; R, setiger 23; S, ventral

I simpl (A-W,e osrietaginals frombysetigYDeKr ) 35; T-W, aciculae: T, sefiger 1; U, setiger 12; V, setiger 9; W, setiger 30.

I

• l 23

I"

I
!

Prostomium wider than long; with 2 pairs of eyes, lenticulate, arranged in trapezoid, anterior pair i

larger; posterior pair often hidden by overlapping tentacular segment; eyes fragment easily, multiple eyes

common; 1 pair of eyespots, situated just below junction of prostomium and palps (Fig. 1.6A,B). Three •

prostomial antennae, median one commonly 4 times longer than laterals but may be up to 10 times longer; all

fusiform (Fig. 1.6A,B). Palps long and pointed, length greater than prostomial length (Fig. 1.6A,B). Pharynx

in setigers 1-4, with distal anterior middorsal tooth (Fig. 1.6A). Proventricle in setigers 5-7, with about 20-23 •

rows of large muscle cells (Fig. 1.6A). One pair of tentacular cirri, smaller than lateral antennae, digitiform I

in shape (Fig. 1.6A,B).

Parapodia stout, conical, with smail lobe where aciculae emerge through body wall (Fig. 1.6C,D). II

Setae are compound bidentate falcigers (Fig. 1.6E-P), 12-15 in anterior fascicles, 6-8 in median fascicles, and

4-6 in posterior fascicles. Blades with subdistal tooth reduced, forming a highly acute angle to long axis of

blade; cutting edges with conspicuously variable teeth of different shape and size, usually slender, spine-like i

proximally, becoming saw toothlike distally (Fig. 1.6E-P); decreasing inferiorly within fascicles, with superior

blades 3 times length of inferior blades (Fig. 1.6C-E,H,I,L); along body axis, blade lengths gradually increasing

to medial segments, decreasing thereafter posteriorly. Dorsal superior simple seta first present from setiger 1, •

distally pointed, unidentate (Fig. 1.6Q,R), becoming slightly notched subdistally in posteriormost setigers

(Fig. 1.6S); all with cutting surface slightly serrated. Ventral inferior simple seta present in posteriormost

setigers, resembling dorsal simple setae, except for subdistal notch (Fig. 1.6T). Aciculae with curved tips, •

numbering 1 per parapodium (Fig. 1.6U-X). Dorsal cirri short, digitiform, present on all setigers except I

setiger 2, shorter than length of parapodial lobes (Fig. 1.6C,D). Ventral cirri short, digitiform, similar in size

to dorsal cirri (Fig. 1.6C). •

Pygidium with paired digitiform anal cirri of variable lengths, each about as long as last 1-3 setigers,

plus smailer ventromedian cirrus.

Remarks. Exogone acutipalpa is most easily confused with E. molesta Banse in the northeastern i

d

Pacific since both species have a long median antenna, long pointed palps, and superior falcigers with i

conspicuously tong blades. Exogone acutipalpa differs from E. molesta in having a pair of eyespots in

addition to two pairs of eyes instead of lacking the paired eyespots, and in having blades of all compound 1

falcigers gradually decreasing in length within all fascicles rather than having two groups of blades having

abruptly different lengths in anterior and median fascicles. i

Habitat. Soft, fine sediments in depths of 91.5-154 m. i

i

Etymology. The specific epithet, acutipalpa, refers to the extremely pointed palps.

Distribution. Southern California. i

Exogone (Parexogone) breviseta Kudenov and Harris, new species 1

Figure 1.7

Exogone sp. C SCAMIT Harris, 1987:1-8. i

Material Examined. California: Santa Maria Basin, off Point Sal, Sta. PJ-7 (1).--Western Santa

Barbara Channel, Goteta, 301(h) Sta. 2, 24-25 m, 14 April 1985, 1986, coll. MBC, 3 paratypes (LACM- i

AHF Poly 1678); Sta. 3, 25 m, 14 April 1986, coll. MBC, 2 paratypes (USNM 170911); Sta. 4, 29 m, 29

i

October 1985, coll. MBC, 1 paratype (USNM 170912); Sta. 6, 22-23.5 m, 14-15 April 1986, 1 paratype I

(SBNMH 142657), 4 paratypes (LACM-AHF Poly 1679-1680); Sta. 7, 23 m, 14 April 1986, coil. MBC, 1 i

paratype (AM W. 22189); Goleta, NPDES, Sta. 4, 29 m, 29 October 1985, coll. MBC (i).--Santa Catalina I

Island, Catalina Ouffail, Sta. 1, 38 m, 2 November 1986, coll. B. Given (4); Sta. 3, 38 m, 2 November 1986, I

coll. B. Given, holotype (LACM-AHF Poly 1666). i

24 i
I Figure 1.7. Exogone B, pharynx(Parexogone) and setiger breviseta 1, ventral Kudenov view; C, anandterior Harris, end,newdorsspecies olateral: viAew, ant ; eDr,iolrefend t para , dors podium al view1;,

dorsal view; E, left parapodium 12, posterior view; F, right parapodium 25, anterior view; G-I,

i susetpeigrior er 4:compound K, superiorfalcige seta;rs L-Mfrom, upset

peigr

erint2;

e

J

rm, samediatee

fromsetae;setigN-P, er 3

c

;

oK-M, mpoundcomfpoalcigundersfalfromcigersse

from tiger

25: N, superior seta; O, intermediate seta; P, inferior seta; Q, superior seta from midbody segment;

R, same from fax posterior segment; S-T, dorsal simple setae: S, from setiger 2; T, from midbody;

I U, from far posterior segment; V-Y, acieulae: V, from setiger 1; W, from midbody segment; X, from setiger 25; Y,from fax posterior segment. (A-Y, originals by JDK)

I
!

am

Description. Holotype 1.8 mm long, 0.1 mm wide without parapodia at level of setiger 6, 0.2 mm •

wide with parapodia atlevel of setiger 6, for 33 setigers. Specimens up to 3.8 mm long. Surface pigmentation

patterns absent; pharynx sometimes golden in color. Eggs first occur in setigers 9-10 and continue towards •

end of body; one complete mature specimen of 28 setigers with natatory setae in setigers 11-27. II

Prostomium wider than long; eyes small, lenticulate, numbering at least 2 pairs, each pair frequently

divided, forming separate multiple pairs, all arranged trapezoidally on posterior part of prostomium; with an n

additional pair of eyespots on anterior surface ofprostomium (Fig. 1.7A,C). Antennae fusiform, close-set, as

transverse group between posterior pair of eyes (Fig. 1.7A,C). Median antenna about twice length of

prostomium; lateral antennae ranging from one-fifth to one-half length of median antenna (Fig. 1.7A,C). n

Palps pointed, longer than prostomium (Fig. 1.7A,C). Nuchal organs ciliated, located laterally atjunction of

prostomium and tentacular segment, continuing internally as bilobed glands. Pharynx extending through

setiger 6, sometimes to setiger 8, with anterior middorsal tooth (Fig. 1.7A,B). Proventriculus present in •

setigers 7-8, sometimes setigers 8-9, with 20-23 rows of large muscle cells (Fig. 1.7A). One pair of tentacular

cirri, similar in shape to lateral antennae, smaller (Fig. 1.7A,C).

Parapodia stout, conical, with small postsetal lobe (Fig. 1.7D-F). All setae compound falcigers with

short, strongly serrated, distally bidentate blades (Fig. 1.7G-R). Blades decreasing in length within fascicles,

superior blades less than twice length of inferior blades; all increasing in overall length from anterior to

median setigers, decreasing thereafter posteriorly (Fig. 1.7C-E). Superior dorsal simple seta present in all •

setigers, inconspicuously bifid in anterior setigers (Fig. 1.7S), becoming thick and strongly bifid with serrated

cutting surfaces in posterior setigers (Fig. 1.7T,U). Inferior ventral simple seta present in median and posterior

setigers, distally bifid, particularly in posterior setigers. Acicula with distally bent tip, 1per parapodium (Fig.. •

1.7V-Y).

Dorsal cirri digitiform, shorter than parapodial lobes, present on all setigers except setiger 2 (Fig.

1.7A,E,F). Ventral cirri similar to dorsal cirri in shape, slightly smaller (Fig. 1.7E,F). I

Pygidium with paired digitiform anal cirri of variable lengths, each about as long as last 1-3 setigers,

plus smaller ventromedian cirrus. n

Remarks, Exogone breviseta has been misidentified as E. molesta Banse by having long pointed I

palps, a long median antenna, and short lateral antennae. Exogone breviseta differs from E. molesta in

lacking superior falcigers with long blades, and in having compound falcigers all subequal in length throughout ID

the body, an anterior pair of prostomial eyespots, and a longer pharynx and a shorter proventriculus. Exogone |

breviseta has also been confused withExogone acutipalpa, described above, in that both have a long median

antenna, long distally pointed palps, two pairs of eyes plus anterior paired eyespots and blades of compound m

falcigers coarsely serrated. Exogone breviseta differs fromE, acutipalpa in having the posterior two pairs of |

eyes small and divided into smaller separate structures, in having superior falcigers all with short rather than

long blades, and in having a considerably shorter pharynx. ID

Exogone breviseta co-occurs with E. lourei, E. molesta and E. uniformis. I

l]abitat. Coarse, mixed and soft sediments in depths of 20-200 m.

Etymology. The species name, breviseta, refers to the short blades on all compound falcigers. I

n

I

Distribution. Southern California.

Subgenus Exogone (Sylline) Clapar&le, 1864 I

Type species: Sylline brevipes Clapar_de, 1864

Diagnosis. Antennae closely set together, usually arising between eyes; compound falcigers with

strongly modified bayonet-shaped blades partially fused to heterogomph joints; shaft tips usually truncate;

dorsal simple setae of one form, generally distally truncate with serrated cutting surfaces. U

:6 |
I

I Remarks. Both species in the key below lack dorsal cirri on setiger 2, and appear to have prostomia

generally fused to the tentacular segment. Exogone (Sylline) naidinoides is not represented in MMS materials,

I although it is reported from the northeast Pacific; E. (SyUine)fustifera Haswell is reported from Australia.

iIll Key to Selected Species of the Subgenus Exogone (SyUine) (after San Martin, 1991b)

I

1A. Bayonet-shaped compound falcigers from median segments including long and short blades ...........

i 1B. Bayonet-shaped ........................................................................................................... compound falcigers from midbedy segments thin withExogone blades of(Sylline unifo)rmnaidinoides length ....

................................................................................................................ Exogone (Sylline) fustifera

!

Genus Sphaerosyllis Clapar_de, 1863

I

'_1 Type species: SphaerosyUis hystrix Clapar_de, 1863

I Diagnosis. antennae. Palps dorsally most or length. One pair

Prostomium with 3 fused for entire of

tentacular cirri. Nuchal organs inconspicuous. Pharynx with single anterior middorsal tooth. Antennae,

tentacular cirri and dorsal cirri usually flask-shaped (pyriform). Integument usually with glandular adhesive

I papillae.

Remarks. The majority of SphaerosyIlis species lack dorsal cirri on setiger 2 (San Martin, 1984a-b,

i 1991a; Riser, 1991). San Martin (1984b) relied on this and other features to define subgenera: Prosphaerosyllis San Martin, 1984b, for species with dorsal cirri on setiger 2; and SphaerosyIlis Clapar&te, 1863, for species

lacking dorsal cirri on setiger 2. The subgenus S. (Sphaerosyllis) is further subdivided into the "'erinaceus"

and "'pirifera" groups. The "erinaceus'" group is partly characterized by having aciculae with tips flared and

I mucronate, posterior eyes supposedly arranged in a straight transverse row, and have not been reported

the

from the eastern Pacific; the "pirifera'" group has aciculae with tips shaped like golf clubs, and eyes arranged

polygonally. Riser (1991 )suggests that the above subgenus concept may be inadequate in view of additional

I described since San Martin first this scheme. Russell does San species proposed (1989a) not recognize

Martin's (1984b) subgenera since they are "...based more on artificial than on important phylogenetic

characters." We concur with both Russell (1989a) and Riser (1991 ), and find the usefulness of San Martin's

I sphaerosyllid subgenera limited.

Riser (1991) also provides an important critique of systematic characters forSphaerosyIlis. Perhaps

i one of his most illuminating statements is that exogonine syllids (including Sphaerosyllis species that have dorsal cirri on setiger 2 as adults) usually Jack dorsal cirri on setiger 2 prior to developmental differentiation

of fore-gut elements. Identification of juvenile specimens will be difficult if one relies primarily on dorsal

cirri. Care must therefore be taken to consider the age of one's specimens, especially sinceSphaerosyUis that

I on setiger as adults have a pair of non-deciduous papillae that are larger than general body

lack dorsal cirri 2

papillae.

i 1966; S Three species of Sphaerosyllis have been encountered in the MMS collection: S. californiensis Hartman, . bilineata, new species; and S. ranunculus, new species. Sphaerosyllis brandhorsti HartmannSchr6der, 1965, is included in the key below, although it was originally described from Chile, and later

i reported from Washington (Banse, 1972). Banse's record is based on an incomplete specimen which differs

|
.

from Hartmann-Schr/Sder's original account in potentially significant ways; it seems unlikely that S. (S.) i

brandhorsti occurs in the northeast Pacific. Unfortunately, we have not examined Banse's specimen. Records

ofS. hystrixfrom the northeast Pacific are probably based on Berkeley and Berkeley (1938), who distinguished i

this species by the presence of glandular pits containing bundles of minute rods in each segment. In contrast,

S. californiensis was traditionally characterized by lacking these parapodial glands. However, subsequent

examinations of S. californiensis type specimens confirms their presence (see below), and worms previously •

identified as S. hystrix from southern California are all S. californiensis. Moreover, Riser (1991) records S. II

californiensis from British Columbia, and it therefore seems likely that all records ofS. hystrix in the northeast

Pacific are either misidentified specimens orS. californiensis, or represent atleast one undescribed species of I

Sphaerosyllis. In view of these findings, S. hystrix is excluded from the following key. !1

Key to Species of SphaerosyUis from i

the Northeast Pacific

!

1A. Dorsal cirri similar to one another .................................................................................................. 2

1B. Dorsal cirri of two forms, not similar to one another ....................................................................... 3 i

2A. Proventriculus usually present through 1.5-2 segments (Fig. 1.8A); dorsum with dense fields of filiform II

and elliptical papillae (Fig. 1.8C-D); ventrum lacking papillae; laterally with 2 pairs of conspicuous Ii

papillae per segment, each pair associated with basal anterior and posterior facies of parapodial lobes

.............................................................................................................. Sphaerosyllis californiensis •

2B. Proventriculus present in 4 segments; dorsum with scattered, sparse fields of filiform papillae; ventrum

with papillae; papillae inconspicuous on parapodia .................................. Sphaerosyllis brandhorsti

!

3A. Dorsum with 2 conspicuous dorsomedial longitudinal rows of distally knobbed macropapillae, 2

pairs per segment from setiger 4 or 5, alternating between a large anterior and smaller posterior pair

per segment (Fig. 1.9A); ventrum with 12digitiform papillae in 4 longitudinal rows of 3 papillae plus i

2 additional longitudinal rows of 4 small, round papillae per segment, these altogether forming 6

i

longitudinal rows (Fig. 1.9C); dorsal cirri flask-shaped anteriorly, becoming cirriform posteriorly. I

Sphaerosyllis bilineata Di

3B. Dorsum without obvious dorsomedial longitudinal rows of papillae; ventrum with 2 pairs of papillae

per segment, all arranged in 2 longitudinal rows; dorsal cirri mammiform in setigers 1-7 or 9 (Fig. i

1.10D) and far posterior segments, digitiform in medial segments from setigers 8 to 10, or all digitiform I

from setigers 8-10 (Fig. 1.10E) ................................................................ Sphaerosyllis ranunculus

Sphaerosyllis californiensis Hartman, 1966 I

Figure 1.8

==

Sphaerosyllis californiensis Hartman, 1966:196-197, pl. 3; 1968:453-454, figs. 1-7. i

Material Examined. California: R/V VeIeroSta. 5028-57, holotype (LACM-AHF Poly 863) and47 •

paratypes (LACM-AHF Poly 864).---Southeast of Point Conception, Sta. 86 (1, USNM 170926).--Santa 1

Barbara, Camby's Reef, rock scraping, 16 m, 15 April 1983, coll. D. B. Cadien (2).---Gulf of the Faraltones,

Cordell Bank, Sta. 831033, coll. B. Schmieder (1). i
I Figure 1.8. Sphaerosyllis californiensis: A, anteriorsegments,dorsal view (Goleta 6/HI);B, prostomium,dorsal

view; C, leftparapodium, setiger 7, anterior view; D, left parapodium, setiger 30, anterior view; E,

fight parapodium, setiger 22, posterior view (LBGS B8/m); F-G, superior compound falcigers, setiger 2; H, inferior compound falciger, setiger 2; I-K, compound falcigers from setiger 3 l: I,

superior seta; J, intermediate seta; K, inferior seta; L-N, dorsal simple setae: L, setiger 28; M,

sefiger 32; N, setiger 33; O, ventral simple seta, setiger 3t; P-Q, aciculae: E sefiger 2; Q, setiger

I 31; R, Posterior end,dorsal view. (A-R, originals by JDK)

I Description. Largest complete MMS specimen 3.7 mm long, 0.9 mm wide excluding parapodia and

setae, for 40 setigers. Preserved specimens lack pigmentation. Integument covered with fine sediment (I-Iamnan,

I short 1966).elliptica Dorsulmpapillae usually(updensely to 8 pmpapil long); late, allincpapillae luding con gradually spicuouins creasing filiform inpapil length lae, posteriorly up to 22 pmalong long,body. and

Ventrum smooth, lacking papillae. Lateral margins of setigers with long papillae, 2 pairs per segment, each

I pair 10 toassociated setiger 22with or sometimes anterior andalmost posteri tooend r faces of body. of basal parapodial lobes. Epitokous segments from setiger

Prostomium oval in outline, 3 times longer than wide, broadly fused dorsally with tentacular segment;

eyes numbering 2 pairs, lenticulate, in trapezoidal to rectangular arrangement onposterior half of prostomium, anterior eyes largest; eyes connate (Fig. 1.8A,B). Median antenna slightly longer than lateral antennae,

similar in shape, arising from anterior emargination of tentacular fold; lateral antennae arising anterolaterally

I from prostomium, with bulbous bases and cylindrical tips, extending to edge of palps (Fig. 1.8A,B). Palps
!

short, dorsally fused; distal median notch as a wide furrow (Fig. 1.8B); palps covered dorsally with short i

papillae. Pharynx usually present in setiger 1 to middle of setigers 3-4 (in setigers 1-3.5 in holotype), with

large anterior middorsal tooth (Fig. 1.8A). Everted pharynx surrounded by 10 widely spaced papillae. In

Proventriculus barrel-shaped, with 13-14 transverse rows of muscle cells, usually present in setigers 5-6 (in •

setigers 3.5-5 in holotype) (Fig. 1.8A), sometimes in setigers 3-5 or 4-6, or in setigers 4-8 in highly contracted

i

specimens. I

Tentacular segment visible dorsally asfold covering posterior half of prostomium, including posterior H

pair of eyes and sometimes anterior pair of eyes. Tentacular cirri similar to lateral antennae, arising

ventrotaterally in relation to anterior pair of eyes (Fig. 1.8A,B). Small papillae present on anterior margin of •

tentacular fold. |

Parapodial lobes conical, distally blunt (Fig. 1.8C-E), each with 3 conspicuous distal papillae as

follows: longest papilla on posterior basal surface of parapodium; shortest papilla on anterior distal surface; •

intermediate papilla on posterior distal surface. Parapodial glands present from setiger 5, conspicuous fi'om II

setigers 7-8, medial to dorsal cirri, containing spherical yellow granules which can fade and be undetectable

in some preserved specimens. •

Compound setae numbering 5-7 per fascicle in anterior segments, 3-5 in posterior segments; blades

unidentate, longer in anterior segments, becoming stouter, more triangular posteriorly. Superior blades of

anterior segments coarsely (sometimes finely) serrated, 2-3 times longer than smooth inferior blades (Fig. II

1.8F-H); those of posterior segments subequal, superior blades finely serrated to smooth, inferior ones smooth

(Fig. 1.8I-K). Ends of shafts with 2-3 denticles on cutting surface plus additional hair like processes (Fig.

1.8F,G). Dorsal simple seta present in all setigers, slightly curved subdistally; anterior simple setae smooth, i

appearing finely hirsute when highly worn; posterior setae shorter with serrated cutting surfaces (Fig. 1.8L,M).

Inferior simple seta small, smooth, present in last 2-19 setigers (Fig. 1.8N,O). Aciculae normally numbering

one per parapodium, thick, tapering abruptly to tip bent distally at a right angle (Fig. 1.8P), gradually becoming •

more stout posteriorly (Fig. 1.8Q). Additional acicula sometimes present in setigers 1-7, slender, distally

pointed. Natatory capillary setae long, smooth, in setal sac located between neuropodia and dorsal cirri (Fig.

1.8E). Notoaciculae absent. •

Dorsal cirri similar to antennae in shape, slightly thinner basally, absent from setiger 2 (Fig. 1.8A);

longer than anterior parapodial lobes, equal to or somewhat shorter than median and posterior parapodial

lobes. Ventral cirri short, cylindrical, sometimes extending beyond parapodial lobes (Fig. 1.8C-E). •

Pygidium hemispherical, subdistally encircled by about 10 long papillae, with 2 long clavate anal

cirri, these slightly larger than nearest dorsal cirri, plus an additional cluster of smaller papillae surrounding

anal opening, sometimes obscured by circlet of longer papillae or anal cirri. H

Remarks. Re-examination of type materials revealed the presence of 2 specimens of an undescribed

Sphaerosyllis species (described below), a specimen of Exogone lourei, and 3 specimens of E. dwisula, as

noted above. While this f'mding does not alter the species concept of S. californiensis, Hartman's original •

description appears to be based on one ortwo type specimens; new illustrations are here provided for additional

information.

Riser (199t) examined west coast specimens of Sphaerosyllis pirifera Clapar_de, 1868, and found i

that nearly all were S. californiensis. SphaerosyIlispirifera appears not to be represented along the western

coast of North America (Riser, 1991:216) as first reported by Berkeley and Berkeley (1948). In addition, i

examination (by LHH) of specimens identified and archived by O. Hartman both as S. hystrix Claparbde, |

1863 and S. pirifera were also found to be S. caIiforniensis. It is likely that S. hystrix is not represented in the

polychaete fauna of southern California. Additional specimens previously identified as S. hystrix from other i

localities along the western coast of North America need to be re-examined to determine whether itis represented •m

in our fauna as also indicated by Berkeley and Berkeley (1948).

Habitat. Silt, mixed soft sediments and rocky subtidal habitats. •

Distribution. Southern California. II

30 !
!

: I SphaerosyUis bUineata Kudenov and Harris, new species

Figure 1.9

!

Sphaerosyllis californiensis Hartman, 1966:196-197, pl. 3; 1968:453 -454, figs. 1-7 (in part).

I Material Examined, California: Santa Mafia Basin, off Purisima Point, Stas. BRA-13 rock, pamtype

(USNM 170924); BRC-13 rock (1); BRC-14 rock, paratype (AM W. 22193); BRA-16 rock, 11 pamtypes

I (LACM Point Conception, -AHF PolySta. 1664, 84, paratype 1665); BRA (USNM-27 (SBMNH 170925); offPoint 142699)..--Western Conception, Sta. SantaBRC-2 BarbarroackChannel, (1); San Francisco south of

Bay, off Hunters Point, ESA Homeporting, Sta. HPIB, 37°43'30.8"N, 122°21'25.8"W, 9 m, 8 September

i pamtypes 1986, coll.(removed Kinnetic Laboratories, from paratype Inc., lot ofholotype Sphaerosyllis (LACMcaliforniensis -AHF Poly; 1673).--R LACM-AHF /V Velero Poly IV1675). Sta 5028-57, 2

Description. Holotype complete with 31 setigers, 3 mm long, 0.3 mm wide without parapodia, 0.4

I mmyellowwideduewitho pencrusted ampodia. silt. BodyOneslender, femaledorswithum dorsally convex, laattacking chedpigmentatio embryos nonpsetigers atterns; dorsum 11-17. Two appears sexually light

mature males with sperm in setigers 8-24 or 26.

i i Prostomium wider than long, with small conical papillae dorsally and laterally; 2 pairs of eyes, large,

• lenticulate, 2 eyes on each side set close together; posterior pair usually covered by dorsal fold of tentacular

segment (Fig. 1.9A); additional pair of anterior eyespots present adjacent to lateral antennae (not illustrated).

I Median antenna arising from posterior margin of prostomium, on border of dorsal fold; lateral antennae smaller than median, cylindrical with bulbous bases (Fig. 1.9B). Palps longer than wide, anterior margins

blunt, directed ventrally. Small, conical papillae present on dorsal and lateral margins of palps, and anterior

I marg 1.9A).in Pharynx of tentacular fold. Ciliated nuchal organs present between prostomium and tentacular segment (Fig. tong, narrow, thick-walled, extending to setiger 3-4, or setigers 2-6 in contracted specimens,

often appearing sinuous inpreserved specimens; middorsal tooth large, positioned one-quarter length of pharynx

I 4behind segments, anterior barrel-shaped pharyngeal with margin 20-23wherows n everted of muscle (Fig.cells 1.9A). (Fig.Proventri 1.9A). cTenta ulus cnormally ular segment extending and prostomium through 3-

fused dorsally. Tentacular cirri similar to prostomial antennae, but smaller (Fig. 1.9B).

I Parapodial lobes triangular, each with 2 digitiform papillae, on anterior face and distally on posterior face (Fig. 1.9C). Setae generally compound falcigers, numbering 6-7 per fascicle anteriorly, decreasing to 5

posteriorly. Blades unidentate, longest anteriorly, gradually decreasing in length posteriorly (Fig. 1.9D);

I superiormost blade (sometimes 2 superiormost blades) in each fascicle with finely serrated cutting margins, others smooth. Superior dorsal simple seta distally unidentate, with smooth cutting surface, in all setigers.

Inferior simple seta unidentate, slightly serrated to smooth, in last 2-8 setigers. One acicula per pampodium,

I slender, subdistaUy enlarged, bent. Dorsal cirri flask-shaped on setiger 1, about equal to or slightly larger than prostomial antennae;

larger, longer from setiger 2, becoming cirriform posteriorly, extending beyond parapodial lobes (Fig. 1.9AI B). Ventral cirri digitiform, not extending beyond parapodial lobes (Fig. 1.9C).

Each segment with 1 or 2 transverse rows of papillae, with 4 papillae per row (Fig. 1.9A). Dorsum

of setigers 1-3 or 4 each with 4 filiform papillae in single transverse row per segment; setigers 4-5 to end of

I each with 8 filiform in transverse Middle 2 of all body papillae an anterior-posterior rOW. papillae segments

distally knobbed, together forming two dorsomedian longitudinal rows along body; those of anterior segmental

rows from setiger 4 largest, alternating with shorter papillae in posterior rows to end of body. Smaller outer

I papillae of anterior segmental rows arising from body wall superior to parapodial lobes and anterior to dorsal

cirri; those of posterior segmental rows located just anterior to intersegmental annuli. AU papillae gradually

i increasing in length posteriorly. ,I
I 

I

I

A B

I

' I

m

Figure 1.9. SplmerosyUisbilineam Kudenov and Harris, new species: A, anteriorsegments, dorsal view; B, •

prostomium and setigers 1-2, lateral view; C, ventrum, median setiger; D, superior compound

falciger,setiger 8. (A-D, originals by J. Dorsey) _.

!

i

Ventrum with 20 papillae per segment in 6 longitudinal rows, including 12 digitiform papillae in 4

longitudinal rows of 3 papillae, and 2 longitudinal rows of 4 short round papillae (Fig. 1.9C).

ll Pygidium with paired anal cirri, these 2-3 times longer, and 3-4 times wider than dorsal cirri of last

setiger; circlet of 8-10 small, digitfform papillae present above anal cirri.

Remarks. SphaerosyUis bilineata, S. IongipapiUata Hartmann-Schr6der, 1979, S. riseri Perkins !

1981, andS. tetalix Eliason, 1920 are similar in having four papiUae per segment, all arranged in two longitudinal

m

rows along the dorsum. All species also have an additional pair of papillae per segment, each in front of the

dorsal cirri on the body wall superior to the parapodia. Sphaerosyllis bilineata differs from the other listed •
!

!

taxa in having one pair of small papillae per segment in setigers 1-3 or 4, and two pairs of different sized

I papillae that alternate regularly from setigers 4-5 to the end of the body; papillae are the same size in all segments for the other three species exceptS, longipapiUata which lacks papillae from the two dorsal rows in

setigers 1-6. Sphaerosyllis bilineata further differs from the above listed species in having an additional pair

i ofshort small papillae papillaearedorsolaterally the most conspicuous along thefeature posteriorof margin S. bilineata of each .segment. The alternating rows of long and

The compound falcigers of both S. bilineata and S. riseri differ from those ofS. longipapillata and S.

I tetralix in having superior blades all serrated instead of all smooth. Sphaerosyllis bilineata differs from S.

riseri in having slightly longer blades, and 20-23 rows of proventricular muscle cells rather than 17-18.

Habitat. Rocky substrates in depths of 89-126 m.

I Etymology. The species name, bilineata, refers to the paired dorsomedial rows of conspicuous papillae.

Distribution. California.

I SphaerosyUis ranunculus Kudenov and Harris, new species

I Figure 1.10

Material Examined. California: off Point San Luis, Sta. R-2 (14, USNM 170930); off Point Sal,

i Stas. 4, 5 paratypes PJ 1(46); PJ-2 (27, including 1 brooding 2 setiger larvae); PJ-3, 3 paratypes (AM W. 22194) + (11); PJ- (SBMNH 142661) + (3); PJ-5 (9); PJ-6 (42); PJ-7 (47); PJ-8, 15 paratypes (USNM 170929)

+ (36); PJ-9 (31); PJ-10 (67); PJ-11, 35 paratypes (LACM-AHF Poly 1668) + (21).--U.S. Army Corps of

i EApplied ngineers,Environmental dump site LSciences, A-5, Sta. holotype BD 5-4(3), (USNM32°32'20"N, 170927).--Between 117°20'35"W,Point 188Sanm, Pedro 12 Dec. and1983, Pescadero coll. Point, MBC

FTSP-Corps of Engineers, Stas. B5-W(3), 37°29'21 "N, 122°55'32"W, 128 m, 10 October 1986, coll. Kinnetic

i Laboratories, Inc. (KLI), (1); B5-W(4), 5 paratypes (USNM 170928); B2-W(5), 37°22'10"N, 122°49'27"W, 128 m, 11 October 1986, coll. KLI (1); B2-N(1) 37°23'09"N, 122°49'34"W, 126 m, 2-3 April 1986, coll.

KLI, 5 paratypes, (LACM-AHF Poly 1667).--Mexico: Tijuana, Tijuana Oceanographic Engineering Study,

Sta. B10(3), 32°35.74'N, 117°10.34_N, 18 m, 18 July 1986, coll. MBC Applied Environmental Sciences, (1).

I Description. Holotype incomplete, 1.3 mm long, 0.15 mm wide including parapodia, for 23 setigers.

Body slender, circular in cross section in atokous specimens; oval to dorsoventrally flattened in epitokous

I specime papillae,n.usually Dorsum with micropapillae (best observed in lateral view). Ventrum with 2 longitudinal rows of two pairs per segment; each lower papilla on ventrolateral margin loosely associated with

parapodia (Fig. 1.10D). Color in alcohol light yellow; pigmentation absent.

I equal inProsize, stomium arrangedsmall, trapezoidally; micropapillanterior ate; 3 pairs pair ofin eyes, front al ofl lateral lenticulatentennae ; posterior (Fig. 21.10A-C). pairs largest, Antennae nearly

small, inconspicuous, rnammiform (basally spherical, abruptly becoming distally papilliform); median antenna

I arising eyes; each between aboutorone-half slightlysize behiofndmedian posteriorantennayes; (Fig. lateral1.10A-C). antennae arising Palps small, betweefused n antcompletely erior and mi(Fig. ddle 1.10A- pair of

C), directed ventrally, micropapillate. Pharynx extending to middle of setiger 3, with middorsal tooth in

i anterior 3to end qofuarter setigerwhen 6 when inverted pharynx (Fig.inverted; 1.10A). inProve 4 segme ntricunltsuswhen in 3.5pharynx segments, everted; extending with from aboutmiddle 20 (19-22) of setiger rows

of muscle cells (Fig. 1.10A). Retractor muscle visible through body wall, especially along right side of body

in PJ 1/3; running from body wall around level of setiger 2 to intestinal region posterior to proventriculus (Fig.

I 1.10A).

!
I

!

A,,_, _ B C I

I

I

TI '

I

I

FGHI J KI ItNO |

LM

!

Figure 1.10. Spprostomium, haerosyUis doranunculus Kudenov and Harris, new species: A, anterior end, dorsal view; B, I

rsal view; C, prostomium, ovigerous female, dorsal view; D, right parapodium,

setiger 7, anterior view; E, right parapodium, setiger 16, anterior view; F-G, compound falcigers

from setiger 4: E superior seta; G, inferior seta; H-I, compound falcigers from setiger 5: H, inferior

seta; I, superior seta; J, compound falciger, setiger 20; K, compound falciger, setiger 22; L, dorsal I

simple seta, sefiger 18; M, ventral simple seta, setiger 23; N, acicula, setiger 15; O, acicula, setiger

19. (A-O, originals by JDK)

i

!

Tentacular segment fused to prostomium, not visible dorsally (Fig. 1.10A,C); 1 pair of tentacular

cirri, similar in form and size to antennae (Fig. 1.10B,C). Parapodia uniramous, triangular, distally pointed

in atokous specimens; stout, thick in anterior segments (Fig. 1.10D), becoming longer, narrower in middle and

posterior segments (Fig. 1.10E). Compound falcigers present in all setigers, all unidentate (Fig. 1.10F-K); J

numbering around 7-8 per anterior fascicle, around 3-5 in posterior fascicles. Blades of setigers I-3 decreasing

in length inferiorly within fascicles, cutting edges all finely serrated; blade lengths of following setigers more I

variable, with tendency for superiormost to have somewhat more coarsely serrated cutting margins, and m

inferiormost blades to be smooth (Fig. 1.10F-I). Blades of posterior sefigers usually smooth, slightly shorter

than those in anterior segments (Fig. 1.10J,K). Superior dorsal simple seta first present from setiger 1 to end I

34 !
!

I of body, distally unidentate, with serrated cutting surfaces (Fig. 1.10L). Ventral inferior simple seta first

present in last 8 or fewer posterior segments, distally unidentate, smaller than dorsal simple seta, generally

i smoot tip tightly h (Fig. covered 1.10M). andOne readily acicula visible per parapodium, through cuticle; distally distallypointed, flexed,projecting with "flat"through surfaceparapodium, representingalthough dorsal

surface (Fig. 1.10N,O).

I 1.10D), Dorsal commonlycirribecoming on all setigers, digitiformsmall,fromapparently stiger contractile; 8-10 to end ofthosebodyof(Fig. setigers1.10E); 1-7 orsometimes 9 mammiformremaining (Fi.

mammiform on all setigers or having mammiform cirri on anterior and posterior segments, digitiform cirri

i medially. 1.10D,E). Ventral cirri small, digitiform, arising basaUy from and not extending beyond parapodial lobes (Fig.

Pygidium micropapillate, with paired anal cirri, surrounded by circlet of small digitiform papillae.

I Length of other specimens at least 2.5 nun, 0.15 mm wide excluding setae, for 24 setigers (PJ-2(3));

all incomplete. Epitokous specimen (PJ- 1(3)) with biramous parapodia beginning in setiger 8 and continuing

through last setiger, each with a dense fascicle of long, fine capillary setae emerging from notosetal sacs

I located between dorsal cirri and parapodia. Gametes first visible from setiger 8; 4 external eggs carried dorsally per segment, with dorsomedial rows easily detached, rarely present on preserved specimens.

t Remarks. Most Phase II MMS specimens here identified as Sphaerosyllis ranunculus were originally identified as S. brandhorsti Hartmann-Schrfder, I965. Based on the present samples, S. brandhorsti is

probably not present along southern California. SphaerosylIis ranunculus differs from S. brandhorsti in

having minute prostomial antennae and tentacular cirri compared to relatively large, conspicuous appendages,

I in clavate dorsal cirri of similar size 7 thereafter instead having through setiger generaIly becoming digitiform

of being clavate in all body segments, and in having blades of inferior compound falcigers smooth rather than

serrated. Sphaerosyllis brandhorsti was originally described from Chile, and was subsequently reported

I from Orcas Island, Washington, on the basis of single specimen Banse (1972:209). Although Riser

a by

(1991) suggests that the degree of fusion between the prostomium and tentacular segment is an unreliable

character in the systematics of Sphaerosyllis, Banse notes that his specimen differs from the types of S.

I brandhorsti by the fusion of these two areas. Banse further notes the absence of epidermal papillae in his specimen. These same differences exist in the present materials compared to the original description of S.

brandhorsti, and lead us to doubt Banse's identification.

I Habitat. Soft sediments, including isolated rocks in soft sediments, in depths of 18-128 m.

Etymology. The term, ranunculus, isLatin for tadpole, and refers to the body shape of these specimens.

I Washington. Distribution. Southern California, San Diego to San Francisco; central Oregon; Puget Sound

I Subfamily Eusyllinae

I Genus Dioplosyllis Gidholm, 1962

Type Species: Dioplosyllis cirrosa Gidholm, 1962

I Diagnosis. Prostomium with 3 antennae. Palps large, lingutate, fused basally. Nuchal organs as

transverse ridges when present. Two pairs of tentacular cirri. Parapodia long. Pharynx with a middorsal

I anterior tooth Remarks and additional smaller teeth, with a smooth rim, or with a few teeth.

. Although Dioplosyllis octodentata Perkins, 1981 is included in the key below, it was

originally described from the east coast of Florida, and may occur at other localities in the Gulf of Mexico

I (Uebelacker, 1984).
!

Key to the Species of Dioplosyllis from California N

1A. Compound falcigers with distally bidentate blades with minutely serrated cutting edges (Fig. 1.11E); I

pharynx in 2 segments; proventriculus in 3-4 segments, with about 90 rows of muscle cells ...........

......................................................................................................................... Dioplosyllis lagunae

1B. Compound falcigers include distally tridentate blades with conspicuously serrated cutting edges (Fig. I

1.12D); pharynx in 7-8 segments (Fig. 1.12A); proventriculus in 5-7 segments with 30-33 rows of

muscle cells (Fig. 1.12A) ................................................................................................................ 2 n

2A. Compound falcigers tridentate blades enshrouded by distal hoods (Fig. 1.12D); distal teeth of blades I

aligned parallel to long axis of blade; shaft tips of compound falcigers distally pointed, entire ........ I

....................................................................................................................... Dioplosyllis tridentata

2B. Compound falcigers include bi-, tri- and multidentate blades, distal hoods absent; distal teeth of I

blades paired to unpaired, depending on state of wear; shaft tips of compound falcigers deeply notched II

and distally bifid ........................................................................................ Dioplosyllis octodentata

im

Dioplosyllis lagunae (Hartman, 1961), new combination m

Figure 1.11 I

u

Orseis lagunae Hartman, 1961:64-65; 1968:371-372, figs. 1-2.

Dioplosyllis broadi Mueller and Fauchald, 1976:19-22, figs. 1-6. N

Material Examined. California: Santa Catalina Island, Big Fishermans Cove Pier, holotype of iln

Dioplosyllis broadi (LACM-AHF Poly 1141).--Laguna B each, intertidal, holotype of Orseis lagunae (LACM- l_

AFH Poly 135). II

Description of Atokous Form. Length at least 15 mm, 2.5 mm wide excluding parapodia, 5.2 mm ll

wide including parapodia; an anterior fragment with 10 segments. Body short, inflated (Hartman, 1961). I[

Prostomium wider than long; 2 pairs of eyes, large, conspicuous; anterior pair set farthest apart,

posterior pair near postectal margins, all trapezoidally arranged. Antennae long, slender, cirriform. Median m

antenna arising just anterior to anterior eyes. Lateral antennae anterior to anterior eyes. Palps long, distally I!

inflated, projecting ventrally.

Parapodia thick, triangular. Setae compound falcigers with distally bidentate blades. Blades decreasing _l

in length inferiorly in fascicles, superior blades twice the length of inferior blades; angle between two distal II

teeth large, with subdistal tooth projecting outward obliquely.

Dorsal cirri long, slender, cirriform. Ventral cirri similar in form to dorsal cirri, extending beyond 1

parapodial lobes. U

Description of Epitoke Form. Length up to 24 mm, 2.1 mm wide excluding parapodia, 6.1 mm 1

wide with parapodia, for 13 setigers (Mueller andFauchald, 1976). Body widest medially, tapering anteriorly I[

and posteriorly; dorsum strongly arched; ventrum flat. In live specimens, pigmentation on dorsum generally

brownish purple, ventrum lighter; segments with paired series of transverse ridges, each purple ridge separated

from neighboring ridges by cream colored grooves, 2 per tentacular segment, increasing to 12 on setiger 4, 17- U

18 on setiger 8 (Fig. 1.11A). Parapodia with ridges running parallel to body axis, each ridge purple with I

white spots (Fig. 1.11A). Dorsal and ventral cirri each with 10 longitudinal stripes, consisting of series of

small purple spots. 1

U

36 I
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| A "

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Figure 1.11. Dioplosyllis lagunae: A, anterior end, dorsal view; B, prostomium and anterior segments, dorsal

I view; setigerC,7,p(hA-Earynxred;D, rawnsetifrger om7,Mante uellerrioanrview; dFauchal E, codm,1976) pound falcigers, setiger 7; F, neuroaciculae,

I

Prostomium wider than long; 2 pairs of eyes, subequal, posterior pair lenticulate, anterior pair set

_: • farthest apart, all arranged trapezoidally (Fig. 1.11A,B). Antennae long, wrinkled to smooth with short

ceratophores; median antenna arising between anterior and posterior eyes, extending posteriorly to setiger 6;

lateral antennae arising from anterior margin of prostomium, extending posteriorly to setiger 5 (Fig. 1.11A).

I Paired twice length nuchaloforgans prostomiurn, at postectal fusedcobasally rners of(Fig. prost1.11A,B); omium (Fig. palpostyles 1.11A,B).present. Palps thick Pharynx , rountodsetiger ed, more2, with than

subdistal anterior tooth plus 5 evenly spaced curved teeth, surrounded by distal circlet of 10 soft papillae

I when everted (Fig. 1.11C). Proventriculus present in setigers 3-6, with about 90 rows of muscle cells. Tentacular segment conspicuous dorsally (Fig. 1.11A,B). Two pairs of tentacular cirri, wrinkled to

smooth, similar in form to prostomial antennae; dorsal pair Longest, each extending posteriorly to setiger 8;

I ventral pair about half as long as dorsal ones; all with cirriphores.

Parapodia long, triangular (Fig. 1.11D). Compound setae bidentate falcigers numbering 25-30 per

fascicle (Fig. 1.11E). Blades longest superiorly, decreasing in length inferiorly within fascicles, with minutely

I serrated cutting edges (Fig. 1.11E). Shaft tips distally bifid, with denticles on subdistal superior surfaces

• (Fig. 1.11E). Neuroacicula varying in thickness, tapering abruptly, 6-7 per parapodium from setiger 6 (Fig.

1.11F). Capillary setae long, from setiger 6. Notoacicular lobe inferior tofascicles of capillary setae, numbering

I 6-7 per fascicle from setiger 6.

Dorsal cirri long, cirriform, smooth to wrinkled, those of setigers 1-2 extending to setiger 11-12; all

with short, conspicuous ceratophores (Fig. 1.11A). Ventral cirri long, cirriform, extending to middle of

I adjacent setigers (Fig. 1.11D).

Pygidium with paired anal cirri, long and cirriform.

!
!

Remarks. The single type specimen described as Orseis lagunae Hartman, 1961 from Laguna i

Beach is not a hesionid and is here referred to Dioplosyllis. Hartman's (1961) original description lacks

illustrations; Hartman (1968) depicted two compound falcigers that looked similar to those of D. broadi i

presented by Mueller and Fauchald (1976). Comparisons of the types of both O. lagunae and D. broadi by g

one of us (L. Harris) confirms their synonymy.

Habitat. Intertidal and shallow subtidal rocky substrata, i

I Distribution. Southern California.

Dioplosyllis tridentata Kudenov and Harris, new species i

Figure 1.12 i

II Material Examined. California: off Purisima Point, Sta. BRC-13 rock, holotype (USNM 170900).

Description. Hoiotype 5 ram, 0.5 mm wide without parapodia, 1 mm wide with parapodia. Body Ill

broken into anterior piece with 21 setigers; posterior piece with 12 setigers of which latter 4 or 5 regenerating. I

Pigmentation patterns absent.

Prostomium wider than long, pentagonal, with deep median cleft (Fig. 1.12A); 2 pairs of eyes, •

lenticulate, anterior pair largest, in trapezoidal arrangement (Fig. 1.12A); a minute pair of eyespots present on II

anterior prostomium margin, medial to lateral antennae. Antennae smooth, cirriform, longer than prostomium;

median antenna about 5 times longer than prostomium, arising between anterior and posterior pairs of eyes; •

lateral antennae about twice length of prostomium, arising medial to each anterior eye. Palps long, flat, thick, II

about 1.5 times longer than prostomium, basally fused, projecting ventrally, lacking palpostyles (Fig. 1.12A).

Nuchal organs paired transverse ridges along postectal border of prostomium (Fig. 1.12A). Pharynx tubular, gin

extending through setiger 8 when inverted, middorsal tooth subdistal (dissected) (Fig. 1.12A); distal circlet of II

soft papillae not observed. Proventriculus extending through setigers 9-15, with about 30 rows of large

muscle cells. II

Tentacular segment narrow, with 2 pairs of long, cirriform tentacular cirri, each with short cirrophores; i

dorsal pair probably longest, each atleast as long as median antenna (remaining right tentacular cirrus distally

broken); paired ventral cirri broken off. i

Parapodia long, slender, distally conical to triangular, gradually increasing in length to middle segments, U

decreasing in length thereafter (Fig. 1.12C). About 15-18 compound setae per anterior fascicle (ca. setiger 5),

increasing to 20-25 per medial fascicle (ca. setiger 15), decreasing to around 17-19 per posterior fascicle (ca. i

setiger 29). Blades distally bidentate, largest denticle strongly curved, enshrouded by distal hood; blades U

longest superiorly, just over twice length of inferior blades within same fascicle, all decreasing in length

inferiorly within fascicles; all with minutely serrated cutting edges. Shaft tips distally entire, with denticles II

scattered on subdistal superior surfaces. Neuroaciculae taper to knobbed tips, not projecting through parapodial i

lobes, numbering 1 per parapodium.

Dorsal cirri long, smooth, cirriform (mostly broken), all with short cirrophores, arising from body H

wall; right dorsal cirrus of setiger 1 present, extending posteriorly to around setigers 13-14. Ventral cirri g

glandular, basally inflated, tapering distally; not extending beyond parapodial lobes, short in anterior segments,

becoming longer, better developed in middle segments, decreasing in length posteriorly, i

i

Pygidium with paired anal cirri, all regenerating.

Remarks. Dioplosyllis tridentata and D. octodentata Perkins, 1981, are similar in having tridentate i

compound falcigers, in contrast to the four other described Dioplosyllis species summarized by Mueller and l

Fauchald (1976), which have bidentate compound falcigers. Dioplosyllis tridentata differs fromD, octodentata

in having distal hoods on blade tips; in having minute rather than coarse serratia on the cutting edges of !

38 !
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II A

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t tl

/

1

i .....

il

A D E-G /

i ._--_-.o_ .O,n_, G F E D

!

I Figure 1.I2. Dioplosyllis prostomium tridentata and setigerKudenov 1, lateral and view;Harris, C, left new parapospecies dium:, mA,edanter ian sior egmen segments t, post,eridoorrsal viewview; ; D-EB,,

supcrior compound falcigers; F, inferior compound falciger; G, acicula. (A-C, E, F, redrawn from

I originals by J.Dorsey; D, original by JDK)

!

i blades; in having shaft tips of compound falcigers distally entire instead of bifid and deeply notched; in having the pharynx extend through eight rather than seven segments; in having the proventriculus extend through

seven segments with around 30 rows of muscle segments, instead of five or seven segments with about 33

rows. The pharyngeal armature of these two species may also differ, although it is difficult to clearly examine

I the pharynx of D. tridentata without damaging the single specimen. For example, D. seems to

tridentata

have a single large subdistal tooth whileD, octodentata has a single large tooth plus seven smaller ones. The

i margin of the pharynx in D. tridentata appears to be smooth.

I 39
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Dioplosyllis tridentata is readily distinguished from D. lagunae from southern California, besides I

the differences in compound falcigers noted above, in lacking palpostyles; the pharynx extending through 8

instead of 3-4 segments and the proventriculus through 6-7 rather than 3 segments; with around 30 instead of

around 90 rows of muscle cells in the proventriculus; and in having shaft ends of compound falcigers distally _l

entire instead of deeply notched or bifid. II

Habitat. Rocky substrata.

Etymology. The name, tridentata, refers to the presence of 3 teeth on blades of compound falcigers. I

Distribution. Southern California.

It

Genus Eusyllis Malmgren, 1867 I

Type species: Eusyllis blomstrandi Malmgren, 1867 I

Diagnosis. Prostomium with 3 antennae. Palps fused basally. Nuchal organs usually as ciliated

ridges between prostomium and tentacular segment. Occipital flap sometimes present. Antennae, tentacular i

and dorsal cirri smooth to indistinctly articulate. Pharynx with middorsal tooth and denticulate rim. Setae i

compound falcigers.

Remarks. The presence of a denticulate pharyngeal margin or rim is critical to the systematics of 1

Eusyllis, and must be determined before specimens can be correctly identified to this genus. Specimens with =l

retracted pharynges must therefore be dissected to avoid confusion with the closely similarPionosyllis.

1

Key to Species of Eusyllis from California HI

(after Imajima, 1966c)

1A. Ventral cirri of setiger 1 foliaceous, much larger than those following (Fig. 1.14K); marginal rim of 1

pharynx smooth dorsally, denticulate ventrally (Fig. 1.14B) ...................................... Eusyllis habei

lB. Ventral cirri ofsetiger 1similar in form and size to those following; marginal rim of pharynx uniformly 1

denticulate ....................................................................................................................................... 2 1

I

2A. Dorsal cirri of middle body segments 2-3 times longer than body width (Fig. 1.15C); blades of •

compound falcigers long and narrow (Fig. 1.15E,F) ....................................... EusyIlis longicirrata

2B. Dorsal cirri of middle body segments two-thirds length of body width (Fig. 1.13A,B); blades of m

compound falcigers short and stout (Fig. 1.13E-G) ......................................... Eusyllis blomstrandi •

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I Eusyllis blomstrandi Malmgren, 1867

Figure 1.13

I Eusyllis blomstrandi Malmgren, 1867:159.--Augener, 1928:721 .--Annenkova, 1938: 153, fig. 6.--Fauvel,

1923:293-294, fig 112 h-m.--Wesenberg-Lund, 1947:11-13, fig. 3; 1950a: 16; 1950b:48; 1951: 37.--

I Berkeley 120, fig. 31andn-p;Berkeley, Imajima, 1948:84-85, 1966c:92-94,fig.text-126.--Pettibone, fig. 29, a-h.--Banse 1954:260-261, and Hobson,fig.1968:15-16, 28 g-i.--1963:119- figs. 4 ac; 1974:56, figs. 14 b-c.--Hartmann-Schr6der, 1971:158, fig. 52.

I Eus Typosyllis yllis bloemstrand collaris Hartman i Friedrich1948:23 1938:, 1fig. 22 6(misspe a-c. lling of E. bIomstrandi).

I Materials D examined, California: Santa Maria Basin, Sta. R-1 (1, USNM 170903). escription. Specimens 10-12 mm long, 0.8-1.2 mm wide, with 50 segments; single MMS specimen

an anterior fragment 3.5 mm long, 0.5 mm wide excluding parapodia, with 28 setigers. Body slender, pale

I brown 1968). in alcohol; each segment with single, dorsal transverse ciliary band (Imajima, 1966c; Banse and Hobson,

Prostomium wider than long, anterior margin somewhat rounded; 2 pairs of eyes, red, in trapezoidal

i arrangement, anterior pair largest; additional pair of red eyespots usually present on anterior prostomium margin (Fig. 1.13A). Antennae smooth to wrinlded proximally, distinctly articulated distally; median antenna

arising between largest pair of eyes, around 5 times longer than prostomium; lateral antennae, arising from

I anterior edge of prostomium, each about one-half length of median antenna (Fig. 1.13A). Palps broad, flat, triangular, slightly fused basally (Fig. 1.13A). Pharynx thick, about as wide as proventTiculus; anterior rim

freely denticulate, with anterior middorsal tooth, surrounded by distal circlet of 10 soft papillae when everted.

I Proventriculus extending from setigers 7-8 to 13-16. Tentacular segment reduced dorsally, forming low occipital flap covering posterior prostomium margin.

Two pairs of tentacular cirri, proximally smooth, weakly articulated distally; dorsal pair longest, each about

I as long as median antenna; ventral pair about half as long as dorsal ones (Fig. 1.13A).

Parapodia bluntly conical (Fig. 1.13B-D). Setae generally compound falcigers (Fig. 1.13E-G); blades

short, distally bidentate with cutting edges serrated. Blades decreasing in length inferiorly within fascicles;

I segments (Fig. 1.13E), becoming more stout posteriorly (Fig. 1.13F,G), especially in

those of anterior slender

inferior pan of fascicles. Superior distal ends of shafts minutely serrated (Fig. 1.13E-G). Dorsal superior

and ventral inferior seta present infascicles of posterior segments. Two to 3 aciculae per anterior parapodium,

I decreasing per posterior parapodium; distally pointed, mallet-shaped (Fig. 3H).

to I curved, 1D 1

Dorsal cirri basally smooth to wrinkled, weakly to distinctly articulated distally (Fig. 1.13A-D).

I Dorsal cirri of setiger 1 longest, longer than median antenna, with 40-50 articles (Fig. 1.13B); setiger 2 dorsal cirri short, about one-half length of first pair; cirri on setigers 3 and 4 long; on setiger 5 short; on setiger 6

long; on setiger 7 to end of body about two-thirds body width. Ventral cirri thick, foliaceous, extending

i beyond parapodial Remarks. lobes The present (Fig. 1.13B-D). record of Eusyllis blomstrandi is based on a single anterior fragment from

Phase 1I. Although this species is widely reported from the west coast of North America (Berkeley and

I Berkeley, pharyngeal1945; morphology Banse andcompared Hobson, 1968, 1974; Hartman, 1968), Banse and Hobson (1968) noted different to that described by both Malmgren (1867) and Imajima (1966c). We

suspect that specimens identified as E. blomstrandi at least from southern California represent an undescribed

species. Efforts to address this issue must await additional, well-preserved specimens and re-examination of

i Malmgren's specimens.

Distribution. California to Alaska; Japan; Labrador Sea to Massachusetts; Spitsbergen; Iceland;

I Ireland; Mediterranean Sea.

I 41
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E F G H I

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Figure 1.13. Eusytlis bloomstrandi: A, anterior end, dorsal view; B, parapodium 1; C, parapodium 15; D,

parapodium 17 from epitokous specimen; E, inferior compound falciger, parapodium 1; F, inferior I

compound falciger, middle body segment; G, superior compound falciger, middle body segment; H, I

aciculae, parapodium 1. (A-H, modified and redrawn from Hartmann-Schr6der, 1971)

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I Eusyllis habei Imajima, 1966

Figure 1.14

I Eusyllis habei Irnajima, 1966c:97-99, text-fig. 31, a-k.

Material Examined. California: between Point San Luis and Point Sal, Sta. BRA-25 rock (5,

I USNM 170904) + (2); off E1 Morro, Sta. BRA-27 (5, SBMNH 142700); east of Point Conception, Sta.

BRC-01 rock (1); off Point Conception, Sta. BRA-02 rock (2); off point San Luis, Stas. R-1 (2, USNM

170905); R-8 (2, USNM 170906); off Point Arguello, Stas. PH-F (5); PH-I (1); PH-U (1); PH-W (4).

I Description. Length up to 5 mm, 0.9 mm wide, for 43 setigers 0majima, 1966c); MMS specimens

all incomplete, 2.1 mm long, 0.2 mm wide excluding setae, for 24 setigers. Body pale yellow white in alcohol.

I Some Japanese specimens with 2 dorsal transverse color bands per segment (Imajima, 1966c); most MMS specimens with 1 black transverse band on anterior edge of each segment, bands sometimes incomplete

middorsally. Body fragile.

i pair Prostomium wider than long; 2 pairs of eyes, each circular in outline, arranged trapezoidally, anterior set farthest apart (Imajima, 1966c); MMS specimens with conspicuous reddish-brown eyes (Fig. 1.14A).

Antennae weakly annulated; median antenna arising between anterior pair of eyes, 4-5 times longer than

! • prostomium; lateral antennae arising from anterior prostomium margin, each about one-half length of median

' I antenna (Fig. 1.14A). Nuchal ridges paired, extending broadly along posterior prostomial border. Palps

about as long as prostomium, distally rounded, basally fused (Fig. 1.14A,K). Pharynx extending through

i setiger 6 when inverted, setiger 3 when everted; ventral rim of everted pharynx with 20-26 denticles, dorsal rim smooth with large, distal, spindle-shaped middorsal tooth, all surrounded by distal circlet of 10 soft

papillae (Fig. 1.14B). Proventriculus present in setigers 6 to 9-10, with 23-24 rows of muscle cells, anterior

i 20 of theseTentacular large, conspicuous. segment reduced dorsally. Two pairs of tentacular cirri; dorsal pair each about as long as

median antenna; ventral pair each about one-third length of dorsal pair (Fig. 1.14A).

I serrated Parapodia cutting edges short,(Fig. thick, 1.14D-G). conical Blades (Fig. 1.decreasing 14C). Compo in length und setae inferiody bidentate withinfaleigers fascicles, withposte minute riorlyly

along body axis. Superior blades of anterior, middle segments oar-shaped, 2-2.5 times longer than inferior

I blades Anterior(Figs. setigers D,E);eachthosewithof 10-15 posterior supesegments rior falcigers shortwith er, just longslightly bladeslonger and 20-30 than infe inferior rior falcigers blades (Figs. with F,G). short

blades; median setigers each with 10 falcigers. Dorsal superior and ventral inferior simple seta present in

I poste distallyriorbidentate setigers; (Fig. dorsalI).seta Onecapillariform to 2 aciculaewith in antecutting rior parapodia surface minutely and 1aciserrated cula in (Fig. medianH),parapodia; ventral setadistally stout,

blunt (Fig. J).

I Dorsal cirri of setiger 1 longest, 2 times longer than median antenna, alternating long and short thereafter; long cirri equal to and short ones one-half body width (including parapodia). Ventral cirri of setiger

I foliaceous, flattened; digitiform from setiger 2 to end of body; all shorter than parapodial tobes.

I Remarks. This is the first record of Eusyllis habei from the western coast of North America. The MMS specimens generally conform to features previously described, although they are smaller.

Distribution. Southern California; Japan. !

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f

K I I

H I

I

i

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Figure 1.14. Eusyllis habei: A, anterior end, dorsal view; B, pharynx, dissected; C, parapodium 18; D, superior 1

compound falciger, middle segment; E, inferior compound falciger, middle segment; F, superior B

compound falciger, posterior segment; G, inferior compound falciger, posterior segment; H, dorsal

simple seta, posterior segment; I, ventral simple seta, same posterior segment as H; J, acicula, m

median parapodium; K, anterior end, ventral view. (A-K, redrawn from Imajima, 1966c) I

I

44 |
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Eusyllis longicirrata Imajima, 1966

I Figure 1.15

EusyUis longicirrata Imajima, 1966c:94-97, text-fig. 30, a-f.

I Material Examined. California: between Point San Luis and Point Sal, Sta. BRA-25 rock (3,

USNM 170907).

I Description. Length upto 18 mm, 1.2 mm wide for 87 setigers (Imajima, 1966c); all MMS specimens anterior fragments up to 2 mm long, 0.5 mm wide excluding parapodia for 21 setigers.

Prostomium wider than long, anterior margin slightly rounded, posterior middorsal cleft deep, extending

I to center of prostomium (Fig. 1.15A) 2 pairs of red, anterior farthest Antennae eyes, pair largest, apart. long,

smooth, slender. Median antenna arising from middle of prostomium, 4 times longer than prostomium; lateral

antennae arising from anterior prostomial margin, each one-half length of median antenna. Nuchal organs

I paired, extending along posterior margin of prostomium (Fig. 1.15A). Palps longer than prostomium,

narrow,

distally rounded, not fused basally (Fig. 1.15A). Everted pharynx extending through 5 or 8 segments in two

MMS specimens, with anterior rim minutely denticulate, with middorsal subdistal tooth, surrounded by distal

i circlet of 10 soft papillae. Proventriculus in setigers 13-21 (Imajima, 1966c; Fig. 1.15B); MMS specimens with proventriculus in setigers 9-17 or 6-14 (8 segments) with 32-34 rows of muscle cells.

Tentacular segment reduced dorsally. Two pairs of tentacular cirri; dorsal pair as long as median

I antenna; ventral pair each one-half length of dorsal tentacular cirri.

Parapodia elliptical in outline, distally rounded (Fig. 1.15C). Setae compound falcigers (Figs. D,E).

i Blades distally bidentate with cutting edges minutely serrated; decreasing in length inferiorly within fascicles, superior blades two times longer than inferior blades. Two to 3 aeiculae per parapodium, distally pointed,

slightly curved (Fig. 1.15F).

I tentacularDorsal cirri (Fig. cirri 1.15A,C); with long alternating cirriphore, long smooth, and short considalong erablybody. moreLong slenddorsal er, longer cirri ofthananterior both antennae, segments

up to 4 times longer than body width, decreasing to 3 times in middle body segments; short dorsal cirri 2 times

I bbeyodoynwidth d parapthroodial ugholout, bes.gradually decreasing in overall length posteriorly. Ventral cirri digitiform, extending

Remarks. This is the first record of Eusyllis longicirrata from the eastern Pacific. The MMS

I specimens generally conform to features previously described, although they are smaller. Distribution. Southern California; Japan.

I Genus Odontosyllis Clapar_de, 1863

i I Type species: Syllisfulgurans Audouin and Miine-Edwards, 1833

g

Diagnosis. Prostornium with 3 antennae. Palps either fused basally or separate. Nuchal organs

I lflap arge,usually curvedprese ciliated nt. Antennae, ridges alontentacular g posteriorcirri margin and dorsal of prostcirri omium. smooth. TwoPharynx pairs of with tentacfewer ular cirri. than 20Occurved cipital

teeth.

I Pacific cRemoast arks of North . TheAmerica key presented from California below inctoludes Alaska. all known Only Odontosyllis species, and phosphorea one new species, and Ofrom . fragilisthe

were encountered in MMS materials.

I
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II

II

II

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Figure 1.15. Eusytlis longicirrata: A, anterior end, dorsal view, distal parts of antennae, tentacular and dorsal i

cirri omitted; B, pharynx dissected; C, parapodium l 1, D, superior compound falciger, parapodium

11; E, inferior compound falciger, parapodium 11; F, aciculae, parapodium l 1. (A-E redrawn from

Imajima, 1966c) i

I

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Key to Known Species of Odontosyllis from the Northeast Pacific

I (after Banse and Hobson, 1974)

I 1A. Median in dorsalantenna view ...................................................................................................... arising from anterior prostomium margin; palps small, projecting ventrally, Odontosyllis not visib parvale

1B. Median antenna arising from dorsal surface of prostomium near anterior pair of eyes (Figs. 1.16A,

I 1.17A); palps large, projecting anteroventrally, visible in dorsal view (Figs. 1.16A, 1.17A) .......... 2

2A. Proventriculus extending through about 20 segments; blades of bidentate compound falcigers short,

I serrated in anterior fascicles and smooth in cutting edges minutely posterior fascicles; preserved

specimens lack color patterns ......................................................... Odontosyllisfulguransjaponica

i 2B. Proventric and short,uwith lus extending cutting edges throughall 10serrated or feweror segments; all smooth;blades preserved of bidespecimens ntate compouusually nd falcigers with dorsal long

pigmentation ..................................................................................................................................... 3

I 3A. Proventriculus extending through 9-10 segments with 60-67 rows of muscle cells; blades of compound

falcigers with serrated cutting edges (Fig. 1.16E,G); dorsum with dark pigment spot every third to

I fourth intersegmental groove or with black transverse line on each of the first 20 segments (Fig. 1.16A); occipital flap with or without black pigment; pharynx with 6 distal teeth (Fig. 1.16B) .......

................................................................................................................... Odontosyllis phosphorea

I 3B. Proventficulus extending through 12 segments with approximately 300 rows of small, dense muscle cells; blades of compound falcigers with smooth cutting edges (Fig. 1.17F,G); middorsum with a dark

pigment spot in middle of every segment including tentacular segment (Fig. 1.17A); occipital flap

I without black pigment (Fig. 1.17A); pharynx with 5 distal teeth (Fig. 1.17E)

......................................................................................................................... Odontosyllis fragilis

I Odontosyllis phosphorea Moore, 1909

i Figure 1.16 Odontosyllis phosphorea Moore, 1909:327.--Berkeley and Berkeley, 1948:82.--Hartman, 1961:76-7;

1968:441-442, figs. 1-3.--Banse, 1972:207-209, fig. 7G-N.--Banse and Hobson, 1974:58, fig. 15cI d.--Blake, 1975:189, pl. 35, fig. 133. Odontosyllis phosphorea nanaimoensis E. Berkeley, 1923:207; 1961 :1321 .--Berkeley and Berkeley,

1948:82.--Pettibone, 1967:5.

I Hesperalia nans Chamberlin, 1919:9. Not Hesperalia californiensis Chamberlin, 1919:9. Fide Hartman, 1961:9.

Not Odontosyllis phosphorea Berkeley and Berkeley, 1938:42. Fide Banse, 1972:207.

i Material Examined. California: Santa Mafia Basin, Sta. BRA-16 rock (1); between Point Estero

and Point Buchon, Sta. 6, rock (4, SBMNH t42701).--Laguna Beach, holotype ofHesperalia californiensis

I Chamberline, dried 1919, ovigerous (MCZ 2149); holotype of Hesperalia rums Chamberlin, 1919, ovigerous, (MCZ 2150).---Santa Catalina Island, type of Odontosyllis phosphorea (USNM 17214).--British

Columbia: Nanaimo, syntypes of O. phosphorea nanaimoensis (15, USNM 32858).

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Figure 1.16. Odontosyllis phosphorea: A, anterior end, dorsal view; B, teeth and lateral chitinized regions of I

pharynx, frontal view; C, anterior parapodium; D, medial biramous parapodium from epitokous

specimen;E, compoundfalciger, setiger 10; F,same, setiger 70; G, same, from inferiorfascicle; H, •

dorsalsimple seta, posterior segment; I, ventral simple seta, posteriorsegment;J, acicula, setiger 1; I

K, same, posterior segment; L, same, setiger 70. (A, modified from Blake, 1975; B, E-F, H-L,

redrawn from Banse, 1972; C-D, G, redrawn from Moore, 1909) i

Description of Epitokous Form. Length 20 to 30 mm, 2.5 mm wide, for about 90 segments (Moore,

1909; Hartman, 1968); MMS specimens up to 9.5 mm long, 0.7 mm wide excluding parapodia, for 69 •

setigers. Dorsum arched, ventrum flattened. Color in life variable: pale yellow to ivory with dark or black I

middorsal spot every third or fourth intersegmental groove or as a black transverse line on each of first 20

setigers followed by faint middorsal pigment spot in each successive medial segment; occipital flap darkly •

pigmented. |

Prostomium small, wider than long; 2 pairs of eyes, black; anterior pair widest apart; posterior pair

lying in crescent-shaped patch of dark pigment; all arranged trapezoidaUy (Fig. 1.16A). Median antenna •

small, smooth, arising between anterior pair of eyes; lateral antennae smooth, subequal in length and arising Hi

slightly anterior to median antenna (Fig. 1.16A). Palps thick, fleshy, projecting ventrally, fused basally.

Everted pharynx extending to setiger 6-8, with circlet of 6 distal teeth (Fig. 1.16B). Proventriculus from •

setigers 7-9 to 17-19; muscle rows ranging from 60-67, depending on body size. I

Tentacular segment reduced dorsally, obscured by prostomium, with semicircular occipital flap

extending forward to cover anterior pair of eyes (Fig. 1.16A). Two pairs of tentacular cirri, smooth to i

wrinkled; dorsal pair longest, about 3 times longer than prostomium; ventral pair about one-half length of II

dorsal pair (Fig. 1.16A).

Parapodia short; uniramous, distally rounded in anterior segments (Fig. 1.16C); biramous in median

segments with notopodial lobes triangular, neuropodial lobes rounded (Fig. 1.16D) Setae including compound

falcigers with distally bidentate blades with serrated cutting margins, about 30 per anterior fascicle at setiger

10 (Fig. 1.16E), decreasing thereafter to about 24 per median fascicle, and 18 per posterior fascicle (Fig. i

1.16F,G). Blade lengths subequal within anterior medial and posterior fascicles; decreasing slightly in overall

length from anterior to posterior setigers. Dorsal simple seta distally unidentate, wide, present in last 2-4

setigers of complete specimens (Fig. 1.16H). Ventral simple seta distally bidentate, present in last few posterior •

48 i
!

I setigers (Fig. 1.16I). Capillary swimming setae long, from setigers 21-22. Neuroaciculae 4 per anterior

parapodia (Fig. 1.16J), decreasing to 2 in median and posterior parapodia (Fig. 1.16K,L). Notoacicula thin,

I tapering Dorsal distally,cirri deeplysmooth embedded, with short 1 percirripbores; notopodiumalternating of median long segments. and short. Ventral cirri thick, distally

blunt, projecting slightly beyond parapodial lobes in median segments.

I Remarks. Odontosyllisphosphorea is a well-defined species commonly associated with rocky habitats from western Canada to southern California that is readily collected using night lights. Banse (1972) reported

the cutting edges of compound falcigers in specimens he examined to be serrated, in contrast to Moore's

I (1909) originally observations and illustration, which is included in this study (Fig. 1.16G); Banse's data are used in the above key.

Hartman (1961) proposed that Hesperalia californiensis Chamberlin, 1919, is a junior synonym of

I O. phosphorea. Re-examination of the of both species by one of us (L. Harris) revealed that H.

types

californiensis is a species of Odontosyllis, but not O. phosphorea. Odontosyllis phosphorea differs from O.

californiensis in having blades of compound setae with secondary teeth located subdistally, rather than atthe

I midpoint of the cutting edge. However, H. nans Chamberlin, 1919, has the identical pigmentation pattern and setae that are characteristic of O. phosphorea. We suggest that the latter two species are synonyms, although

the type ofH. nans has dried out, and precise comparisons are not presently feasible.

I Distribution. coast, Mexico to British Columbia.

Pacific

I OdontosyUisfragiUs Kudenov and Harris, new species

Figure 1.17

I Material Examined. California: Western Santa Barbara Channel, east of Point Conception, Sta. BRA-1 rock, holotype (USNM 170918).

Description. Holotype complete specimen 5.4 mm long, 0.3 mm wide excluding setae, with 65

I in 3 anterior with 19 middle with with 30. Color in alcohol segments parts: part setigers; part 15; posterior

generally white, except for middorsal pigment patch on each segment, including patch under occipital flap on

tentacular segment. i

m=

I Prostomium wider than long, with deep medial cleft along posterior margin; 2pairs of eyes, lenticulate, !

i

i connate, arranged trapezoidally; anterior pair largest (Fig. 1.17A). Antennae smooth; median antenna inserted

i on anterior prostomium margin, longer than prostomium, just anterior to anterior pair of eyes; lateral antennae inserted near anterior edge of prostomium, slightly shorter than median antenna (Fig. 1.17A). Palps large,

globular, directed ventrally, basally free, visible in dorsal view (Fig. 1.17A). Pharynx extending to setiger 4

when inverted; with at least 5 curved, subequal chitinous teeth (dissected), 1 middorsal, 2 dorsolateral and 2

I ventrolateral in position (Fig. 1.17E). Proventriculus in setigers 5-16, with approximately 300 rows of minute,

densely packed muscle cells (dissected) barely visible using 250x magnification.

i Tentacular segment dorsally distinct from prostomium and setiger 1, with tongue-shaped occipital flap extending over prostomium, not covering eyes (Fig. 1.17A). Two pairs of tentacular cirri, subequal,

smooth and longer than prostomial antennae; dorsal pair longest, each nearly 2 times longer than median

i antenna Parapodia (Fig. 1.17Atapering, ). with distally truncate presetal lobe extending beyond parapodial lobe in anterior

segments (Fig. 1.I7B); gradually becoming smaller, distally incised into 2 unequal lobes, superior lobe smallest,

I as1.17F, sociated G), around with acic10-15 ula inanteriorly, medial anddecreasing posterior segments to about(Fig. 7-121.17C,D,I posteriorly. ). AllBlades setae compou similarndinfalcigers size, distally (Fig.

bidentate, increasing slightly in length inferiorly within fascicles (Fig. 1.17F,G). Superior blades with distal

t tooth smaller than subdistal tooth, becoming larger in inferior blades; smooth cutting edges, although 2 additional,

I 49
I

I

'

FG I-I l E

I

Figure 1.17. OdontosyUisfragilis Kudenov and Harris, new species: A, anterior segments, dorsal view; B, left i

parapodium, setiger 11, setae omitted, anterior view; C, left parapodium, setiger 11, including |

dorsal cirrus and setae, anterior view; D, left parapodium, setiger 36, anterior view; E, pharynx,

dissected, dorsolateral view; F,superior compound falciger, sefiger 11; G, inferior compound falciger,

setiger 11; H, paired aciculae, setiger 11; I, paired aciculae embedded in acicular lobe, setiger I 1. R

(A-I, originals by JDK) i

i

small denticles sometimes present on largest inferior blades. Shafts distally inflated, with subdistal spines on

superior surfaces, these best developed on superior setae (Fig. 1.17F, G). Two aciculae per parapodium, not •

projecting tl_rough parapodial lobes or cuticle; distally pointed, tip circumscribed by flange or collar (Fig. II

1.17H,I).

Dorsal cirri smooth, cirriform, arising from body wail. Ventral cirri thickly digitiform, inflated, not i

extending beyond parapodium. U

Pygidium unknown.

Remarks. Odontosyllisfragilis is characterized by the presence of a dorsomedian row of pigment i

spots from the tentacular segment to the end of the body, 5 subequal chitinous teeth in the pharynx of which

J

one is middorsal, pharynx in setigers 1-4, proventriculus in setigers 5-16 with perhaps 300 rows of small,

tightly packed muscle cells that are poorly discemable using 250 x magnification; blades of compound falcigers H

all distally bidentate with smooth cutting edges, and two acicula per parapodium. The proventriculus is i

particularly unusual, and is not like those described for known Odontosyllis species.

Etymology. The name,fragilis, alludes to the fact that specimens are extremely delicate, and seldom i

remain intact. m

Distribution. Southern California. i

g

50 I
I L

I Genus Opisthodonta Langerhans, 1879

i Type species: Opisthodonta morena Langerhans, 1879

Diagnosis. Prostomium with 3 antennae. Palps fused basally. Nuchal organs large, ciliated ridges

between prostomium and tentacular segment. Two pairs of tentacular cirri. Antennae, tentacular cirri and

I dorsal cirri smooth, slender. Pharynx armed with a large middorsal posterior tooth. Some anterior parapodia with thick aciculae.

i Remarks. Hartmann-Schrrder (1971) reviewed the genus Opisthodonta which includes the type species O. morena (Langerhans, 1879) and O. pterochaeta (Southern, 1914). Opisthodonta is polytypic as

presently defined: O. pterochaeta is more closely allied to the Streptosyllis-complex in having huge aciculae

i terminating in flat, enlarged ends plus thick compound falcigers with short blades confined to anterior segments,

and will be assigned to a new genus elsewhere (Dorsey and Kudenov, in preparation); it will not be included

in this analysis.

I Opisthodonta mitehelli Kudenov and Harris, new species

I Figure 1.18

Material Examined. California: Santa Mafia Basin, off Point Arguello, Sta. BRA-6, rock, holotype

i (USNM BRC-13, 170919) + 2 paratypes (LACM-AHF Poly 1674); Stas. BRA-13, rock, paratype (AM W. 22191); rock (1); BRC-14, rock (1); BRA-16 rock, paratype (USNM 170920); BRA-21 rock (1); between

Point San Luis and Point Sal, Sta. BRA-25 (2); off E1 Morro, Sta. BRA-27 (1); off Point Estero, Sta. 1, 2

i paratypes Arguello, Sta. (LACM-AHF PH-N Poly 1669); off E1Morro, Sta. 6, 6 paratypes (LACM-AHF Poty 1670); off Point (1).--Western Santa Barbara Channel, east of Point Conception, Stas. BRA-1 rock (1);

BRC-1 rock, paratype (SBMNH 142659); south of Point Conception, Stas. BRA-2, rock (1); BRC-2, rock

(1).

I Description. Holotype 9.5 mm long, 0.5 mm wide without parapodia at setiger 7, with 60 setigers.

Body coiled, strongly arched dorsally, flattened ventrally; fragile. Body wall pigmentation absent inpreserved

I specimens; P pharynx darkreddish brown. rostomium subrectangular to pentagonal; 2pairs of eyes, anterior pair largest, lenticulate, widely

separated, in trapezoidal arrangement; additional pair generally present medial to and near bases of lateral

I antennae, from posterior minute, margin lenticulate, of prostomium, absent in hjust olotype anterior (Fig.to1.18A). nuchal Antennae organs, extending long, smooposteriorly th; mediantoanten setiger na arising 10-11,

2.5-3 times longer than lateral antennae; lateral antennae arising from anterior margin of prostomium, anterior

I tconspicuously o largest pair ciliated, of eyes; present each 2.5ontime posterior s l nger prostomial than prostomi margnum(Fig. (Fig.1.18A). 1.18A).PalpsNuchal square,ridgeextremely continuowell us,

developed, fused basally; shorter, wider than prostomium (Fig. 1.18A). Pharynx wide, thick-walled, with

I middorsal to setigers tooth t0-11inwith postedistal rior half; circlet exteofnding 10 sotoftsetiger papillae12(Fig. when1.18B). retractedProventri (Fig. 1.1c8ulus A); wlong, hen everted, barrel-shaped, extendingin

setigers 13-20 (extending to setigers 16-20), with approximately 23-27 rows of muscle cells.

I Dorsal Tentacular segment dorsally distinct from prostomium, not as wide asfollowing segments (Fig. 1.18A). tentacular cirri three-fourths length of median antenna; ventral tentacular cirri one-third length of

dorsal cirri (Fig. 1.18A).

I Parapodia with single anteriorly placed node, usually located over tips of aciculae (not illustrated). Parapodial lobes of setigers 1-12 thick, fleshy, broadly truncate with pronounced ventral lip, appearing crowded

and together conveying impression of a modified anterior region (Fig. 1.18A,B); parapodia of median and

I posterior setigers longer, thinner. Setae generally compound falcigers (Fig. 1.18D,E); blades all bifid, each
I

I

A I

I? ,'

!

I

!

I

Figure 1.18. Opisthodonta mitchelli Kudenov and Harris, new species: A, Anteriorend,dorsalview;B, fight 1

parapodium,setiger4, anteriorview; C, superiorcompoundfalciger,setiger4; D,inferiorcompound II

falciger,sctiger 4; E,shafttip of compoundfalciger,posteroventralview; F,shafttipof compound

falciger, ventral view; G, shafttip of compoundfalciger,dorsalview; H, Acicula. (A-G, redrawn I

from originalsby J. Dorsey) I

with fine, long serrations on cutting edges, suggesting presence of sheath at low magnification. Setigers 1-12 1

with 3-5 superior falcigers with blades (Fig. 1.18C) 2 times longer than middle and inferior blades (Fig. l

1.18D) within fascicles, latter short and numerous, arranged in close-set dorso-ventral rows. Setae of median,

posterior setigers consisting of 1-2 superior falcigers with blades slightly longer than others within fascicles, I

all gradually decreasing inferiorly; median setigers with 7-9 compound setae per fascicle, decreasing to 4-5

posteriorly. Shaft endings all strongly serrated, distally excavate (Fig. 1.18E-F). Dorsal simple seta present

in median setigers to end of body, long, slender, tips appearing flattened, distally bifid, gently curved; cutting 1

surface serrated. Ventral inferior seta present in last 10-13 setigers, stout, distally bifid with small superior II

tooth; cutting surface serrated. Three aciculae per anterior parapodia, of these 2 slightly enlarged, all with

bent tips, decreasing to 1 per pampodium in posterior segments (Fig. 1.18H). 1

Dorsal cirri inserted on small elevations above parapodia, cirrophores conspicuous; alternating in

length along body, longest anteriorly, decreasing in overall length posteriorly (Fig. 1.18A). Cirri of setiger 1

largest, extending posteriorly over 17 segments, those of setiger 2 over 6-7 segments (Fig. 1.18A). Longer I

dorsal cirri of median and posterior setigers extending over 2-3 segments; shorter cirri over 1-2 segments. J

Ventral cirri large, foliaceous in anterior segments, extending beyond tip of parapodial lobes (Fig. 1.18B),

gradually become smaller, digitiform, not extending beyond parapodia in median and posterior segments. 1

II

52 I
!

I Pygidium semicircular, with 2 long, smooth anal cirri.

Additional details are here provided for small specimens measuring 1.8 mm long, with 22 setigers:

I pharynx to setigersextending 1-6. Atocomplete setiger 6;juvenile proventriculus measurespresent 1 mmin long, setigers12.512-19; om broadly wide excluding truncate parapodia, parapodia confined with 13

setigers. Six eyes present. Pharynx inverted, extending to setiger 4. Proventriculus in setigers 5 to anterior

I setiger faicigers7,numbering with around1025perrows fascicleof muscle in setigers cells.1-4,Truncate blades longest parapodia in superiormost in setigers 1-4 2 toenlarged. 3 setae; numbe Compound ring 6-

7 per fascicle in median parapodia; and 3 in posterior parapodia. Dorsal superior simple seta beginning in

I setigers compared9-13. to adults; Ventralanterior inferiorones simple as long setaasbeginning body width; in setigers posterior11-13. ones Dorsal 2 timescirri lengthproportionately of the body. smaller Ventral

cirri foliaceous in setigers 1-6, transitional in form in setigers 7-8, becoming digitiform in setigers 9-13.

I long spiniger-like Remarks. falcigers, Opisthodonta and instead mitchelli havingdiffers short-bladed from Opisthodonta falcigers enclosed sp. B by(Uebelacker, a sheath; continuous 1984) in lacking instead

of paired nuchal organs; aciculae tapering and slightly bent distally instead of being straight and terminating

I rather in a distthan al knob auricular and subdistal ventral cirflange ri. that is marginally dentate; and foliaceous and digitiform ventral cirri

Opisthodonta mitchelli also differs from Opisthodonta sp. A (Uebelacker, 1984) in lacking aciculae

I with distally rounded ends and characteristic subdistal clefts, in having the proventriculus longer, extending through 6-7 segments with 23-27 rows of muscle cells, instead of 2-4 segments with 19-23 rows; and in

having ventral cirri both foliaceous and digitiform rather than auricular.

I mitchelli differs from O. in aciculae in anterior Opisthodonta morena lacking enlarged segments;

superior compound blades of anterior setigers two times instead of four times longer than middle and inferior

blades within fascicles; 23-27 instead of 18-24 rows proventricular muscle cells; and simple setae in median

I and posterior

segments.

One female specimen ofO. mitchelli (Sta. PH-N) has gametes beginning in setiger 20; all others noni reproductive. Habitat. Soft bottoms and rocky substrata, in depths of72-111 m.

Etymology. This species is named in fond memory and honor of Charles "Mitch" Mitchell, St., who

I was a special Distribperson to both L. Harris and John H. Dorsey, and whose absence is deeply regretted. ution. Southern California.

I Genus Pionosyllis Malmgren, t 867

I Type species: Pionosyllis compacta Malmgren, 1867

Diagnosis. Prostomium with 3 antennae. Palps fused basally or separate. Nuchal organs usually as

I ccirri liatedandridorsal dges alcirri ongsmooth postectaltomargins weakly articulated. of prostomiumPharynx . Twowith pairsa osingle f tentacmiddorsal ular cirritooth; . Antenn rimae,smooth. tentacular

Remarks. The presence of a smooth, edentate pharyngeal margin or rim is critical to the systematics

I ogenus. f PionosyIl Specis species. This character mustbe determined before specimens can be correctly assigned to this imens with retracted pharynges must therefore be dissected to avoid confusion with Eusyllis.

Although Pionosyllis gigantea Moore, 1908, was not encountered in MMS materials examined, it is

I widely distributed both latitudinally and bathymetrically from Alaska to southern California (Hamnan, 1968),

and is included in the key below. Pionosyllis uraga Imajima, 1966, has also been found in southern California

(L. Harris, personal observation).

I
!

Key to the Species of Pionosyllis from California I

1A. Blades of compound setae at least 3 times longer than basal width (Fig. 1.120C-E) ..................... 2 I

I

IB. Blades of compound setae short,less than3 times longer than wide (Fig. 1.19C); antennaearise from

anterior prostomium margin (Fig. 1.19A) ...................................................... Pionosyllis magnifica l

2A. Superior compound falcigers with extremely long, spiniger-like bidentate blades in all segments, 6-10 m

times longer than inferior blades in fascicles (Fig. 1.20C-E); antennae distally articulated (Fig. n

m

1.20A,B); dorsal cirri irregularly alternating in length along body (Fig. 1.20A) ...............................

........................................................................................................................ Pionosyllis articulata 1

2B. Superior compound falcigers with blades of uniform length within fascicles; antennae distally smooth; I

dorsal cirri usually long, not alternating in length along body ........................... Pionosyllis gigantea

I1

Pionosyilis magnifica Moore, 1906 U

Figure 1.19 I

I

Pionosyllis magnifica Moore, 1906:223-225, pl. 10,figs. 9-11; 1908:325.--Annenkova, 1938:152.--Hartman,

1968:447-448, figs. 1-3. •

Eusyllis magnifica Pettibone, 1954:261-262.--Banse and Hobson, 1974:56, fig. ! 4f. I

I

Material Examined. California: Santa Maria Basin, between Point Estero and Point Buchon, Sta. 6 •

rock (t, USNM 170923).--Washington: Puget Sound, holotype (USNM 5533). II

Description. Length 48 mm, 2 mm wide excluding dorsal cirri and setae, 150 segments (Moore, 1

1906; Hartman, 1968); MMS specimen 2.5 mm long, 0.3 mm wide for 17 setigers. Body widest in middle n

body segments, flattened, tapering markedly both anteriorly and posteriorly. Pigmentation absent in preserved

specimens. I1

Prostomium small, depressed, wider than long, subrectangular; 2 pairs of black eyes, anterior pair I

largest, arranged trapezoidally (Fig 1.19A). Antennae slender, long, arising near anterior prostomium margin;

median antenna 3-3.5 times longer than prostomium (lost in MMS specimen); paired lateral antennae each 2- •

3 times longer than prostomium (Fig. 1.19A). Palps large, projecting anteroventrally, basally fused (Fig. I

1.19A). Everted pharynx with subdistal, middorsal tooth plus distal circlet of 10 soft papillae. Proventriculus

of MMS specimen present in 4.5 segments. I

Tentacular segment distinct, reduced dorsally (Fig. 1.19A), well-developed ventrally as lips. Two I

pairs of tentacular cirri, long, slender, smooth; dorsal pair 4 times and ventral pair 2.5 times longer than

prostomium (Fig. 1.19A). •

Pampodia stout, fleshy, distally rounded (Fig. 1.19B), becoming more elongate, conspicuous posteriorly.

Setae compound falcigers with bidentate blades (Fig. 1.19C), about 40 per fascicle in median segments where

best developed. Blade lengths decreasing inferiorly within fascicles; all with serrated cutting edges. Notoaciculae n

slender, curved, acutely pointed, 1 per parapodium in median segments, ventral to dorsal cirriphores (Fig. U

1.19B); neuroaciculae stout, distally knobbed, 3-4 per pampodium.

Dorsal cirri cirriform, smooth to wrinkled (not annulated), resembling one another throughout; longest I

in anterior segments, decreasing to about one-half body width in median segments; all with long, pronounced

dorsal cirrophores. Ventral cirri short, stout and enlarged, distally blunt, not extending beyond parapodial.

I

Pygidium small, with paired anal cirri, each the length of the last 12 segments. I

54 I
I

!

|

!

I Figure 1.19. Pionosyllis redrawn frommagnifica Moore,:1906) A, anterior end, dorsal view; B, parapodium; C, compound falciger. (A-C,

!

Remarks. Pionosyllis magnifica is a long established species normally associated with muddy,

I sandy shelly Alaska, British Columbia and Washington (Pettibone,

and sediments in shallow habitats of

1954; Hartman, 1968; Banse and Hobson, 1974). The single MMS specimen is poorly preserved. Pettibone

(1954) referred Moore's species to Eusyllis on the basis of its denticulate pharyngeal margin. Examination of

I Moore's by of (L. Harris) confirmed the of smooth and

type one US presence a pharyngeal margin, supports

the original placement of this species in Pionosyllis.

i HDistr abitat ibutio . Mun. d,Csaanlifodranniad; shelly Washibot ngtontom;s,Altaska. o a depth of 109 m.

I PionosyUis articulata Kudenov and Harris, new species

Figure 1.20

I Material Examined. California: Santa Mafia Basin, off Point Estero, Sta. 2, 3 paratypes (SBMNH

142660) + (4); off Purisima Point, Sta. 42, 1 paratype (USNM 170922) + (9); off Point Buchon, Sta. 12 (6);

I off Arguello, Point San Sta. L64uis,(2);Sta. off 22Point (1);Buchon, off PointSta. Sal,BRC-26 Sta. 30Extra, , 4 paratypes rock, holotype (AM W(USNM . 22192);170921); Sta. 43 (1off);E1offMorro, Point

Sta. BRA-28 Extra, rock, 6 paratypes (LACM-AHF Poly 1671); off Point Sal, Stas. PJ-1 (3); PJ-2 (4); PJI Western 5 (1); PJSanta 6 (5);Barbara PJ-7, 13Channel, paratypesoff(LACM-AHF Point Conception, Poly Sta. I672)BRA-2 + (24);(1).--Point PJ-8 (2); Loma, PJ-10 (22); head PJ-16 of Coronado (1).-

Canyon, Army Corp of Engineers, Stas. BD5-2 (3).

I parapodiaDesc(atrisetiger ption. 7);Holotype 68 setigers. measuBody res 6 mmcoiled, long,t_eadlike; 0.3 mm wide setigers excluding 1-9 eachparapodi shorta,,u0.5 suallymmcont wideractewithd,

parapodia closely spaced, setigers becoming longer from setiger 10(Fig. 1.20A); dorsum with paired transverse

I bands pharynxof cilia on each segment, conspicuous in reproductive individuals. Pigment absent in preserved specimens; reddish-brown. Sexually mature specimens with swimming setae beginning on setiger 21, continuing

to near end of body; with 2-4 eggs per segment in setigers to far posterior.

!

| "
!

Figure 1.20. Pionosyllis articulam Kudenov and Harris, new species: A, anteriorend, dorsalview; B,pharynx

andsetiger I, ventralview; C,superior compoundfalciger,setiger6; D, inferiorcompoundfalciger,

setiger 6; E, superiorcompound falciger,with detailof tip,setiger 40;F,inferiorcompound falciger,

setiger 40; G, dorsal simple seta, setiger 40; H, ventral simple seta, setiger 40. (A-H, redrawn from

originals by J. Dorsey)

!

Prostomium wider than long (Fig. 1.20A); 2 pairs of eyes, in trapezoidal arrangement, anterior pair

largest, lenticutate; additional pair of eyespots sometimes present, usually on anterior prostomium margin, I

anterior and medial to lateral antennae (Fig. 1.20A). Antennae slender, basally smooth (Fig. 1.20A); median

antenna long, about 5 times width of prostomium, with about 16 distal articles; arising just posterior to

midpoint of prostomium; lateral antennae about one-half length of median antenna, each with 6distal articles, I

arising near anterior border of prostomium. Paips large, flattened, rounded anteriorly, slightly fused basally III

(Fig. 1.20A); distal one-third to one-half usually projecting ventrally. Nuchal organs not observed. Pharynx

extending to setigers 6-7, with anterior middorsal tooth; chitinized margin smooth, surrounded by circlet of 10 B

soft distal papillae, with small papiUae-tike thickenings on proximal circumoral ring (Fig. 1.20B). Proventficulus II

extending from setigers 7-8 to 10-12, usually 5 segments in length, with 22-24 rows of muscle cells (Fig.

1.20A). I

Tentacular segment as long as setiger 1, usually contracted, with 2 pairs of tentacular cirri, similar in

form to prostomial antennae, distal articles sometimes present; dorsal pair longest, each about same length as

lateral antennae; ventral pair each about one-third to one-half length of dorsal pair (Fig. 1.20A,B). I
!

!

Parapodia conical, distally rounded, with small round pre-, and post-setal lobes. Compound setae

I bidentate falcigers. Blades decreasing in length posteriorly along body and inferiorly within fascicles; blades in setigers 1-19 thin, becoming more stout from setiger 20 to end of body. Distal denticle of short blades in

setigers 1-19 somewhat larger than subdistal denticle (Fig. 1.20E); from setiger 20 becoming smaller than

I subdistal becomingdenticle, coarser, sharply bidentate (Fig. 1.2OF). Cutting edges finely serrated in blades of anterior fascicles, less numerous in blades of posterior fascicles. Superiormost falcigers with extremely long

blades in all setigers, numbering 4-5 in anterior segments, decreasing to 1 posteriorly (Figs. C,D); blades of

I C,E), superior increasing 4-5 compound after setiger falcigers 20 upwithin to 10fascicles times length of setigers (Figs. 1-19 D,F).sixDorsal times length simple ofsetathose frominferiormost setigers 21-22 (Figs. to

end of body, initially unidentate, becoming bidentate posteriorly, with smooth cutting surfaces (Fig. 1.20G).

I AVentral ciculaesimple numbering seta from 2 per aboutnteriorsetiger parapodia, 36 to increasing end of body, to 3-4bifid, inposterior with smooth segments; cuttinganterior surfacesones(Fig. terminating 1.20H).

in diamond-shaped tips, generally becoming distally blunt posteriorly except for one enlarged geniculate

I form. Dorsal cirri present on all segments, smooth except for those of setiger 1, which are slightly articulate

distally (Fig. 1.20A); irregularly alternating in length all along body (short-long, two short-one long). Those

I ofwhere setigerlongest 1 1.3-1.5 up totimes 2.5 times longerthethanwidth dorsaloftentacular median segment cirri; alls, generally shortest one-half increasingthisin length width (both to middle estimbaotesdy

excluding parapodia). Ventral cirri of anterior segments digitiform (Fig. 1.20B), not extending beyond

I parapodial lobes, decreasing to about one-half the length of median, posterior parapodia. Pygidium with 1 pair of anal cirri, each long, smooth; median digitiform cirrus short.

Remarks. Pionosyllis articulata is characterized by the presence of superior compound falcigers

I extremely long spiniger-like setigers; change shape offalcigers setiger 19; prostomial

with blades in all in after

antennae distally articulated; dorsal cirri that alternate long and short; acicula increasing in numbers in posterior

parapodia; presence of simple superior and inferior setae; and setigers 1-9 apparently shorter than those

I following. Although the small paired usually the anterior prostomium, they apparently eyespots are present on

can occur nearly anywhere on the prostomium or be absent. For example, they may either be located next to

the median antenna, or along the posterior prostomium, in addition to the usual location noted above.

I Superior compound falcigers with extremely long blades are present in fascicles of P. a_'culata, P.

ehlersiaeformis Augener, 1913, P.weismannioides Augener, 1913, P. augeneri Hartmann-Schrtder, 1979, P.

ll longisetosa Hartmann-Sclaroder, 1965, and P. uraga Imajima, 1966c. Pionosyllis articulata differs from

II these species in having antennae distinctly articulated distally rather than smooth or indistinctly wrinkled.

i

, " Pionosyllis articulata, P. longisetosa andP. uraga differ from species listed above in having compound setae

with long blades in all setigers; these setae are first present in median setigers of P. ehlersiaeformis, P.

I P. Both P. articuIata and P differ from P. in that the

weismannioides, augeneri. longisetosa uraga long

blades of compound setae are falcigerous and distally bidentate rather than spinigerous and distally tapered to

fine points. Pionosyllis articulata differs fromP. Iongisetosa in having long, thin, whip-like blades each 6-10

I times longer than inferior setae in the same fascicles, rather than wide blades each around 3 times longer than

inferior setae; spiniger-like blades with freely serrated cutting edges in conWast to ones with coarsely serrated

cutting edges; blades with subdistal tooth well clef'reedinstead of being weakly defmed; falcigers becoming

I stout in medial segments rather that being of uniform throughout; simple setae distally bidentate with serrated cutting surfaces instead of distally rounded and unidentate with smooth cutting surfaces; and having dorsal

cirri alternating irregularly in length, rather than regularly. Pionosyllis articulata further differs from P.

I uraga in having blades of compound falcigers from median segments short, sharply bidentate, rather than butter knife-like and weakly bidentate; and having digitiform ventral cirri instead of foliaceous ones that are

broadly attached to parapodial lobes.

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The tips of long spiniger-like blades from compound falcigers must be examined using oil immersion.

It may be possible to subdividePionosyllis into a subgenus of species all having long spiniger-like compound •

setae similar to the way Ehlersia has functionally been separated from Typosyllis (Uschakov, 1955; Day, 1

1967; Banse and Hobson, 1974; Uebelacker, 1984; Dorsey andPhillips, 1987). Although such an investigation

falls well outside the scope of the present study, it is likely to be both interesting and productive. •

Habitat. Soft bottoms and rocky substrata, in depths of 60-205 m.

Etymology. The species name, articulata, refers to the characteristic distal articulations of the Ilb

prostomial antennae, and dorsal cirri of setiger 1, 1

u

Distribution. Southern to central California.

Genus Syllides Orsted, 1845 I

llll

Type species: Syllides longocirrata Orsted, 1845 I

I

Diagnosis. Prostomium with 3 antennae. Tentacular segment frequently collar like and ciliated. •HI

Two pairs of tentacular cirri. Some dorsal cirri usually articulated. Setae and aciculae of anterior segments

not enlarged. Pharynx usually unarmed, with smooth anterior marginal rim. 1

Remarks. Banse (1971) reviewed the genus, and provided descriptions, illustrations and a key. The I

key below includes both Syllidesjaponica Imajima, 1966c, and S. longocirrata, which have been reported

from Washington and British Columbia (Banse and Hobson, 1974). I

Key to Known Species of SyUides from the Northeast Pacific I

1A. Two kinds of dorsal simple setae present: thick with bent tips in setigers 1-5;thin and distally tapering []

from setiger 6 to end of body ........................................................................... Syllides longocirrata II

lB. One kind of dorsal simple seta present in all segments (Figs. 1.21D, 1.22R,S) ............................. 2

I

2A. Dorsal cirri with bright, reflective golden yellow pigment in most articles; 20 or fewer articles per

dorsal cirrus (Figs. 1.21C, 1.22D-G) .............................................................................................. 3 I

2B. Dorsal cirri lack reflective pigment; 20-30 articles per dorsal cirrus .................... Syllidesjaponica 1

3A. Dorsal cirri with 15-20 spherical articles (Fig. 1.21A,C); median antenna shorter than lateral antennae 1

(Fig. 1.21A); integument with epidermal papillae (Fig. 1.21A); compound and dorsal simple setae

lack distal hoods; dorsal simple setae capillary tipped (Fig. 1.21D) ............................ Syllides reishi

I

3B. Dorsal cirri with 5-8 tear drop-shaped articles (Fig. 1.22D-G); median antenna longer than lateral 1

antennae (Fig. 1.22A); integument lacking papillae; compound and dorsal simple setae enshrouded

n

by distal hoods (Fig. 1.22R-T); dorsal simple setae distally blunt (Fig. 1.22R-T) .... Syllides mikeIi

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I Syllides reishi Dorsey, 1978

I Figure 1.21

Syllides reishi Dorsey, 1978:24-26, figs. ld, 2 a-f.

I Material Examined. California: Santa Maria Basin, Sta. BRA-13 rock (1, USNM 170934); off E1

Morro, Sta. 6 rocks (1).---San Clemente Island, holotype (LACM-AHF Poly 1255) and paratypes (9, LACMI AHF Poly 1256).

Description. Length 2 mm, 0.4 mm wide excluding parapodia, for 30 setigers (Dorsey, 1978). Body

golden brown in life, epidermal papillae dark brown. Epidermal papillae small, irregularly shaped, multilobate,

I abundant on dorsum, becoming numerous on ventrum, infrequent on prostomium, palps, parapodia; absent

from intersegmental regions. Articles of dorsal cirri contain golden yellow pigment (Fig. 1.21C).

Prostomium wider than long with pharynx inverted (Fig. 1.21A), pentagonal when everted (Fig.

I 1.21B); 2 pairs of eyes, lenticulate, subequal in size, anterior pair set farthest apart, all arranged trapezoidally;

additional pair of eyespots present on anterior prostomium margin (Fig. 1.21A,B). Antennae smooth to

wrinkled; median antenna arising between posterior pair of eyes, up to 1.5 times longer than prostomium;

I antennae arising front of and medial to anterior pair of eyes, each 3 times longer than prostomium,

lateral in

two times longer than median antenna (Fig. 1.21A,B). Pharynx extending to setiger 3, unarmed; lacking

distal circlet of soft terminal papillae when evened. Proventriculus extending through setigers 4-7 when

I pharynx inverted.

Tentacular segment distinct dorsally (Fig. 1.21A). Tentacular cirri cirriform, wrinkled, 2 pairs;

dorsal pair 1.2 times longer than lateral antennae, nearly 3 times longer than prostomium; ventral pair about

I as long as dorsal pair.

Parapodia short, conical (Fig. 1.21C). Compound setae bidentate falcigers (Fig. 1.21D,E), 8-9 in

I anterior inferior blades fscicles, withindecreasi fascicles, ng tgradually o 2-5 in posterior decreasingfascicle posteriorly s. Blades alonglongest body; allsuwith periorly, minutely 2 times serrated longercuttinghan

edges (Fig. 1.21D). Dorsal simple seta with capillary tip, present from setiger 1, with serrated cutting surface

I (Fig. 1.21F). DorsalAcic cirriulaeof with distally enlarged tips, 1 per pampodium (Fig. 1.21G). setiger 1inflated, wrinkled; strongly articulated from setiger 2 to end of body, with 15-

20 articles anteriorly, decreasing to about 10 articles posteriorly. Articles each with 1-2 vesicles, variably

I present, parapodialcontaining lobes, alsogocontaining lden-yellowgolden-yellow granules (Fig.granules. 1.21C). Ventral cirri digitiform, as long as or longer than

Pygidium with paired anal cirri plus median unpaired ventral cirrus; all spherical, equal in size.

I Remarks. reishi is well defined described from San Clemente

Syllides a

species originally Island,

that seems to be present only in rocky habitats along southern California. Refer to the discussion ofS. mikeli

below for additional comments concerning S. reishi.

I Distribution. Southern and central California.

I

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G

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Figure 1.21. Sytlides reishi: A, anterior end, dorsal view; B, prostomium and everted pharynx, dorsal view; C, I

parapodium 16, posterior view; D, dorsal simple seta, parapodium 10; E, compound falciger,

parapodium 10; F, shaft end of compound falciger, ventral view; G, acicula, parapodium 10. (A-G,

redrawn from J. Dorsey, 1978) I

I

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Syllides mikeli Kudenov and Harris, new species

Figure 1.22

l Material Examined. California:SantaMaria Basin, off Point Sal, Stas. PJ-1, I paratype (SBMNH

142662) + (1); PJ-2 (1); PJ-3 (1); PJ-7, holotype (USNM 170931), 1paratype (USNM 170932), 4 paratypes

,I ((2); LACM-AHF PJ-16, 3 paratypes Poly 1676); (USPNMJ-8, 170933). 1 paratype (LACM-AHF Poly 1677); PJ-10, 1 paratype (AM W. 22195)+

Description. Holotype complete, 1.4mm long, 0.2 mm wide excluding parapodia, 0.3 mm wide with

I parapodia, for 26 setigers. Body excluding dorsal cirri white in preserved specimens; articles of dorsal cirri with golden-brown pigment granules in transmitted light, yellow-brown in reflected light. Epidermal papillae

absent. Each segment with a dorsal transverse row of cilia.

Prostomium wider than long with pharynx inverted; pentagonal when everted; 3 pairs of eyes,

lenticulate, middle pair largest and set farthest apart, posterior 2 pairs arranged trapezoidally; anterior pair

associated with lateral antennae (Fig. 1.22A,B). Antennae smooth to wrinkled; median antenna arising between

I posterior pair of to 2 times longer than prostomium; lateral antennae arising in front of and medial to

eyes, up

anteriormost pair of lenticulate eyes, each 1.5 times longer than prostomiurn, two-thirds length of median

antenna (Fig. 1.22A). Pharynx extending to junction of setigers 3-4 or through setiger 4, unarmed (Fig.

I 1.22A); subdistal circlet of I0 soft terminal papillae (Fig. 1.22C), uni- to trilobed when pharynx everted, shape apparently associated with extent of pharyngeal eversion. Proventriculus extending through setigers 4

to 7-8 when pharynx inverted (Fig. 1.22A), from junction of setigers 1-2 through setiger 6 when pharynx

I everted, with about 46 rows of small muscle cells.

Tentacular segment distinct dorsally, glandular, with anterior margin conspicuously ciliated (Fig.

1.22A,B). Tentacular cirri cirriform, wrinkled to smooth, 2 pairs; dorsal pair about as long as lateral antennae;

I ventral about one-half of dorsal cirri 1

pair length (Fig. ,22A_B),

Parapodia narrow, conical (Fig. 1.22D-G). Compound setae bidentate falcigers with minutely sensed

i cutting edges and hoods covering both cutting edges and distal tips (Fig. 1.22H-Q); hoods well-developed in setae of anterior fascicles. Setae number 10 per fascicle each in setigers 1-2, increasing to 12-14 each in

setigers 5-6, decreasing to 8-9 to end of body. Superior blades of anterior setigers narrow, longest, distally

i twisted (Fig. 1.22H,I,K), 2 times longer than inferior blades within anterior fascicles (Fig. 1.22J,L), gradually decreasing posteriorly along body (Fig. 1.22P,Q); 1-2 per fascicle in setiger 1; 2-3 in setiger 2; 3 in setigers 5-

6; 2 from setiger 7 to end of body. Dorsal superior simple seta slender, capillary, present from setiger 1 with

1 minute denficles of long serrated cutting surface covered by long hood (Fig. 1.22R,S), terminating in bubblel shaped capsule around unidentate tip posteriorly (Fig. 1.22T). Ventral inferior simple seta long, capillary,

lacking distal hood, present in last few posterior parapodia (Fig. 1.22U). Aciculae distally blunt anteriorly

(Fig. 1.22V), with distal knobs posteriorly (Fig. 1.22W), 1 per parapodium.

I Dorsal cirri of setigers 1-2 cirriform (Fig. 1.22A), wrinkled to smooth, strongly articulated thereafter

to end of body, articles teardrop-shaped, increasing in length to setigers 3-6 (Fig. 1.22I)), decreasing gradually

I thereafter to end of body. Dorsal articles of setiger 3 (= 5); setiger 4 (= 5-6); setiger 5 (= 5-8); setiger 7 (= 4- 6); setiger 9 (= 5-6); setiger 19 (= 4-5); each with 1-2 vesicles containing reflective golden-yellow pigment.

Ventral cirri digitiform, extending almost to but not beyond end of pampodial lobes (Fig. 1.22D-G).

• Pygidium with paired anal cirri plus median unpaired ventral cirrus.

I il Remarks. SylIides mikeli, S. reishi, S.japonica Imajima, 1966c, and S. longocirrata Orsted, 1845

sensu Banse (1971) are reported from the western coast of North America. Syllides mikeli is superficially

similar mikeli differs toS. reishi in that both have golden-yellow pigment granules in the articles of the dorsal cirri. Syllides from S. reishi in having a long median and short lateral antennae rather than a short median

antenna and longer laterals; lacking epidermal papillae; having both compound falcigers and dorsal simple

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Figure 1.22. Syllides mikeli Kudenov and Harris, new species: A, anterior end, dorsal view; B, prostomium •

and pharynx everted, dorsal view; C, pharynx, dorsal view; D, left parapodium, setiger 5, dorsal

view; E, left parapodium, setiger 7, anterior view; E right parapodium, setiger 9, anterior view; G, l

right parapodium, setiger 19, anterior view; H-J, compound falcigers from setiger 1; H-I, superior I

setae; J, inferior seta; K-M, compound falcigers from setiger 3: K, superior seta; L, intermediate

seta; M, inferior seta; N-O, inferior compound falcigers, setiger 6; P-Q, compound falciger from

setiger 20; P, superior seta; Q, inferior seta; R-T, dorsal simple setae: R, setiger 1; S, setiger 16; T, 1

setiger 20 with distal hood, dorsal view; U, ventral simple seta, setiger, far posterior segment; V- I

W, aciculae: V, setiger 2; W, setiger 19. (A-W, originals by JDK)

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62 I
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setae with distal hoods; and in having dorsal cirri with tear drop-shaped rather than spherical articles, and

i these numbering 5-8 per dorsal cirrus instead of 15-20. Syllides mikeli is characteristically associated with soft bottom habitats, in contrast to S. reishi which is associated with rocky substrata. Syllides mikeli differs

from S.japonica in lacking 3-4 coarse basal serratia on the blades of superior compound falcigers; in having

I 5-8 tear drop-shaped instead of 20-30 spherical articles per dorsal cirrus; and in having digitiform rather than

l ellipsoidal ventral cirri. Syllides mikeli differs from S. longocirrata as described by Banse (1971) in having

a single kind of dorsal superior simple seta instead of two kinds including a thick, stout and distally blunt form

in setigers 1-5; in having bidentate rather than unidentate compound setae; and in lacking basal spurs on the

I superior compound falcigers.

blades of

We suspect the five specimens described by Banse (1971) as S.japonica represent an undescribed

i species. For example, the blades of superior compound falcigers of Banse's specimens are five times the length of inferior blades within fascicles rather than two times as reported by Imajima (1966c); the number of

setae per fascicte number 10-20 rather than 5-12; the proventriculus extends through 6 segments and not 4-5;

and that number of articles in dorsal cirri number 10-20 instead of 20-30. Banse's materials need to be reI examined with these differences in mind.

Etymology. This species is named for Tim Mikel, ABC Laboratories, Ventura, California, in

appreciation Distriofbuhis ton.friendship CentraltoCalif the ofirst rnia.author.

Subfamily Syllinae

i Genus Ehlersia Quatrefages, 1865

Type species: Syllis sexocuIata Ehlers, 1864

I Diagnosis. Prostomium with 3 antennae. Two pairs of tentacular cirri. Antennae, tentacular cirri

and dorsal cirri of anterior segments articulated. Pharynxwith subdistal middorsal tooth andsmooth anterior

I marginal longer thanrim.inferior Compoblades und setae withinfalcigero fasciclesus,ofwmedian ith superior and posterior blades spiniger-like parapodia. inTwo appearan simplece,setae at least per median 4 times

and posterior setal fascicles.

I minutelyRemarks. Compound setae with extremely long blades invariably have blunt unidentate, knobbed or bidentate tips, are present in the superiormost region of fascicles, and are most appropriately described

as compound falcigers. Previous reports of compound spinigers tend to refer to these setae (Uebelacker,

1984; Dorsey and Phillips, 1987). Dorsey and Phillips (1987 )note that Ehlersia and Typosyllis differ primarily in the respective presence

or absence of falcigers with long blades, and this trait has been used by other workers (Imajima, 1966e;

I Gardiner, 1976; Ben-Eliahu, 1977; Fauchald, 1977). Banse and Hobson (1974) first used blade ratios (4.0) within fascicles to separate species of these genera, as do Dorsey and Phillips (1987); Uebelacker (1984) uses

a higher ratio (5.0). Preliminary morphometfic data provided by Dorsey and Phillips (1987:155, 159) seem

I tmore o supprigorous ort the use multivariate of setal rati analysis os, althofousetal gh itvariation is not poss in both ible tofo these assesstaxa theirmaydatasubstantiate with regardthetovalidity body siofze.thisA

approach (see Vogt and Kudenov, 1994).

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Key to the MMS Species of Ehlersia from

the Northeast Pacific I

1A. Prostomium with 3 pairs of eyes (Fig. 1.23A); blades of superior compound falcigers from midbody I

segments 4 times longer than inferior blades within fascicles (Fig. 1.23G,E); proventriculus in 6-9 ll

segments with 30-36 rows of muscle cells ..................................................... Ehlersia heterochaeta

lB. Prostomium without eyes (Fig. 1.24A,C); blades of superior compound falcigers from midbody !

segments 10 times longer than inferior blades within fascicles (Fig. 1.24G,H); proventriculus in 4-7

u

segments with 40-43 rows of muscle cells .......................................................... Ehlersia hyperioni

!

Ehlersia heteroclmem (Moore, 1909)

Figure 1.23 I

Syllis (Ehlersia) heterochaeta Moore, 1909:322-325, pl. 15, figs. l-5.--Treadwell, 1914:176.--Rioja, I

1941:694-695.--Berkeley and Berkeley, 1948:76, fig. 113.--Banse and Hobson, 1974:62, fig. 16 k1.--Dorsey and Phillips, 1987:153-156, fig. 1 a-h.

Syllis heterochaeta Moore, 1923:256. I[

Ehlersia heterochaeta: Hartman and Barnard, 1958:21, 55.

Langerhansia heterochaeta: Hartman and Barnard, 1960:90, 276.--Hartman, 1963:17; 1968:435-434, figs.

1-7. I

Syllis (Ehlersia) cornuta: Pettibone, 1954:253, fig. 28f (in part). Not Rathke, 1843.

Syllis cornuta: Pettibone, 1963:118, figs. 3 lid (in part). Not Rathke, 1843.

!

Material E_nmed. California: Santa Maria Basin, Sta. BRA-16 lock (1, USNM 170935); off

Point Buchon, Sta. 12 (1, SBMNH 142702); off Point Sal, Sta. 30 (2); off Purisima Point, Sta. 42 (1; USNM I

98785); Santa Maria Basin, Sta. 52 (1); off Point Arguello, Stas. 64 (1); Sta. 72 (1; USNM 98784); offPoint I

Sal, Stas. R-8 (4, USNM 170936); PJ-1 (1); PJ-2 (2); PJ-3 (2); PJ-6 (7); PJ-7 (5); PJ-8 (5); PJ-10 (2); J-11

(2).--San Nicolas Island, holotype (USNM 17400). []

Description of Atoke. Length about 30 nun, 2 mm wide, for 80 segments (Moore, 1909; Dorsey and 1

Phillips, 1987); MMS specimens 12 mm long, 0.6 mm wide without setae, for 110 segments. Large black

pigment patch on posterior prostomium sometimes absent.

Prostomium pentagonal, wider than long; 3 pairs of eyes, middle pair largest, posterior pair lenticulate, •

all in trapezoidal arrangement; additional pair of eyes present as pigment specks on anterior margin (Fig.

1.23B,C). Antennae strongly articulated; median antenna long, with 13-26 articles, arising between 2 posterior 1

pairs of eyes; lateral antennae arising from anterior prostomium margin, in front of, slightly medial to largest

pair of eyes, each with 10-17 articles (Fig. 1.23B,C). Palps large, triangular, distally round, not fused

basally. Everted pharynx with anterior unpaired tooth, surrounded by distal circlet of 10 soft papillae when II

everted. Proventriculus present in 6-9 setigers, with 30-36 rows of muscle ceils. []

Tentacular segment dorsally distinct from prostomium and setiger 1; 2 pairs of tentacular cirri with 

numerous articles, resembling antennae; dorsal pair each with 17-21 articles; ventral pair each with 9-12 1 

articles (Fig. 1.23B,C). !1

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I Figure 1.23. Ehlersia heterochaeta: A, anterior end, male epitoke, dorsal view; B, anterior end, atoke, dorsal

view; C, anterior end, atoke, dorsal view; D, inferior compound falciger, anterior segment; E,

inferior compound falciger, posterior segment; F, superior compound falciger, anterior segment,

I with detail of tip; G, superior compound falciger, posterior setiger; H, ventral simple seta, posterior setiger. (A-H, after Dorsey and Phillips, 1987).

I

I
I

Parapodia distally rounded. Compound setae bidentate falcigers (Fig. 1.23D-G), with 10-16 to 21- I

28 setae per anterior fascicle, decreasing to 6-7 per posterior fascicle. Blades decreasing in length inferiorly

within fascicles. Superiormost falcigers with longest blades 3-4 times length of inferiormost blades within I

fascicles, numbering 3-4 per anterior fascicle decreasing to 2 per posterior fascicle, each with subterminal

tooth round, poorly defined and finely denticulate cutting edges (Fig. 1.23EG); middle, inferior blades short,

sharply bidentate, with cutting edges coarsely serrated (Fig. 1.23D,E). Dorsal simple seta distally unidentate, i

present in median segments to end of body. Ventral simple seta distally bidentate, present in last few posterior I

segments (Fig. 1.23H). Five to 7 aciculae per anterior parapodium, decreasing to 1-2 per posterior parapodium.

Description of Epitoke. Male epitoke 8.7 mm long, 0.8 mm wide including parapodia (Dorsey and i

Phillips, 1987). Prostomium oval; 1pair of eyes, lenticulate; antennae smooth with median antenna arising II

anterior to eyes, about two times longer than lateral antennae, which arise anterolaterally from prostomial

border (Fig. 1.23A). ParapodiaI lobes conical. Compound falcigers distally bidentate, resembling those i

described for atokous form; 4-7 per fascicle including 1-2 superior setae with long blades 3-4 times length of I

inferior ones, latter with short blades strongly bidentate. Cutting edges of superior blades freely serrated;

inferior blades coarsely serrated. One acicula per parapodium, thick, distally pointed, yellow. Long capillary i

swimming setae from setiger 3. Dorsal cirri articulated, long cirri with up to 11articles alternating with short

cirri with up to 8 articles. Ventral cirri digitiform, not extending to ends of parapodial lobes. Pygidium with

paired anal cirri, each with 21 articles; ventromedian cirrus digitiform. •

Remarks. Ehlersia heterochaeta is widely distributed along the Pacific coast of North America

where it commonly inhabits mixed sediments. Dorsey and Phillips (1987:153) state that the holotype of E.

heterochaeta is missing and established a neotype (ANSP 3328). In actuality, the holotype is at the National i

Museum of Natural History, Smithsonian Institution (USNM 17400; Loi, 1980). Ehlersia heterochaeta is u

also discussed below in the remarks for E. hyperioni (Dorsey and Phillips).

Habitat. Mixed sediments in shelf and canyon depths, 15-409 m. i

Illl

Distribution. Western Mexico to Western Canada.

Ehlersia hyperioni (Dorsey and Phillips, 1987) I

Figure 1.24 l

Syllis (Ehlersia) hyperioni Dorsey and Phillips, 1987:156-159, fig. 2 a-c, fig. 3 a-k. i

II

Material Examined. California: western Santa Barbara Channel, Sta. 46 (I, USNM 000000); east

of Point Conception, Sta. 94 (I, SBMNH 142904).---Santa Monica Bay, holotype (LACM-AHF Poly 1436); i

3 paratypes (LACM-AHF Poly 1437-1439). g

Description. Length up to 21.6 mm, 0.4 mm wide including parapodia at level of proventriculus, for

122 setigers (Dorsey and Phillips, 1987); MMS specimens anterior fragments, up to 3 mm long, 0.3 mm wide 1

excluding parapodia and setae, for up to 22 setigers. Body long, threadlike; tan in color, occasionally with

specks of dark pigment ha dorsal cirri of anterior and median segments, parapodia, ventral cirri; pharynx

orange-brown, i

Prostomium wider than long, anterior margin with palps indistinct; eyes absent (Fig. 1.24A-C).

Antennae with short ceratophores; median antenna arising from middle of prostomium, with up to 26 articles; il

lateral antennae two-thirds length of median, with 11-13 articles (Fig. 1.24A-C). Palps long, twice length of 1

prostomium, fused basaily (Fig. 1.24A,B). Retracted pharynx extending to setiger 13, with anterior middorsal u

tooth; surrounded by distal circlet of 10 soft papillae when everted (Fig. 1.24C). Proventficulus extending

I
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I I GF H J K

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, Eigure 1.24. Ehtersia hyperioni: A, anterior end, holotype, dorsal view; B, anterior end, holotype, ventral view;

• i C, anterior end, lateral view; D, left parapodium, setiger 9, posterior view; E, right parapodium, I setiger 78, posterior view; F, inferior compound falciger, anterior setiger; G, superior compound

I

falciger, posterior setiger; H, inferior compound faiciger, posterior setiger, I, superior compound

I falciger, ventral simple posteriorseta,setiger, posterior with setiger; detail ofL,tippaired not to aciculae, scale; J, danterior orsal simple setiger; seta,M,posteri acicula, or setiger; posteriorK,

setiger, N, pygidium, holotype, dorsal view. (A-N, modified from Dorsey and Phillips, 1987)

i

through setigers 10-16 when pharynx inverted, 13 segments long when everted; 40-43 rows of muscle cells.

I cirrophores, Tentadorsal cular pair segmelongest, nt distinct each frwith om 7-15 prostomiu articles; m, nventral ot fusedpairtoeach setigerwith1;4-92 pairs articlesof (Fig. cirri, 1.24A-C). with short

Anterior setigers 1.5 times wider than long (Fig. 1.24A). Anterior parapodial lobes conical, distally

I blu falcigers nt, withdistally small pbidentate re- and postse (Fig.tal1.24F-I), lobes, gradually 8-11 per ante becrioming or fascicle, longer deposteri creasing orly to(Fig. 5-7 1per .24D, poste E). riorCompound fascicle.

Blades decreasing markedly in length inferiorly within fascicles. Superiormost blades 2-3 per fascicle, increasing

in length posteriorly along body, each 6-8.5 times longer than inferior blades, with fmely denticulate cutting

I 67
I

edges (Fig. 1.24G,I); inferior blades with cutting edges more coarsely serrated (Fig. 1.24F,H). Dorsal simple i

seta from middle setigers to end of body, unidentate, with coarsely serrated cutting surfaces (Fig. 1.24J).

Ventral simple seta haposterior setigers, sigmoid, distally bidentate, with finely serrated cutting surface (Fig. 'i

1.24K). Two aciculae per anterior parapodium, each distally blunt, sometimes dark (Fig. 1.24L), decreasing |

to 1 per posterior parapodium, stouter with pointed tips (Fig. 1.24M).

Dorsal cirri longest anteriorly, gradually decreasing posteriorly, not alternating in length, with short I

cirriphores; anterior cirri with 7-13 articles, middle and posterior cirri with 7-9 articles. Ventral cirri digitiform, I

short anteriorly, becoming longer posteriorly, extending just to end of parapodial lobes.

Pygidium with paired anal cirri, each with basal cirrophores and up to 18 articles; rnidventral unpaired 'i

papilla digit[form (Fig. 1.24N). n

Remarks. Ehlersia hyperioni and E. heterochaeta are sympatric. Ehlersia hyperioni differs from am

E. heterochaeta in lacking eyes, and in having superior blades of compound falcigers from middle body il

II

segments up to 10 times longer than inferiormost blades of the same fascicle in contrast to four times longer

in the same body region. Dorsey and Phillips (1987) provide morphometric data describing blades of compound

setae for both of these species, i

llll

Habitat. In mixed to fine sediments on the continental shelf, 12-146 m.

Distribution. Southern California to Washington. I

Genus Eurysyllis Ehlers, 1864 i

l1

Type species: Eurysyllis tuberculata Ehlers, 1864

Diagnosis. Body short, flattened. Prostomium with 3 globular antennae. Palps globular, not fused I

basally. Two pairs of globular tentacular cirri. Pharynx with a trepan of 10 teeth and a middorsal tooth.

lU

Dorsal cirri globular. Dorsum covered with longitudinal rows of globular papillae. I

Remarks. Hartman (I965:28) assigned EurysyIlis, without comment or subsequent discussion, to i

her new subfamily Eurysyllinae, from the Syllinae. The genus has also been associated with the Exogoninae.

Hartmann-Schrfder (1971) noted Hartman's subfamily and that Eurysyllis differs from Exogoninae species i

in palp morphology and pharyngeal armature, although its reproduction is more characteristic of the Syllinae. !i

Garwood (1991) synonymized the Eurysyllinae with the Syllinae, formally transferring EurysyUis to the

Syllinae primarily on the basis of reproductive similarities. Garwood's revision is followed here. i

Eurysyllis spicum Kudenov and Harris, new species

Figure 1.25 I

Material Examined. California: Santa Maria Basin, off Purisima Point, Stas. BRA-20, paratype

(SBMNH 142655); BRC-13, holotype (USNM 170901); between Point San Luis and Point Sal, Sta. BRA- II

25, paratype (USNM 170902).--Western Santa Barbara Channel, off Point Conception, Sta. BRA-2, paratype

!11

(LACM-AHF Poly 1660). I

Description. Holotype 2.2 mm long, 0.4 ram wide, without parapodia, 0.6 mm wide with parapodia, 1

for 49 setigers; paratypes with 56-66 setigers. Body ribbon-like, dorsum slightly arched in cross section,

i

ventrum flat; pigmentation absent in preserved specimens. Integument encrusted with fine sediment; thin, 1

delicate, transparent, internal septa visible. Golden-brown granules in antennae, cirri and papillae of freshly 1

preserved specimens (Fig. 1.25A). Sexual reproduction by stolons, beginning on setiger 40; sperm fill all but

I

last 2-3 segments in males. Ii

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I

!

I E

I

I

'

I

H I J KL M N O P Q

I

I Figure 1.25. EurysyUisspicum Kudenovand Harris,new species:A, anteriorsegments,dorsalview (SBMNH); B,prostomium,dorsalview (SBMNH);C, anteriorsegments,dorsalview (Sta.BRC-I3); D, anterior

segments, ventralview (Sta. BRC-13); E, right parapodium,setiger 11, posteriorview; E right

parapodium,setiger25,posteriorview; G,right parapodium,setiger50, posterior view; H,superior

I compoundfalciger,setiger 11;I, inferior compoundfalciger,setiger 11;J-K, superiorcompound falcigers,sefiger25; L-M, superiorcompoundfalcigers,setiger 50;N, inferiorcompoundfalciger,

setiger 50; O, acicula,setiger 11;P, acicula,setiger 25; Q, acicula,setiger 50. (A-Q, originalsby

I JDK)

Prostomium subrectangular, with large round lateral lobes, anterior middorsal region elevated; 3

i pairs palps;ofnumerous eyes; 2 pairs additional large, leeye nticula spotste, indispersed trapezoidal overarprostomium rangement, plus (Fig.add1.25A-D). itional ventral Antennae pair posspherical, terior to

papilliform; median antenna arising from anterofrontal swelling; lateral antennae arising from anterolateraI

I prostomial lobes (Fig. 1.25A-D). Palps spherical, papilliform, larger than antennae; basally free, projecting ventrally (Fig. 1.25D). Prostomium, antennae and tentacular cirri covered by thick cuticle; latter otherwise

thin. Pharynx from setigers 2-3, extending to middle of setiger 7; middorsal tooth at anterior pharyngeal

i border (Fig. 1.25A); encompassed by distal circlet of I0 large, soft lobes when everted. Trepan of 10 small, : sharply pointed, light-colored teeth. Proventriculus cylindrical, from middle of setiger 7 to 11, approximately

i 23 rows of muscle cells (Fig. 1.25A); attenuated extension lacking muscle cells in setiger 12.

i Tentacular segment narrower than following segments; dorsal tentacular cirri spherical, slightly larger than either lateral antennae or following dorsal cirri; ventral pair one-half length of dorsal pair, attached

ventrally, not visible from above (Fig. 1.25A-D).

I (Fig. 1.25E), Setigersbecoming 6-7 timesquared-off wider than loposteriorly ng (Fig. 1.25A). (Fig. 1.25F, Parapodia G). small, Setae generally distally rounded compound to conical falcigers anterior withly

short unidentate blades (Fig. 1.25H-N); 18-20 per fascicle in anterior segments, decreasing to 13-15 posteriorly.

I Number of setae per fascicle increase with body size. Blades longest in anterior fascicles (Fig. 1.25H,I),

I 69
!

!

decreasing in length both inferiorly within fascicles (Fig. 1.25H,I) and posteriorly along body, becoming more I

strongly curved posteriorly (Fig. 1.25J-N). Cutting edges of superiormost 5-8 blades of fascicles with long, It

fine teeth (Fig. 1.25J,K); remaining setae smooth (Fig. 1.25I,N). Shaft tips of superiormost 5-8 setae within

fascicles finely denticulate (Fig. 1.25H-I); remainder normally smooth (Fig. 1.25L-N). One acicula per

parapodium, stout with bent, blunt tip (Fig. 1.250-Q). Ventral simple seta present in 8 or fewer posteriormost n

setigers.

Dorsal cirri with single distal papilliform article attached to stout cirrophores, extending beyond n

parapodial lobes (Fig. 1.25E-G). Ventral cirri conical anteriorly (Fig. 1.25E,F) to digitiform posteriorly (Fig.

U

1.25G), extending beyond parapodial lobes, especially in posterior segments. m

Dorsum with 4, occasionally 5, large globular papillae per segment, forming 4 longitudinal rows; I

Fifth papilla located next to cirrophore (Fig. 1.25A).

Pygidium with 2 anal cirri similar in form to dorsal cirri, larger, distally pointed; cirrophores

conspicuously developed. I

Remarks. Four species of Eurysyllis are presently recognized: E. spicum, E. tuberculata (Ehlers,

1864), E. ehlersi Benham, 1927, and E. pacificus (Hartman, 1954). Eurysyllis spicum, E. tuberculata and E. B

pacificus all have longitudinal rows of conspicuous dorsal papillae, and are here considered members of this U

genus. In contrast, E. ehtersi clearly lacks dorsal papillae based on Benham's description, and most likely

should be transferred to Plakosyllis Hartmann-Schr6der, 1956. This decision, however, will be deferred i

pending examination of Benham's type, and E. ehlersi will continue to be considered a congener of the other II

three species treated here.

Eurysyllis spicum differs fromE, tuberculata in having aciculae distally blunt rather than mucronate,

the pharynx present in four to five segments instead of two to three segments, the proventriculus with 23 rows II

of muscle cells present in four segments in contrast to 13-20 rows in three segments, ventral cirri conical to

digitiform instead of auricular, and unidentate blades of compound falcigers within the same fascicles both

serrated and smooth instead of all serrated. In this latter feature, E. spicum also differs from E. pacificus in

which the unidentate blades are smooth, and more noticeably curved distally. Additional published information

providing detailed descriptions of E. pacificus are apparently nonexistent, and efforts to locate Hartman's

paratype (originally described as Sphaerodorum pacificum) inthe LACM-AHF collections were unsuccessful. l!

Ltitzen (1961) first recognized the aberrant nature of Hartman's species, and Laubier (1968) strongly suspected

it was a species of Eurysyllis. More detailed comparisons between E. spicum and E. pacificus must await

examination of the latter's holotype. Eurysyllis spicum differs from E. ehlersi in having the pharynx in four II

to five rather than six segments, the proventriculus in four instead of eight segments, blades of compound

falcigers unidentate and not bidentate, and ventral cirri conical to digitiform rather than ovoid and projecting

beyond parapodial lobes.

Finally, Hartman (unpublished) seems to have suspected thatPlakosyllis americana Hartman, 1961,

had longitudinal rows of papillae on the dorsum, and should be assigned toEurysyllis. Re-examination of the

Plakosyllis americana types (LACM-AHF Poly 0865, 0866) did not confirm this suspicion, although the

dorsal papillae are rather easily detached during the examination of preserved specimens ofE. spicum.

Etymology. The term, spicum, is Lat/n for "ear of grain," and refers to the kernel-like arrangement i

I

of large dorsal papillae. !

Distribution. Southern and central California. I

!

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I Genus Geminosyllis Imajima, 1966d

Type species: Trypanosyllis (Trypanedenta) ohma Imajima and Hartman, 1964

I Diagnosis. Body subcylindrical. Prostomium with 3 antennae. Two pairs of tentacular cirri. Antennae,

tentacular cirri and dorsal cirri slender, articulated. Pharynx with a trepan of 10 teeth with or without a single

I large middorsal tooth. All setae simple.

I Geminosyllis ohma Clmajima and Hartman, 1964) Figure 1.26

I TrypanosyIIis (Trypanedenta) ohma Irnajima and Hartman, 1964:129-130, pl. 31, figs. a-d. Syllis (Haplosyllis)spongicola: Uschakov, 1955:179, fig. 50, A.

Geminosyllis ohma: Imajima, 1966d:233-235, text-fig. 43, a-g.

I Material Examined. California: Santa Maria Basin, between Point San Luis and Point Sal, Sta.

BRA-25 (1); off Point Arguello, Sta. BRA-6 (1, SBMNH 142704).--Western Santa Barbara Channel, east

I of Point Conception, Sta. BRC-1 (6).--Japan: Shifikishinai, 20 July 1959, holotype (LACM-AHF Poly 0857).

i Description, Length up to 32 mm, 2 mm wide including parapodia, with 82 setigers (Imajima and Hartman, 1964); MMS specimen (SBMNH) complete, 3.5 mm long, 0.2 mm wide excluding parapodia and

setae, for 31 setigers. Body subcylindrical; preserved specimens lacking color patterns.

i anterior Prostomium pair pentagonal in outline, posterior margin entire, lacking median cleft; 2 pairs of eyes, reniform, largest, farthest apart; posterior pair round (Fig. 1.26A,B). Median antenna inserted

between posterior pair of eyes, with 30 annulations (18 in MMS specimen); lateral antennae arising from

i anterior prostomium margin, each with about 25 annulations (11-13 in MMS specimen) (Fig. 1.26A,B). Palps thick, large, conspicuous, broadly triangular (Fig. 1.26A). Everted pharynx long, cylindrical, with

distal circlet of 10 soft papillae, chitinized lining dark brown (inverted pharynx in setigers 1-8 in MMS

i specimen); trepan with 10triangular teeth plus a sharp subterminal middorsal tooth (Fig. 1.26B). Pmventriculus present in setigers 12-21 (in setigers 9-12 in MMS specimen), with 40-50 rows of muscle cells 0aolotype with

44 cells; MMS specimen with 50).

I dorsal tentacular Dorsal tentac cirriu(Fig. lar cirri 1.26A,B). about as long as lateral antennae; ventral tentacular cirri one-half length of

Parapodia short, distally blunt, with presetal lobe (Fig. 1.26C). Setae simple, numbering 6-7 per

I fascicle; (Fig. 1.26D,E). distally Ventr bifid,alsusimple bdistallseta y enlarge presentd oinr spoon-sh far posterior aped,segments with smooth (in lastto 10finelsetigers y serrated in MMS cuttingspecimen), surfaces

distally bifid (Fig. 1.26F). Aciculae distally rounded, 4 per median parapodia, decreasing to 1 in posterior

i segments (Fig. 1.26G). Dorsal cirri of anterior segments alternating between long ones with 35 articles and short ones with

about 30 articles (Fig. 1.26A,C); in MMS specimens, dorsal cirri of setiger 1 with 16 articles, short dorsal

I cirri Ventral of setiger 2 with 8-9, long alternating with 2 short of midbody segments with 8 and 4, respectively. cirri digitiform, extending just beyond parapodium (Fig. 1.26C).

Pygidium with long anal cirri (with 11 articles in MMS specimen).

I Remarks. This is the first formal record of this species from the northeastern Pacific. The MMS

specimens are generally consistent with the features previously described for Geminosyllis ohma, although

they are significantly smaller in body size and sexually immature.

I Distribution. Japan; Russia (Kurile Islands); southern California.
I

I

I

I

B I

A

- I

I

I

i

I

I

// c I

If !

D E F G II

Figure 1.26. Oeminosyllis ohma: A, anterior end with pharynx everted, dorsal view; B, anterior end, detail of i

pharyngeal armature, dorsolateral view; C, parapodium 18; D, bifid simple seta with smooth cutting

edge from anterior parapodium; E, bifid simple seta with minute serrations along cutting edge from I same anterior parapodium as D; F, ventral simple seta, posterior parapodium; G, acicula, posterior

parapodium. (A-G, redrawn from Imajima, 1966d)

I

I

72 I
I

I Genus Syllis Lamarck, 1818

i Type species: Syllis monilaris Lamarck, 1818

Diagnosis. Prostomium with 3 antennae. Two pairs of tentacular cirri. Antennae, tentacular cirri

and dorsal cirri all strongly articulated. Pharynx with a single middorsal anterior tooth, smooth marginal rim

I and distal circlet of 10 soft terminal papillae. Compound falcigers present in all setigers and pseudocompound

setae and a superior and/or inferior simple seta variably present from median segments.

I pseudocompound Remarks. Species of SyUis are readily distinguished from those of TyposyIlis by the presence of setae. Refer to the Remarks section following the diagnosis of Typosyllis for further comments.

I SyUis spongiphila Verrill, 1885

Figure 1.27

I Syllis spongiphila Verrill, 1885: 435.--Hartman, 1944: 339, pl. 24, fig. 10.--Pettibone, 1963:114-115, fig.

30 g, h.--Banse and Hobson, 1974: 61, fig. 16p.--Imajima, 1966d:250-251, text-fig. 491-s.

Syllis sclerolaema Ehlers, 1901: 86, pl. 10,figs. 1, 2.--Monro, 1930: 102, fig. 35.--Berkeley and Berkeley,

I 40, fig. 5.--Hartrnan, 20, fig. 2; 92.--Uschakov, 179, fig. 50, d.--Imajima

1938: 1953: 1964: 1955:

and Hartman, 1964: 122-124, pl. 28, figs. i,j, pl. 29, figs. a-i. Fide Imajima, 1966d.

i Syllis (Typosyllis) sclerolaema Wesenberg-Lund, 1962:54, fig. 15. Fide Imajima, 1966d.

Material Examined. California: Santa Mafia Basin, Stas. BRA-20 rock (3, SBMNH 142705);

I B170938).--Western RA-13 (1); BRC-13Santa (1); BRBarbara A-16 Channel, (3); betweeneastPoin of Point t SanConception, Luis and PointSta.Sal,BRSta. C-1 Brock RA-25(4);rock south(1,ofUSNM Point

Conception, Sta. BRC-2 (1).

I Description. Length about 27 mm, 1.5 mm wide with parapodia, excluding setae, with about 90 segments (I-Iartman, 1944; Imajima, 1966d); complete MMS specimens (Sta. BRC-1) up to 13.5 mm long,

0.4 mm wide excluding parapodia and setae, for 68 segments. Body pigmentation patterns absent in preserved

I specimens. Mature specimen with a few oocytes in setigers 17-42 (Sta. BRC-1).

Prostomium slightly wider than long, subglobular; posterior margin entire; 2 pairs of eyes, set in

trapezoidal arrangement, anterior pair largest; additional pair of eyespots absent (Fig. 1.27A). Median antenna

I with 25-30 articles (MMS specimens with 18-22), arising from center of prostomium; lateral antennae with

slightly fewer articles (MMS specimens with 15), each arising anterior to anterior eyes (Fig. 1.27A). Palps

large, subtriangular, basally free, about as long as prostomium (Fig. 1.27A). Everted pharynx in setigers 1-

I 13 with middorsal anterior tooth; with distal circlet of 10 soft papillae and smooth anterior margin.

• Proventriculus extending from setiger 14 through setigers 22-24, with 47 rows of muscle cells.

Twp pairs of tentacular cirri, resembling antennae; dorsal pair about as long as median antenna

I (MMS specimens with 17-22 articles); ventral pair about two-thirds length of dorsal pair (MMS specimens

with 9-14 articles) (Fig. 1.27A).

i obviouslyParapodi compound a conical falcigers (Fig.with1.27B). bidentateSetaeblades generaandlly serrated pseudocompo cuttingundedges(Fig.usually 1.27C-Epresent ), although in anterior 3-5

fascicles (Fig. 1.27C). Pseudocompound setae with distally bidentate tips numbering 6-8 per fascicle in

m median (Fig. 1.27E) andposterior fascicles, sometimes present also in anterior fascicles (Fig. 1.27D). Ventral

i I simple seta slender, distally bidentate with subtly serrated cutting surfaces present in posterior parapodia

(Fig. 1.27F). Aciculae distally curved, numbering 3-4 per parapodium in median segments (Fig. 1.27G).

i

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I

I

I

i # II

.,oo i

G I

I

Pigu_ 1.27. Syllis c

F,

om

ventral

posite spongiphila f

simple

alciger,:

seta,

setig A, anterior

posterior

er 1; D, enp

parapodium;

seud d, dorsal ocomposite vie

G,

w;

aciculae;

falciger, B, parapodium setiger

H, short

1;21E,

dorsal

w

same ith

cirrus,

,

long mediandorsal

parapodium

parapodium; cirrus;

22.

C, I

(A-H, redrawn from Imajima, 1966d) I

I

Dorsal cirri articulated, alternating in length on median and posterior segments, and occasionally also won anterior segments (Fig. 1.27B,H); long cirri on setiger 1 each with 30-36 annulations (MMS specimens I

with 15-20); short cirri on setiger 2 each with 20 (MMS specimens with 9-10) (Fig. 1.27A); thereafter I

alternating with long cirri having 20-40 articles (MMS specimens with 8), and short ones with 16-30 (MMS I

specimens with 5). Ventral cirri slender, digitiform, extending to ends of parapodial lobes (Fig. 1.27B). i

Pygidium with long paired anal cirri. I

Remarks. As presently recognized, Syllis spongiphila is a widespread species associated with

poriferans, mussels, mud, sand, gravel, and hard surfaces such as shells and rocks. I

l]a_stribution. Southern California; Washington; British Columbia; Japan; Sea of Okhotsk; Chile; I

Falkland Islands; Massachusetts.

!
!

I Genus Trypanosyllis Clapar_de, 1864

I Type species: SyUis zebra Grube, 1860.

I ProstomiumDiagnosis. with 3 antennae. Body long,Palps usually separate dorsoventrally basally. Nuchal flattened, organswithconspicuous numerous short, ridges wi along de segments. postectal

margin of prostomium. Two pairs of tentacular cirri. Antennae, tentacular cirri and dorsal cirri strongly

I articulated Setae simple . Pharynx or compound with terminal falcigers.trepan of 10 teeth, with or without an additional middorsal anterior tooth.

Remarks. Imajima and Haxtman (1964) distinguish two major groups representing three subgenera

I of Trypanosyllis on the basis of the pharyngeal middorsal tooth and setae. For example, the first major group, represented by the stem genus Trypanosyllis (Trypanosyllis) is defined by the presence of a pharyngeal trepan

with a middorsal tooth. The other major group has a trepan and lacks a middorsal tooth, and is characterized

I byhastwo subgenera: TrypanosyIlis (Trypanedenta) has compound setae only; and Trypanosyllis (Trypanobia) simple setae only. This scheme is followed below.

I Subgenus Trypanosyllis (Trypanosyllis) Imajima and Hartman, 1964

i Type species: Trypanosyllis (Trypanosyllis) zebra (Grube, 1860)

I cirri longDiagnos or shortis,.closely Trepanarticulated. usually suSetae rroundusually ed by flesh compound y distalfalcigers circlet owith f 10uni- soft ortermin bidentate al papillae. blades; Dsimple orsal

setae may be present in posterior segments. Tetraglene individuals produced asexually by budding of posterior

I segments.

I Key to the Species of Trypanosyllis (Trypanosyllis) from California

I 1A. Blades of superior compound falcigers distally bidentate with serrated cutting edges, becoming unidentate and smooth inferiorly within fascicles (Figs. 1.28B-D, 1.29D-G) ................................. 2

lB. Blades of all compound falcigers distally bidentate, cutting edges of superior blades smooth,

I becoming serrated inferiorly within fascicles (Fig. 1.30F-H); proventriculus extending through 4-5 segments ............................................................................. Trypanosyllis (Trypanosyllis) sp. B

I 2A. Proventriculus in 7 segments; parapodia distally conical (Fig. 1.28A); aciculae not projecting through

parapodia; ventral cirri not projecting beyond parapodial lobes; occipital flap absent ......................

I ............................................................................. Trypanosyllis (Trypanosyllis) coeliaca nipponica 2B. Proventriculus in 3.5 segments; parapodia distally rounded (Fig. 1.29B,C); aciculae projecting through

parapodia (Fig. 1.29B,C); ventral cirri projecting beyond parapodial lobes; occipital flap may be

I present (Fig. 1.29A) .................................................................. Trypanosyllis (TrypanosyIIis) sp. A

!

i 75
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Trypanosyllis (TrypanosyUis) coeliaca nipponica _aj_a and Ha_man, 1964

I

Figure 1.28 I

Trypanosyllis (Trypanosyllis) coeliaca nipponica Imajima and Hartman, 1964:124-125, pl. 29, fig. j; pl. 30,

figs. a-e.--Imajima, 1966d:236-237. I

Material Examined. California: Santa Maria Basin, Sta. BRA-16 (1, USNM 170940).--Japan: •

Shirikishinae, holotype (LACM-AHF Poly 858) and paratypes (LACM-AHF Poly 859).

Description. Length up to 17 mm, 2.2 mm wide; with 140 setigers Clmajima and Hartman, 1964);

MMS specimens about 4.5 mm long, 0.5 mm wide without parapodia, 0.7 mm wide with parapodia, for 63 I

setigers. Body strongly flattened, lacking color patterns in alcohol. Prostomium rectangular, 2 times wider I1

than long, lacking obvious median cleft; 2 pairs of eyes, anterior pair largest (Fig. 1.28A). Palps conical, not

fused (Fig. 1.28A). Median and lateral antennae stout, short, each with 10-12 distinct articulations; all

arising from anterior margin of prostomium (Fig. 1.28A). Everted pharynx with distal circlet of 10 soft II

papillae; trepan with 11teeth including middorsal one. Proventriculus short, extending through setigers 13 to

junction of setigers 18-19. Parapodia short, conical (Fig. 1.28B). Setae generally compound falcigers (Fig. n

1.28C-E). Blades with finely serrated cutting margins; superior blades within fascicles distally bidentate m

(Fig. 1.28C); middle and inferior blades distally unidentate with single accessory tooth (Fig. 1.28D); inferior

blades unidentate (Fig. 1.28E); (uncommon in MMS specimens). Shafts with subdistal serratia on superior l

surfaces (Fig. 1.28C-E). Posteriormost parapodia each with a ventral simple seta with subdistal tooth. Dorsal U

cirri thick, short, with 8-14 annulations (MMS specimen with 10-13). Ventral cirri digitiform, not extending

beyond parapodial lobes (Fig. 1.28B). Aciculae numbering 1-2 per parapodium, tapering to pointed tips (Fig. I

1.28F). Pygidium with pair of short anal cirri.

Remarks. This is the first formal record of Trypanosyltis (Trypanosyllis) coeliaca nipponica from

the northeastern Pacific. The MMS specimens are generally consistent with the features previously described 1

for the subspecies, although they are significantly smaller in body size. It

Distribution. Northern Japan; southern California. U

°,

U

l

Figure 1.28. Trypanosyllis (Trypanosyltis) coetiaca nipponica: A, anterior end, dorsal view; B, median _l

parapodium; C, superior compound bidentate falciger, setiger 6; D, median compound falciger, II

setiger 6; E, inferior compound unidentate falciger; F, acicula, distal end. (A-F, redrawn Imajima

and Hartman, I964) 1

1

76 !
I

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I TrypanosyUis (Trypanosyllis) sp. A

Figure 1.29

I Material Examined. California: Santa Maria Basin, Stas. BRA-13 (1, USNM 170941), (I); BRA16 (3); BRA-20 (4)--Western Santa BarbaraChannel, south of Point Conception, Sta. BRC-2 (2).

I Description. Complete specimens 4 mm long, 0.7 mm wide excluding setae, for 61 setigers. Body strongly flattened dorsoventrally; lacking pigmentation patterns, pale yellow in alcohol.

Prostomium subrectangular, shaped somewhat like a bowtie; 2 pairs of eyes, connate, lenticulate,

I anterior pair all 1 Antennae arise from anterior frontal of

largest, arranged trapezoidally (Fig. .29A). margin

prostomium; median antenna slightly longer than prostomium, with about 8 articles; lateral antennae smaller,

with 5-6 articles (Fig. 1.29A). Palps small, globular, shorter than prostomium, separate (Fig. 1.29A). Pharynx

I extending to setiger 10. Proventriculus present in setigers 11-14 or 15. Trepan partly visible through body wall, with obvious middorsal tooth plus 10 irregular denticles, all sharply pointed (not dissected).

i Tentacular segment reduced dorsally; occipital flap small, marginally ciliated (Fig. 1.29A). Two pairs of tentacular cirri; dorsal pair largest, each with 3 distal articles and long cirrophore together nearly as

long as prostomium; ventral pair smaller, each with 1-2 articles, about one-half length of dorsal pair (Fig.

1.29A).

I Parapodia well developed, broadly rounded distally (Fig. 1.29B,C). Setae compound falcigers (Fig.

1.29D-G), numbering 7-8 per anterior fascicles, decreasing gradually to 3-5 in far posterior fascicles. Blades

i dedge ecreasing each inlong, lengthcoarse inferior (Fig. ly within 1.29D);fascicles: intermediate superior blades onesalso longest, bidentate, distallycutting bidentateedges withwith serraatiafewof cutting poorly

defined beadlike enlargements or smooth (Fig. 1.29E,F); inferior blades unidentate, smooth (Fig. 1.29G).

i One to 3Dorsal aciculaecirriperofparapodi setiger u1m,withthick, 8 distinct sgmoid, articles projecti (Fg. ng 1.29A), throughthose parapodial following lobeseach(Fig.generally 1.29H). with 7,

decreasing to 6 posteriorly (Fig. 1.29B,C), last few segments each with 4 articles (Fig. 1.291); all with

I cirrophores, parapodial lobes, each present around on2 time posterior s longer facethan of parapodia wide. Ventr (Fig.al 1.29B,C). cirri bluntly conical, extending just beyond

Pygidium with paired anal cirri, tapered distally, each with 2 large articles (Fig. 1.291).

I Remarks. The MMS specimens have both bidentate and unidentate compound falcigers within the

same fascicle, a feature characteristic of Trypanosyllis (Trypanosyllis) coeliaca nipponica. In addition, the

median and lateral antennae of both species also arise from the frontal margin of the prostomium. Trypanosyllis

I (TrypanosylIis) sp. A differs from T. (Trypanosyllis) coeliaca nipponica in having an occipital flap and

generally having compound falcigers withdifferent blade morphology, although the superior compound falcigers

of the two taxa are quite similar. It is probable that present materials of T. (Trypanosyllis) sp. A are all

I juvenile specimens and that the above differences are transient juvenile features: the specimens all are rather small and sexually immature. It therefore seems appropriate to note the apparent similarities between the

present materials and T. (Trypanosyllis) coeliaca nipponica, and to tentatively identify these specimens as T.

I Trypanosyllis sp. A, pending acquisition of additional specimens. Habitat. Rocky substrata, in depths of 75-237 m.

i Distribution. Southern and central California.

i
!

!

B !

I

B.C I

_._ .05 mm

D-G

D E F G H 0t._, i

l

Figure 1.29. Trypanosyltis (Trypanosytlis) sp. A: A, prostomium and setiger 1, dorsal view, (USNM); B, right

parapodium, setiger 13; E,setiger intermediate 13, anterior compound view;falciger, C, same,setiger posterior 13; Eview; low intermediate D, superior ccompound ompound falciger, falciger, I

setiger 13; G, inferior compound falciger, setiger 13; H, acicula, setiger 13; I, Posterior end, dorsal

view. (A-I, originals by JDK) i

!

Trypanosyllis (TrypanosyUis) sp. B i

Figure 1.30 l

II

Material Examined. California: Santa Maria Basin, Stas. BRA-16 rock (1, SBMNH 142707);

BRA-20 rock (1, USNM 170942); west of Point Conception, Sta. BRA-4 (1, SBMNH 142708); between n

Point Estero and Point Buchon, Sta. 6, rocks (1). l

llll$

Description. Length up to 2 turn, 1 mm wide excluding setae, for 29 segments including 10 regenerating

posterior setigers (SBMNH voucher); USNM voucher an anterior macerated fragment, poorly preserved, 1.5 •

rnm long, 0.8 mm wide excluding setae, for 18 setigers; all other specimens smaller. Body dorsoventraIly |

flattened; pigmentation patterns absent in alcohol.

Prostomium wider than long, with posteromedian cleft; 2 pairs of eyes, lenticulate, anterior pair

largest, all trapezoidally arranged (Fig. 1.30A,B). Antennae usually arising from anterofrontal margin of II

prostomium, all slightly longer than prostomium; median antenna with 9-12 distinct articles; lateral antennae

each with 7-9 articles (Fig. 1.30A,B). Palps small, globular, one-half length of prostomium, separate (Fig. •

1

78 |
I

I

I A

I F

C D

'

C.D

I .o, o FG H J K

I

Figure 1.311. Trypanosyllis (Trypanosyltis) sp. B: A, prostomium and setigers 1-2, dorsal view (USNM); B,

i prostomium, dorsal view (SBMNH); C, everted pharynx showing trepan, dorsal view (SBMNH); D, everted pharynx showing trepan, ventral view; E, median segment, anterior view; F, superior

compound falciger, median setiger; G, intermediate compound falciger, median setiger; H, inferior

compound falciger, median setiger; I, ventral simple seta, setiger 25; J, inferior acicula, median

I setiger; K, superior acicula, median setiger.(A-K, originals by JDK)

I .30A,B). Pharynx extending to setigers Trepan plus (Fig..30C,D).

1 7-8. with middorsal tooth 10 distal teeth 1

Proventficulus extending through 4-5 segments, present in setigers 8-12 or 9-14.

i

short cirrophores Tentacularcharacteristically segment distinct,vesiculate; reduced dorsally dorsal pair (Fig.longest, 1.30A,B). longer Twothanpairsmedian of tentacular antenna,cirri, withall12-14 with

articles; ventral pair one-half length of dorsal pair, each with about 5-7 articles (Fig. 1.30A,B).

I 1.30F-H),Parapodia numberingtapering aoundslightly, 8-10 perdistally anteriortruncate fascicles,(Fig. decreasing 1.30E). inCompound median andsetae posterior distallyfascicles. bidentateBlades (Fig.

subequally long within fascicles (Fig. 1.30F-H), length decreasing gradually posteriorly. Cutting edges of

I superior superior blad endsesofsmooth shafts dentate (Fig. 1.(Fig. 30F),1.30F-H). becoming Ventral conspicuously simple seta serrated distallyinferiorly bidentate, (Fig.sigmoid, 1.30G,H). sometimes Distal

present in far posterior segments (Fig. 1.30I; illustrated for specimen from Sta 6). Aciculae numbering 2-3

per parapodium, projecting through distal lobe; each with a characteristic ventrally thickened, hook-like subdistal

I collar (Fig. 1.30J,K).

Dorsal cirri short, each with 10-12 articles, not looping back on themselves; cirrophores short, each

I characteristically 1.30E). vesiculate (Fig. 1.30A,E). Ventral cirri digitiform, extending beyond parapodial lobes (Fig.

Pygidium (SBMNH secondary voucher) with paired anal cirri, regenerating, each with 5-6 articles.

I
1

!

Remarks. MMS materials of Trypanosyllis (Trypanosyllis) sp. B were originally identified in part

as T. (Trypanosyllis) brevicirrata Uschakov, 1950, based on the presence of short dorsal cirri and the apparently I

unidentate falcigers in all setigers. All present MMS materials have bidentate compound falcigers, have a

IB

small body size, and lack gametes. It seems likely that specimens here identified as T. (Trypanosyllis) sp. B

are unidentifiable juveniles. All are substantially smaller than mature forms of described Pacific west coast D

species which normally have large body sizes and numerous segments. For example, T. (Trypanedenta)

ingens Johnson, 1901, is 130 mm long for 476 segments; T. (Trypanedenta) gemmipara Johnson, 1901, is 40-

68 mm for 385 segments; and T. (Trypanedenta) intermedia Moore, 1909, is 60 mm for 230 segments. •

However, Perkins (1981) described sexually mature specimens of T. (Trypanosyllis) savagei from Florida II

less than 5 mm for 80 segments. All MMS specimens here referred to T. (TrypanosyUis) sp. B are less than

2 mm long, and are sexually immature. Two of these specimens are regenerating body parts: one a head plus

anterior segments, the other posterior segments; none represent stolons. II

Habitat. Rocky substrata, in depths of 91.5-237 m.

Distribution. Southern and central California. I

Subgenus Trypanosyltis (Trypanedenta) Imajima and Hartman, 1964 1

I

Type species: Trypanosyllis (Trypanedenta) gemmipara Johnson, 1901

Diagnosis. Trepan with 10 teeth, middorsal subdistal tooth absent. Dorsal cirri thick, heavily I

articulated. Setae compound falcigers with uni-or bidentate blades. Tetraglene individuals produced asexually

by budding of posterior segments. I

HI

Trypanosyllis (Trypanedenta) sp. A

Figure 1.31 I

Material Examined. California: Santa Maria Basin, Sta. BRA-21 (1, USNM 170939).--

Trypanosyllis (Trepanedenta) gemmipara: Tomales Bay, 9 June 1941, from bryozoan-sponge masses, coll.

O. Hartman (LACM-AHF, Hartman lot N47).--La Jolla, Bird Rock, 30 May 1938, coll. O. Hamnan (1,

w

LACM-AHF, Hartman lot NI031); north of Scripps Pier, 13Feb 1938, coll. O. Hartman (1, LACM-AHF,

Hartman lot N1032).--Japan, Shirikishinai, October 1955, coll. M. Imajima (1, LACM-AHF, Hartman lot

Nl1086).

I

Description. Length approximately 22 mm, 2 mm wide excluding parapodia, for around 120 segments, m

Body twisted into spiral, poorly preserved anteriorly, lacking antennae, tentacular and dorsal cirri;dorsal cirri 1

present posteriorly; dorsum flesh colored, lacking color patterns; sexually mature.

Prostomium wider than long, with conspicuous posteromedian cleft, trapezoidal in shape, anterior •

margin widest. Two pairs of eyes, all lenticulate, anterior pair largest and farthest apart, all in subrectangular U

arrangement. Antennae lost. Palps about as long as pmstomium, free basally, projecting ventrally. Pharynx

long, extending to midsetiger 16; distally with 5 ventral and 7 dorsal terminal papillae, each group separated •

from the other by lateral gaps (dissected). Trepan with 10 pale brown teeth, largest appearing midventral; I

middorsal tooth absent. Proventriculus present from setiger 16 to 28; 39 muscle rows, each with conspicuous

muscle ceils. IB

Tentacular segment incomplete dorsally. Tentacular cirri lost. n

I

|
!

!

!

I A _

.lmm .01o . D E F G

I

Figure 1.31. Trypanosyllis(Trypanedenta)sp. A:left parapodiumfrom far posterior segment, posterior view; BI C, compound falcigers from setiger 15; D-F, compound falcigers from far posterior segment: D, superior seta; E, intermediate seta; F,inferior seta; G, aciculae from far posterior segment. (A-G,

originals by JDK)

!

Parapodia long, distally pointed (Fig. 1.31A). Setae generally delicate, bidentate compound falcigers

I (Fig. 1.31B,C). Blades long, narrow with secondary tooth subdistal to primary tooth in most all falcigers (Fig. 1.3 IB,C), becoming more widely spaced in far posterior segments (Fig. 1.31D-F). Blades with cutting

edges minutely serrated (Fig. 1.31B-F). Aciculae straight, projecting through parapodial lobes, up to 5 per

I anterior parapodium, decreasing to 1-2 posteriorly (Fig. 1.31G).

Dorsal cirri thick, alternating long and short along the body; in midposterior segments long cirri each

with around 45 articles, short cirri each with 27. Ventral cirri short, smooth, digitiform, may extend to and

I sometimes lobes in anterior and beyond parapodial midbody segments.

Remarks. TrypanosylIis (Tranpanedenta) sp. A was collected from one location in Santa Mafia

i Basin. midsetiger Although poorly preserved, the species is partly characterized by having a pharynx that extends to 16, with 2 distal groups of terminal papillae (5 ventral and 7 dorsal ones, observed by dissecting

the inverted pharynx), which are separated from one another by lateral gaps, and a trepan with I0 teeth, the

i largest tooth appears to be midventral; a proventriculus in 13 segments (setiger 16 to 28) with 39 rows of

conspicuously large muscle cells; delicate bidentate compound falcigers with blades long and narrow, secondary

teeth all subdistal, and cutting edges minutely serrated in all fascicles; and ventral cirri extending to or slightly

beyond parapodial lobes.

I Trypanosyllis (Trypanedenta) sp. A is similar to specimens of T. (T.) gemmipara Johnson, 1901,

noted above (see Materials Examined) in having bidentate compound falcigers in all setigers; it differs from Z

i (Tand.) gemmipara narrow withincutting havingedges all secondary minutely teeth serrated, subdistal insteadrather of short, than stout in thetriangular middle ofblades, the blade; withblades cuttingalledges tong

smooth or serrated; 12 terminal pharyngeal papillae arranged in a ventral and dorsal group, in contrast to 8 or

i 11; a trepan of 10 teeth,the largest one of which appears to be midventral, instead of 8 or 11; and a pmventriculus present in 13 segments rather than 4 or 8 segments.

The compound falcigers of the above MMS specimen are similar to those described by Imajima

I (1966d) is sexuallyfrommature, inferiorandandis posterior not a juvenile fascicles T. (T.of)gemmipara T. (T.)gemmipara . In addition, from Japan. an examination However,oftheT. MMS (T.) gemmipara specimen

specimens from California and Japan revealed that all had well-developed compound falcigers with stout,

I smooth triangular blades with the secondary tooth located about midway along the cutting edge; none had

|
I

I

long, narrow, distally bidentate blades with serrated cutting margins characterized by Imajima (1966d). Although •

the poorly preserved MMS specimen precludes a more detailed analysis, these findings suggest that the taxon

it represents is not conspecific with T. (T.) gemmipara. Resolution of this question must await the acquisition _l

of additional materials. II

Finally, Imajima and Hartman (1964) do not describe the presence of a large midventral pharyngeal

tooth in the genus Trypanosyllis. Such a character may be important at the level of subgenus. If this is true, B

then the species represented by the MMS specimen described above may represent a new subgenus if the

pharynges of all known T. (Trypanedenta) species are found to lack a midventral tooth.

Habitat. Associated with rocky substrates in depths of 75-90 m. !

In Distribution. Southern and central California.

Genus Typosyllis Langerhans, 1879 I

Type species: Syllis krohnii Ehlers, 1864 I

W

Diagnosis. Prostomium with 3 antennae. Two pairs of tentacular cirri. Palps broadly triangular,

fused basally. Antennae, tentacular cirri and dorsal cirri strongly articulated. Pharynx with single middorsal 1

anterior tooth, with smooth marginal rim, and distal circlet of 10 softterminal papillae. Setae usually compound II

falcigers with uni- or bidentate blades, with a superior and/or inferior simple seta variably present in posterior

setigers.

Remarks. Fauchald (1977 ) notes 89 described species of TyposyUis,which is one of the most complex II

and systematically challenging of syllid genera. Some workers include Typosyllis (andEhlersia, Haplosyllis)

as subgenera ofSyllis (Fauvel, 1923; Day, 1967; Uebelacker, 1984; Dorsey and Phillips, 1987). Haplosyllis •

has simple setae, and is readily separable from the other three taxa. Note that the separation of Syllis sensu

lato into three separate groups is based on setal traits not normally used to separate other syllid genera, and

fails to reduce the nearly incomprehensible complexity ofTyposyllis (Uebelacker, 1984). This strongly suggests •

that traditional means of identifying and separating Typosyllis species are inadequate, and that additional I

characters are needed before many of the systematic issues concerning TyposyIlis can be resolved. Such an

undertaking is well beyond the scope of this chapter. However, we prefer to recognize four separate but •

closely related genera for matters of practicality. Besides the approximate number of Typosyllis species noted |

above, at least 45 SyUis species, 16 Ehlersia species, and 10 Haplosyllis species have also been reported

(Fauchald, 1977; Pettibone, 1982; Dorsey and Phillips, 1987). A single large genus (SylIis) with over 150 •

species tends to obscure the critical and urgent need to revise this taxon based on combinations of both 1

traditional and new characters. To this end, Piltz (1980) provides an example of the kinds of characters that

can be used to further resolve the complexity surrounding Typosyllis and Syllis. I

m

Key to Species of Typosyllis from California 1u

1A. Dorsum with 2 reddish brown transverse lines per anterior segment inlife (instead of faded brown in 1

preserved material) (Fig. 1.33B); palps not fused basally (Fig. 1.33A); dorsal cirri thick, alternating 1

in length, long cirri of median segments each with 25 and short cirri with 14 articles (Fig. 1.33C)..

m

.................................................................................................................................. Typosyllis bella I

lB. Dorsum with transverse bands of dark pigment on every 1-3 segments (Fig. 1.34A), or lacking any I

pigment pattern; palps fused basally (Figs. 1.3213,1.34A); dorsal cirri usually slender, alternating in

length, each with fewer articles in midbody segments ...................................................................... 2 I
!

I 2A Dorsum with broad transverse bands of broken brown 1-3 usually pigment on every segments (Fig.

1.34A), or lacking pigment pattern; long dorsal cirri of median segments each with 14 (Fig. 1.34C)

and short cirri with 11 articles (Fig. 1.34B); compound falcigers with short, stout blades (Fig. 1.34DI ......................................................................................................................... Typosyllis hyalina

G)

2B. Dorsum without pigmentation patterns; long dorsal cirri of median segments each with 25 (Fig.

i 1.32C) and short cirri with 18 articles; compound falcigers with longer, more narrow blades (Fig. 1.32F-I) ............................................................................................................... Typosyllis alternata

I TyposyUis alternata (Moore, 1908)

Figure 1.32

I Syllis alternata Moore, 1908:323-325, figs. a-f; 1909:321.--Annenkova, 1938:148.--Berkeley and Berkeley,

1938:37-38; 1948:77-78, fig ll5.----_oja, 1941:691-692, pl.3, figs. 1-9.

I Syllis Typosyllis (Typosyllis) alternataalternata : Hartman, : Uschakov, 1948:21; 1968:479-480, 1955:180, fig. figs. 50, z-l.--Banse 1-5.--Imajima, and Hobson, 1966e:273-275, 1974:64,text-fig. fig. 1658a-1. a.--

Day, 1973:30.---Gardiner, 1976:141, fig. 13 b-c.

I Syllis cornuta Pettibone, 1954:253 (in part).

Material Examined. California: off Point Arguello, Sta. BRA-6 rock (3, USNM 170944) + (3,

I SBMNH 142709); off El Morro, Sta. BRA-27 rock (1, USNM 170945); between Point Estero and Point Buchon, Sta. 6, rocks (5, USNM 170943).--Alaska: Behm Canal, holotype (USNM 5542).

Description. Length up to 44 mm, with 125-160 segments (Moore, 1908; Imajima, 1966e); MMS

I specimens 10 mm long, 0.4 mm wide excluding parapodia, for 88 segments. Glandular tissue on body wall

present on every segment just posterior to dorsal cirri.

I farthest Prostomium apart; posterior wider2 pairs than long, set insome trapezoidal what pentagon arrangement; al; 3 pairsanterior of eyes,pair all lenticulate, as eyespots,middle eachpair present largest, on

anterior margin ofpmstomium, just medial to paired lateral antennae (Fig. 1.32A,B). Median antenna inserted

I between 2 posterior pairs of eyes, with up to 30 articles (14-19 in MMS specimens); lateral antennae inserted far forward on anterior margin of prostomium, each with 15-18 articles (9-12 in MMS specimens) (Fig.

1.32A,B). Palps broad, distally rounded, fused basally (Fig. 1.32A,B). Nuchal organs injuvenile specimen

i with 27 setigers small, paired, at posteetal corners of prostomium (Fig. 1.32B). Pharynx usually present in setigers 1-8 or to setigers 10-12, anterior middorsal tooth large, conspicuously visible through body wall in

setiger 3; with distal circlet of 10 soft papillae when everted. Proventficulus beginning from setigers 9-19 (9-

16 in MMS materials), depending on extension of pharynx, extending through 12 segments, with an average

I of 39 rows muscle cells (34-40 in MMS specimens).

Tentacular segment forming complete, distinct ring (Fig. 1.32A). Two pairs of tentacular cirri,

i resembling median antenna, prostomi withal15-23 antennae articles; in form, ventralshorter; pair eachdorsal aboutpairone-half longest, each about three-quarters length of length of dorsal pair, with 8-17 articles

(Fig. 1.32A,B).

I blades (Fig. Pampodia 1.32F-J), thicaround k, fleshy8-12lobes per fascicle, (Fig. 1.32C-E). thickest Setae in medial generally segments. compound Blades falcigers longest superiorly with bidentate (Fig.

1.32G), about 2 times longer than inferior blades in anterior segments decreasing to 1.5 in medial segments

I (Fig. cutting1.32F); edges. allDorsal decreasing and ventral i lengthsimpleinferiorly setae present within infasposterior cicles (Fig. segments, 1.32I,J);eachall with distallycoarsely bidentatesrrated with

serrated cutting surfaces (Fig. 1.32K-M). Up to 5 aciculae per anterior parapodium, decreasing to 1 in

I posterior parapodia; distally pointed, slightly curved distally, thickest emerging through parapodial lobes (Fig. 1.32N,O).

|
!

!

• !

!

!

E

!

!

jJ/, I

F G H I J K L M N 0 g

II

Figure 1.32. Typosytlis atternata: A, anterior segments, dorsal view; B, prostomium, dorsal view; C, left HI

parapodium, median segment, anterior view; D, left parapodium, median segment, dorsal cirri

omitted, anterior view; E, right parapodium, setiger 14, anteriorview; F, superior compound falciger, i

median segment; G-H, J, compound falcigers from setiger 14: G, superior seta; H, intermediate Hi

seta; J, inferior seta; I, inferior compound falciger, median segment; K-L, dorsal simple setae: K,

median segment; L, setiger 14; M, ventral simple seta, setiger 14; N-O, aciculae: N, median segment; mm

O,

setiger 14; P, pygidium, dorsal view. (A-P, originals by YDK) D
!

!

I Dorsal cirri strongly articulated, alternating tong and short, although longest overall in anterior 15 segments, decreasing gradually in medial and posterior segments. Dorsal cirri of setiger 1 longest, with 35

articles; setiger 2 with 25 articles; long ones in medial segments with 25, short ones with 18 articles. In

I specimens with about 60 segments, longest anterior dorsal cirri each with 25-31 articles, altemating with shorter ones having 17-23 articles; long ones of middle segments with 15-26, short ones with 8-21 articles. In

juvenile specimen measuring 1.3 mm long for 27 setigers, dorsal cirri with limited number of articles in all

I segments; those of setiger 1 with 8 articles; setiger 2 with 7; setigers 3-4 each with 6; setigers 5-12 with 5; setigers 13-14 with 4; setigers 15-22 with 3; setigers 23-26 with 2; setiger 27 with 1. Ventral cirri digitiform,

extending beyond parapodial lobes.

I Pygidium with paired anal cirri, each having up to 15-20 articles (Fig. 1.32P).

Remarks. The MMS specimens are generally consistent with the description ofTyposyllis alternata,

particularly in the pattern of alternating long and short dorsal cirri, although the dorsal cirri of the MMS

I tend to have numbers of articles than described Moore 1

specimens slightly greater originally by ( 908), they

also have fewer numbers of aciculae per parapodium, some of which project through the body wall; venlxal

cirri do not project beyond parapodia lobes; and the dorsal simple setae appear to be unidentate. Moore

I (1908) originally reported the ventral cirri to project beyond the parapodial lobes, although they do not in Japanese specimens (Imajima, 1966d).

i Habitat. DistributioMixed n. California; sediments,Washington; gravel, rockyBritish substrColumbia; ata, to depths Alaska; of 2560 Japan. m.

I Typosyllis bella Chamberlin, 1919

Figure 1.33

I Typosyllis bella Chamberlin, 1919:7.

I BRC-1, Ma rockteri(1,al USNM Examine170946). d. California: Western Santa Barbara Channel, east of Point Conception, Sta.

Description. Length about 20 mm, 1.25 mm wide in anterior region excluding parapodia, for about

I 145 segments (Chamberlin, 1919); MMS specimen incomplete, 5 mm long, 0.7 mm wide excluding parapodia, for 48 setigers. Color in life of mature female yellowish, each anterior segment with 2 transverse reddishbrown lines (Fig. 1.33B; Chamberlin, 1919); 2 transverse rust colored lines per segment through setigers 23-

24 in MMS specimen. Prostomium widest anteriorly; 2 pairs of eyes, elliptical; anterior pair largest; alltrapezoidally arranged

(Fig. 1.33A). Median antenna arising between anterior pair of eyes, with 21 distinct articles, nearly 2 times

I longer than prostomium; lateral antennae arising from anterofrontal margin of prostomium, about same length as median antenna, each with 15-18 articles (Fig. 1.33A). Palps nearly as long as prostomium, narrow,

distally rounded, not fused basally (Fig. 1.33A). Pharynx extending through setiger 13, with an anterior

I middorsal tooth. Proventriculus present in setigers 14-23; rows of muscle cells not counted.

Tentacular segment reduced. Two pairs of tentacular cirri, similar in form to median antenna; dorsal

pair longest, each longer than median antenna with 32-34 articles; ventral pair slightly less than one-half

I length of dorsal pair, each with about 15 articles.

!
1

I

A I

C I

I

B .01nm_

2_1

°

D E _ I

I

Figure 1.33. Typosyllis bella: A, pmstomium, dorsal view; B, median segments with paired transverse bars of

pigment per segment, dorsal view; C, right parapodium, setiger 2, anterior view; D, superior I

compoundfalciger,setiger 2;E, inferior compoundfalciger,setiger2; E,pairedaciculae,emerging

throughacicularlobe,superioracicula on fight. (A-E, originals by JDK)

I

Pampodia thick, fleshy, conical in dorsal view; distally incised, forming small superior and larger I

round inferior lobe (Fig. 1.33C). Setae compound bidentate falcigers (Fig. 1.338-E), 8-12 per anterior

fascicle (11-13 in MMS specimen), decreasing posteriorly. Blades longest superiorly (Fig. 1.33D), 2-2.5

times longer than inferior blades within fascicles (Fig. 1.33E), all decreasing gradually in length inferiorly I

within fascicles, and posteriorly along body; all with wide angle between the distal and subdistal teeth and I

finely serrated cutting edges (Fig. 1.33D,E). Setal shafts with numerous subdistal denticles, these gradually

decreasing in numbers inferiorly within fascicles (Fig. 1.33D,E). Two to 3 aciculae per parapodium (2 in •

MMS specimen), inferiormost usually large, pointed, superior ones smaller, distally blunt, middle one enlarged

in posterior segments; all projecting through parapodial lobe at and just below distal incision (Fig. 1.33F).

Dorsal cirri extremely well developed, longest in anterior segments (Fig. 1.33C), alternating long I

with 32 articles, and short with about 18 articles, and gradually decreasing in overall length posteriorly

I

(Chamber[in, 1919); those of MMS specimen generally lost, with dorsal cirri of setiger 1 longest, with 34

articles; setigers 2 and 4 each with 27; setigers 3 and 5 each with 16-17. Ventral cirri slender, digitiform, I

extendinbgeyondparapodiallobes(Fig.1.33C).

unknown. I

I
!

!

Remarks. The present specimen is remarkably consistent with Chamberlin's (I919) original

I description, especially for the prostomium, dorsal cirri and dorsal segmental pigmentation pattern. The holotype of this species is apparently deposited in the Museum of Comparative Zoology, Harvard, and has not been

illustrated or redescribed since first reported, although Hartman (unpublished) did examine the holotype in

I 1937. The MMS specimen is tentatively referred to Typosyllis bella pending re-examination of the holotype.

Habitat. Originally reported from the low intertidal zone of Laguna Beach (Chamberlin, 1919); also

associated with rocky substrata in depths of 73.5-78 m.

I Distribution. Southern California.

I TyposyUis hyalina (Grube, 1863)

Figure 1.34

I Syllis (Typosyllis) hyalina Fauvel, 1923:262-263, fig. 98a-c.--Rullier, 1964:159-160.

SyUis hyalina Grube, 1863:45.--Ehlers, 1901:86.--Monro, 1933:30.--Berkeley and Berkeley, 1948: 74,

i fiWestheide, gs. 107-108.---_ 1974:47-51, oja, 1941:692.-- pl. 20.--Ben-Eliahu, Banse and1977:9-10.--Nfifiez, Hobson, 1974:62, San fig. Mart 16m.--Reish, 1968:214.-- fn and Brito, 1992:120.

TyposyUis hyalina: Imajima, 1966e: 271-273, text-fig. 57, a-k.--Hartman, 1968:487-488, figs. 1-3.---Gardiner,

i TyposyU1976:140, fig. 12v-w.--Tebble, 1959:17.--Day, 1967:246, fig.12.2.v-x; 1973:29. is (TyposyUis) hyalina Hartmann-Schr6der, 1965:95; 1979:89, figs. 57-61; 1980:50.

Typosyllis aciculata orientalis Imajima and Hartman, 1964:130-132, pl. 3, figs. e,f, pl. 32, figs. a-t. Fide

i Westheide, 1974.

Material Examined. California: Santa Maria Basin, Sta. BRA-21 (1, SBMNH 142710).--Western

Santa Barbara Channel, east of Point Conception, Sta. BRC-I (7, USNM 170947) + (1, USNM 170948);

I BRC-2 (5); BCC-1 (1).

Description. Body 10-35 mm long, 2 mm wide including parapodia; with over 100 segments (Imajima,

i • 1966e; Hartman, 1968). Color in life variable: translucent, white or occasionally with dorsal pattern of

II broken or continuous transverse dark lines alternating on every 3 anterior segments (Fig. 1.34A).

Prostomium wider than long, subglobular; 2 pairs of black eyes, setin trapezoidal arrangement, plus

I anbeyond additional palps;pair median of froantenna ntal eyespots arisingbetween from center basesofofprostomium, frontal antennae with(Fig. 13-141.34A). articles;Antennae lateral antennae long, extenarising ding

from anterior margin of prostomium, each with 10-12 articles (Fig. 1.34A). Palps large, thick, subtfiangular,

I anterior 1.5 timesmiddorsal longer thantoothprostomium, conspicuous;fusedwithbasally a distal(Fig. circlet 1.34A). of 10Everted soft papillae. pharynxProventriculus extending to present setiger 9, in

segments 10-21.

I Tentacular cirri longer than antennae; dorsal pair each with 20 articles; ventral pair each about as long as median antenna (Fig. 1.34A).

Parapodia fleshy, bluntly conical (Fig. 1.34B,C,H). Setae generally compound falcigers (Fig. 1.34DI G), numbering around 10 per fascicle. Blades distally bidentate. Superior and inferior blades within fascicles subequal in length (Fig. 1.34D-G); those in anterior segments long, slender, conspicuously serrated (Fig.

1.34D,E); in median and posterior segments short, stout, finely serrated (Fig. 1.34F,G). A dorsal superior

I and a ventral inferior simple seta present in posterior fascicles, both distally bidentate with serrated cutting surfaces (Fig. 1.34I,J). Aciculae numbering 4 per anterior parapodium, decreasing to 1 posteriorly, each

distally resembling bird's beak (Fig. 1.34K).

!
I

I

i

I

A B li

f' t

!

I

I

I

!

I

D E F G I J K |

|

Figure 1.34. Typosyllis hyatina: A, anterior end, dorsal view; B, median parapodium with short dorsal cirrus, u

posterior view; C, median parapodium with long dorsal cirrus, posterior view; D, superior compound

falciger, parapodium 10; E, inferior compound falciger, parapodium 10; F, compound falciger, OR

median parapodium; G, compound falciger, posterior parapodium; H, posterior parapodium; I, II

dorsal simple seta, posterior parapodium; J, ventral simple seta from same parapodium as I; K,

aciculae from median parapodium. (A-K, redrawn from Imajima, 1966e) n

i

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88 il
I

I Dorsal cirri thickest basally, varying in length, strongly annulated; those of setiger 1 longest, with 20

annulations; setigers 2, 3, 5 short, each with 11-14 armulations; setigers 4, 6 long, with 16 annulations;

i setigers (setiger 7, 11,8 sh14oannulations) rt; setiger 9 lotong;endthereafter of bodyregularly (Fig. 1.34A). alternating Ventral shocirri rt (setiger digitiform, 10, 11not annuextending lations)andbeyond long

parapodial lobes (Fig. 1.34B,C,H).

I Pacific andRemarks Atlantic . Typosyllis basins andhyalina the Medite is a rwell-recognized, ranean Sea. Local widespread populationsspecies of bothreported T. hyaIina from and bothT.thepulchra north

were studied extensively by Piltz (1980).

I MMS Habitat. Associated with algae, poriferans and mussel beds in intertidal zones, and hard substrata at stations in depths of 69-90 m.

Distribution. Panama; California north to British Columbia; Japan; North Atlantic Ocean;

I Mediterranean Sea.

I Acknowledgements

I are particularly to editors, Brigitte Hilbig during preparation

We indebted James A. Blake and of this

chapter, and for their critical editorial reviews; Kristian Fauchald and Linda Ward (USNM) for providing

access to and copying literature for both of us, for allowing LHH access to their collections, and for valuable

I Kirk for his careful editorial loans of and for LHH discussions; Fitzhugh (LACM) review, syllids, allowing to

co-author this chapter; both Marian Pettibone (USNM) and David H. Russell (Dept. of Biology, Washington

College, Maryland) for their editorial reviews; and John H. Dorsey (City of Los Angeles, Environmental

I Monitoring Division) for use and modification of his original illustrations, and early discussions with and

encouragement to LHH. Ardis Johnson (MCZ) arranged loans of type materials to LHH; and David George

and Alex Muir (BMNH), Pat Hutchings (AMW), Mary Petersen (Zoological Museum, University of

I Copenhagen), Gesa Hartmann-Schr&ier (ZMH), Paul Scott (SBMNH), and Robert VanSyoc (CASIZ) provided

access to their collections. Charles Mitchell (MBC Applied Environmental Sciences, Inc.) encouraged LHH

in her early efforts to work on syllids collected by MBC; Kinnetic Laboratories, Inc. also provided specimens.

I Assistance given to LHH by Ron Velarde, Larry Lovell, Tony Phillips and other SCAMIT members is also gratefully acknowledged.

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Curiosum, Halle 20:1-264.

I Reish, D. J. 1968. The polychaetous annelids of the Marshall Islands. Pacific Science 22(2):208-231.

i Rioja, E. 1925. Anelidos poliquetos de San Vicente de la Barquera (Cantabrico). Trabajos del Museum National Ciencias de Naturales 53:1-62.
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Rioja, E. 1941. Estudios annelidologicos II. Datos para el conocimiento de la fauna de Poliquetos de las I

costas del Pacifico de Mexico. Anales del Instituto Biologia de la Universidad Nacional de Mexico

12: 669-746. I

Rioja, E. 1943. Estudios annelidologicos VII. Aportaciones al conocimiento de los Exogoninos (Anelidos

Poliquetos) de las costas Mexicanas del Pacifico. Anales del Instituto Biologia de la Universidad i

Nacional de Mexico 14: 207-227. H

Riser, N.W. 1991. An evaluation of the taxonomic characters in the genus SphaerosyIlis (Polychaeta: Syllidae).

Proceedings of the Second Ilatemational Polychaete Conference, Copenhagen, Denmark, August, •

1986. Systematics, biology and morphology of world Polychaeta. Ophelia, Supplement 5:209-217. II

Rullier, E 1964. Rfsultats Scientifiques des Campagnes de laCalypso. Annales de l'Institut Oc6anographique i

du Monaco 6:113-218, figs. 1-23. i

i

Russell, D.E. 1987. The taxonomy and distribution of Syllidae (Annelida: Polychaeta) inhabiting mangrove

and adjacent shallow-water habitats of Twin Cays, Belize. Doctoral Dissertation. The George m

Washington University, Washington, D.C. 388 pp. II

Russell, D.E. 1989a. Three new species of Sphaerosyllis (Polychaeta, Syllidae) from mangrove habitats in

Belize. Zoologica Scripta 18(3):375-380. i

i

Russell, D.E. 1989b. A new species ofOdontosyllis (Polychaeta: Syllidae) from Twin Cays, Belize. Proceedings

of the Biological Society of Washington 102(3) :768-771. B

Russell, D.E. 1991. Exogoninae (Polychaeta: Syllidae) from the Belizean barrier reef with a key to the species

of Sphaerosyllis. Journal of Natural History, London 25:49-74. fit

San Martin, G. 1982. Estudio biogeogrfi:fico, fannfstico y systerr_tico de los Poliquetos de la familia Sflidos B

(Polychaeta: Syllidae) en Baleares. Tesis Doctoral. Universidad Complutense de Madrid. 529 pp.

San Martin, G. 1984a. Estudio biogeogrfifico, faunfstico y systerlfitico de los Poliquetos de la familia Sflidos B

(Polychaeta: Syllidae) en Baleares. Tesis Doctoral. Publicaciones de la Universidad Complutense de m

Madrid. 529 pp.

i

San Martin, G. 1984b. Descripci6n de una nueva especie y revisi6n del g6nero Sphaemsyllis (Polychaeta: i

Syllidae). Cahiers de Biologie Marine 15:375-391.

San Martfn, G. 1990. Eusyllinae (Syllidae, Polychaeta) from Cuba and the Gulf of Mexico. Bultetin of &

Marine Science 46:590-619. J

San Martin, G. 1991a. Sphaerosyllis and Parapionosyllis from Cuba and Florida. In: Petersen, M.E. and J.B. •

Kirkegaard (eds.). Proceedings of the Second International Polychaete Conference, Copenhagen, I

Denmark, August, 1986. Systematics, biology and morphology of world Polychaeta. Ophelia,

Supplement 5:231-238. i

San Martfn, G. 1991b. Grubeosyllis andExogone (Exogoninae, Syllidae, Polychaeta) from Cuba, the Gulf of

Mexico, Florida and Puerto Rico, with a revision of Exogone. Bulletin of Marine Science 49:715-

740. i

Sardfi,R. 1984. La subfamilia Exogoninae (Polychaeta: Syllidae) de Gibraltar, con descripci6n dePseudobrania

euritmica n. sp. Publicacion Departmento Zoology de Barcelona 10:7-13. •

i Schroeder, P.C. and C.O. Hermans. 1975. Armelida: Polychaeta. In: Giese, A.C. and J.S. Pearse (eds.)

Reproduction of Marine Invertebrates. Vol. HI. Academic Press, New York. pp. 1-213.

I
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I Southern, R. 1914. Archiarmelida and Polychaeta. Proceedings of the Royal Irish Academy of Dublin 31(47): 10-

60 (Clare Island Survey).

I Tebble, N. Research 1959. OnCouncil a collection of polychaetes from the Mediterranean coast of Israel. The Bulletin of the of Israel B8:9-30.

i Treadwell, A.L. 1914. Polychaetous annelids of the Pacific Coast in collections of the Zoological Museum of the University of California. Univeristy of California Publications in Zoology 13:175-234.

Uebelacker, J.M. 1984. Family Syllidae Grube, 1850. Chapter 30 In: Uebelacker, J.M. and P.G. Johnson

I (eds.), Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Vot.IV. Barry A. Vittor

' and Associates, Inc. Mobile, Alabama.

t Uschakov, P.V. 1950. [Polychaeta of the Sea of Okhotsk]. Akademiya Nauk SSR, Issled Dal'nevost Morei SSSR 2:140-237. [In Russian.]

Uschakov, P.V. 1955. Polychaeta of the far eastern seas of the USSR. Akademiya Nauk SSR, Opredeliteli po

Faune Translations, SSR 56:1-445 for the National [In Russian; Sciencetranslated Foundation, in 1965419bypp.]Jerusalem, Israel Program for Scientific

i Vogt, K.D. and J.D. Kudenov. 1994. Morphometric variation in bifurcate notosetae of two Euphrosine species (Polychaeta: Euphrosinidae). In: Dauvin, J.C., Laubier, L. and D.J. Reish (eds.) Actes de la

4_me Conf6rence intemationale desPolych_tes. M6moires de Mus6um National D'Histoire Naturelle

i (A) 162: 291-298.

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I 2. FAMILYAPHRODITIDAE]VIALMGREN, 1867

! uy

James A. Blake 1

I Introduction

i The aphroditids are stow-moving scaleworms commonly called sea mice because of a feltlike dorsal

surface that is formed by fine, silky setae and entrapped particles. Aphroditids, because of their size, do not

I occur in dense populations. They are adapted to burrowing in mud or creeping over the bottom, where they feed on detritus and on sessile or slow moving animals, such as polychaetes (Day, 1967; Fauchald and Jumars,

1979).

!

Morphology

!

The body of aphroditids is large, oval, and has relatively few segments (usually less than 60) (Fig.

2.1A, B). Typically, the ventral side is flattened, while the dorsum is arched. The prostomium is spherical

I and bears a single medial antenna and a pair of long palps (Fig. 2.1D). A well-developed facial tubercle is

present above and anterior to the mouth (Fig. 2.1D). Jaws are lacking or poorly developed, but the pharynx

is muscular and eversible with several rows ofpapiUae around the opening. Internally, the alimentary canal

I extends between the form The first bears two of

out septa to segmentally arranged caeca. segment pairs

tentacular cirri and is uniramous with numerous capillary setae. Parapodia of subsequent segments are

biramous, with large internal aciculae. Notosetae include both t'me silky setae which entrap silt particles and

I form the characteristic dorsal felt and stout spines, some of which be harpoon-shaped (as inLaetmonice)

may

(Fig. 2.1C). Neurosetae are simple and sometimes forked. The dorsal surface is covered with 15-20 pairs of

elytra (overlapping scales) which occur on segments 2, 4, 5, 7, and alternate segments to 25, then on every

I third segment. Long dorsal cirri are present on segments which lack elytra (Fig. 2.1E). The pygidium is very small and lacks anal cirri.

i Biology

I Aphroditids occur from the shallow subtidal to the deep sea. They usually inhabit sandy or muddy

bottoms where they are adapted to burrowing or creeping over the surface.

I Mettam The (1980) y havobserved e a large, A, maculeata uscular eversible Linnaeus pharynx feeding onorNephtys proboscisandandNereis are .thoSpecimens ught to be acti ofNereis ve carnivore varianss.

about three times longer than theAphrodita were swallowed head first indicating an active predatory behavior.

I Laetmonice filled with parts speciesof may the brittlestar, also be actiOphiomusium ve predators, bec sp.aus in eALn.tarctic productawaters Grube(Hartman, has been 1967; observedJ.H.wDearborn, ith its gut

personal communication). According to Pettibone (1953), aphroditids eat a great variety of prey items including

I other annelids, small crustaceans, hydroids, sponges, small molluscs, holothurians, and diatoms. Other animals that use the bodies of aphroditids for support andrefuge include other annelids, protozoans, sponges, hydroids,

I *ENSRConsultingandEngineering,89 WaterStreet,WoodsHole,Massachusett0s2543

I 99
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I

I

A C

i

I

pa

noS I mAn dC

".- . . .

felt ' """ " "':: !

dC

ne I

D E

no vC I

el/

Figure 2.1. kphroditidae: A, Aphrodita, entire animal, dorsal view; B, Laetmonice, entire animal, dorsal view; I

C, harpoon seta; D, Aphrodita, anterior end, dorsal view, felt removed; E, parapodium of cirrigerous

segment, anterior view. (A-B, after Mclntosh; C, after Hartman; D, after Fordham; E, afterPettibone). I

100 I
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I echinoderms, bryozoans, molluscs, crustaceans, and tunicates. One particularly interesting association is

with the bivalve, Pseudopythina rugifera (Carpenter) that lives attached to the ventral surface of Aphrodita

i California refulgida Moore. This worm and its commensal clam were collected from off Tomales Point in northern and studied in the laboratory by Narchi (1969). This small clam attaches to the worm with its

byssal threads. Its digestive morphology is well adapted to handling the fine silt particles found inthe worm's

i habitat (Narchi, Little is known 1969). about the reproduction and development of aphroditids. Paired nephridial orsegmental

papillae on the posterior face of parapodia between rami serve for the passage of genital as well as excretory

I products (Pettibone, 1953).

i Taxonomic History

In general, there has never been acomprehensive review of the aphroditid polychaetes. Major works

i by Horst (1916a-b, 1917) on the aphroditids of the Siboga Expedition remained for many years the only summaries of this family. Pettibone (1966) redefined some genera, described one new genus, and provided an

important key. Some eastern Pacific species were reviewed by Hartman (1939) and Pettibone (1953) and the

I aphroditids from California were summarized and keyed out by Hartman (1968). Few of these species, however, including most described by Moore (1903, 1905, 1910), have ever been redescribed since their

original account. More recently, Watson Russell (1989) revised the genus Palmyra and transferred it from the

I Chrysopetalidae to the Aphroditidae and Hutchings and McRae (1993) published a monograph on the aphroditids of Australia and reviewed the species collected as part of the Siboga Expedition. The Hutchings and McRae

effort included definitions of five genera and 34 species, seven of which were new to science. This paper is

I also an important resource and guide to aphroditid morphology and systematic characters.

I Key to the Genera of Aphroditidae from California

1A. Dorsal felt well-developed, completely covering etytra with thick mat (Fig. 2.1A); harpoon-shaped

i notosetae lacking; neurosetae pointed or slightly curved, or fringe

with without lateral of soft bristles

or basal spur (Fig. 2.2B, D) .................................................................................... Genus Aphrodita

t lB. Dorsal felt poorly developed, usually not covering elytra (Fig. 2.1B); hartx_on-shaped notosetae present

(Fig. 2.1C); neurosetae with lateral fringe of long, stiff bristles and basal spur ................................

.............................................................................................................................. Genus Laetmonice

I Descriptions of Species

i Genus Aphrodita Linnaeus, 1758

Type Species: Aphrodita aculeata Linnaeus, 1758.

I Diagnosis. Body oval, arched dorsally and tapered posteriorly, with 25-45 segments. Prostomium

with a single medial antenna and a pair of large sessile eyes. Dorsum covered with tough felt; ventrum usually

with numerous small papillae. Dorsum with 15 pairs of elytra under felt. Notosetae including stout acicular I

i 101
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spines and long, fme capillaries which form felt; harpoon setae absent. Neurosetae very stout, acicular, arranged Illl

in 3 tiers, n

,u

Remarks. Eight species were recorded from California by Hartman (1968) and readers should refer

to her work for keys to species. Earlier, Pettibone (1953) redescribedA, japonica Marenzeller (1879) and A. flU

negligens Moore (1905) from the Puget Sound. As part of that work, she included A. refulgida Moore (1910) I

in the synonymy of A. japonica andA. castanea Moore (1910) andA. brevitentaculata Essenberg (1917) in

the synonynmy ofA. negligens, ff followed, these synonymies would reduce Hartman's list of species present II

in California from 11 to 5. Regardless of how many species are actually in the California fauna, all occur on II

muddy bottoms and should be more commonly taken in trawls than in grabs and box cores. One species,

i

Aphrodita parva Moore has been collected in the Santa Mafia Basin. n

i

Aphrodita parva Moore, 1905

Figure 2.2 I

Aphrodita parva Moore, 1905:529-532, pl. 34, figs. 3-7.--Moore, 1908:339; 1910:385-386.--Treadwell, il

1914:178.--Berkeley, 1923:211.--Haxtman, 1939:22, pl. 1, figs. 9-10; 1968:27, figs. 1-5. i

Material Examined. Santa Mafia Basin, off Point Estero, Sta. 2, 200 m (1 juvenile); off Point Sal, I[

Sta. PJ-1 (1); offPoint Arguello, 900 m (1). II

Description. A relatively small species, body short and broad, up to 24 mm long, 15 mm wide for 27 Ill

segments. Color in alcohol: flesh. Dorsum covered with greenish silt and debris; ventrum covered with In

numerous small, spherical papillae. I

Prostomium wider than long, bearing 2 pairs small eyes on short frontal peaks; medial antenna slender,

about 2.5 times as long as prostomium (Fig. 2.2A); palps thick, about twice as long as medial antennae, i[

Elytra numbering 14-15 pairs, each smooth, thin, transparent without papillae. Parapodia prominent with

mR

elevated notopodium and elongated neuropodium (Fig. 2.2C); dorsal cirrus long, threadlike; ventral cirrus ill

thin, tapering, about as long as neuropodium. Notosetae including heavy bristled copper-colored glossy 1

spines and thin pointed setae (Fig. 2.2D, E); neurosetae in 3 rows: dorsalmost row with 1-2 thick spines,

middle row with 2-3 spines, and lower row with 3-4 spines; setae diminishing in size ventrally, with lowermost /

setae bearing prominent spurs (Fig. 2.2B). Hi i

Biology. Inhabits bottoms with fine silt.

Remarks. The specimen from Sta. 2 had been identified as Pontogenia sp. A in the Phase I

reconnaissance. However, the specimen is clearly a juvenile Aphrodita, probably A. parva. Although A. 1

parva is the name applied to these specimens here, it is possible that the species itself is actually a juvenile

form of A. japonica.

Type Locality. British Columbia, Halibut Bank, Gulf of Georgia, 203-331 m (syntypes, ANSP I_

2061; USNM 4194).

Distribution. Alaska to western Mexico, 6-1220 m. _l

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II

B ,,c C

I

18 D. E

i Figure 2.2. Aphrodita parva: A, anterior end, dorsal view; B, three types of neurosetae with spurs; C, right parapodium from segment 10; D, bearded neuroseta; E, notoseta. (after Moore, 1905).

I Genus Laetmonice Kinberg, 1855

i Type Species: Laetmonicefilicornis Kinberg, 1855.

Diagnosis. Body oval, depressed, with 32-46 segments. Prostomium with papillose facial tubercle;

I rounded with single medial antenna and 2 stout ommatophores. Feltage poorly developed, not completely covering elytra; elytra smooth, coveting dorsum, numbering 15-20 pairs. Notosetae of segments with elytra

including harpoon-shaped setae, acicular setae, and capillaries; notosetae of cirrigerous segments without

i harpoon setae, but silky capillaries may be present; neurosetae bifurcate with fringe bristles on edge.

Remarks. Two species, Laetmonice pellucida Moore and L. producta wyvillei McIntosh have been

recorded from off California in deep water (Hartman, 1968). None have been encountered in the Santa Mafia

i Basin and Santa Barbara Channel.

western

i Literature Cited

i Berkeley,Contributions E. 1923. Poltoychaet Biology, ous anneli new series, ds from1:203-218, the Nanaimo1 pl.District. 1. Syllidae to Sigalionidae. Canadian

Day, J.H. 1967. Polychaeta of Southern Africa. Part I, Errantia. Publication of the British Museum of

i Natural History 656:1-458.

i 103
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Fauchald, K. and EA. Jumars. 1979. The diet of worms: A study of polychaete feeding guilds. Oceanography m

and Marine Biology Annual Review 17:193-284.

Hartman, O. 1939. Polychaetous annelids. PartI. Aphroditidae to Pisionidae. Allan Hancock Pacific Expedition n 

7:1-156, 28 pls. i

Hartman, O. 1967. Polychaetous annelids collected by the USNS Ettanin and Staten Island cruises, chiefly

from Antarctic seas. Allan Hancock Monographs in Marine Biology 2:1-387. H

II

Hartman, O. 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation,

University of Southern California Press, Los Angeles. 828 pp. il

Horst, R. 1916a. On a genus of Aphroditidae from the Netherlands East Indies. Zoo]ogische Medeleelingen- n

Deei II. Leiden 2:63-64.

Horst, R. 1916b. Malayan species of the genera AphroditeUa, Hermione, Laetmonice, and Aprogenia. I

Zoologische Medeleelingen-Deei IT.Leiden 2:65-77.

w

Horst, R. 1917. Polychaeta Errantia of the Siboga-Expedition. Pt. 2. Aphroditidae and Chrysopetalidae. •

Siboga-Expedition Leydon 24b: 1-140. II

Hutchings, P.and J. McRae. 1993. The Aphroditidae (Polychaeta) from Australia, together with a redescription i

of the Aphroditidae collected during theSiboga Expedition. Records of the Australian Museum 45:279- i

363. i

Mettam, C. 1980. On the feeding habits of Aphrodita aculeata and commensal polynoids. Journal of the

Marine Biological Association of the United Kingdom 60:833-834. n

Marenzeller, E. 1879. Stidjapanische Anneliden, I. Denkschriften der Akademie der Wissenschaften Wien

41(2):109-154, 6 plates, i

Moore, J.P. 1903. Polychaeta from the coastal slope ofJapan and from Kamchatka and Bering Sea. Proceedings

of the Academy of Natural Sciences of Philadelphia 55:401-490, pls. 23-27. i

II

Moore, J.E 1905. New species of polychaetes fromthe North Pacific, chiefly from Alaskan waters. Proceedings

of the Academy of Natural Sciences of Philadelphia 57:525-554. i

Moore, J.P. 1908. Some polychaetous annelids of the northern Pacific coast of North America. Proceedings U

of the Academy of Natural Sciences of Philadelphia 60:321-364.

Moore, J.P. 1910. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern H

California in 1904. Polynoidae, Aphroditidae and Segalionidae. Proceedings of the Academy of u

Natural Sciences of Philadelphia 62:328-402, pls. 28-33. I

Narchi, W. 1969. On Pseudopythina rugifera (Carpenter, 1864) (Bivalvia). The Veliger 12(1):43-52. i

Pettibone, M.H. 1953. Some scale-bearing polychaetes of Puget Sound and adjacent waters. University of i

Washington Press, Seattle. 89 pp. + 40 pls. H

Pettibone, M.H. 1966. Heteroaphrodita altoni, a new genus and species of polychaete worm (Polychaeta,

Aphroditidae) from deep water off Oregon, and a revision of the aphroditid genera. Proceedings of i

the Biological Society of Washington 79:95-108. l

Treadwell, A.L. 1914. Polychaetous annelids of the Pacific coast in the collection of the zoological museum

of the University of California. University of California Publications in Zoology 13:175-234, 2 pls. i

m

Watson Russell, C. 1989. Revision of Palmyra Savigny (Polychaeta: Aphroditidae) and redescription of

Palmyra aurifera. The Beagle. Records of the Northern Territory Museum of Arts and Sciences i

6:35-53. i

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I 3. FAMILYPOLYNOIDAE MALMGREN, 1867

! by

I R. Eugene Ruff 1

I Introduction

i The family Polynoidae is the largest and most commonly encountered group of scaleworms. The

approximately 700 species in this family are characterized by dorsoventrally flattened bodies, simple setae in

both notopodial and neuropodial fascicles, and scales alternating with the dorsal cirri down much of the length

of the body. Although a few species become quite large (up to 250 mm), the majority of the scaleworms are

I only a few centimeters in length.

I Morphology

I In most polynoid species the prostomium is bilobed, with a median furrow between the anterior lobes;

the anterolateral corners are sometimes more or less developed into distinct cephalic peaks, or they extend

Ill anteriorly to form the ceratophores of the lateral antennae (Fig. 3.1A). There are typically two pairs of eyes

i arranged in a trapezoidal pattern, although the eyes in deep-water species may be absent. Most species have

a median and a pair of lateral antennae which are smooth or covered to a lesser or greater extent with papillae.

I Aantennae; pair of tapering these structures palps areusually attachedhaveventrally numerous to thelongitudinal prostomium,rowsandofareminute normally sensorythicker papillae. and longer The eversible than the

pharynx is large and muscular, with two pairs of curved, dark, keratinous jaws surrounded by a circlet of

i marginalThe papillae. tentacular segment (segment 1) has two pairs of tentacular cirri supported on large, forwardprojecting basal lobes. These tentaculophores have an internal supporting acicula, and sometimes on the

I anoftthe eriorperistomium face there areforms additional the upperrojecting lip of thesetaemouth that are (Fig.usu3.1allyB).similar This istooften the notosetae. produced The intoventral a ridgeportion whch

sometimes bears a distinct conical facial tubercle.

I be developed The buccal into a nuchal segmentfold(segthat ment partly 2) bears coversthethefirst prostomium, pair of elytraandandventrally the f'n'stitparapodia. forms the lateral Dorsaland ly itlower may

portions of the mouth. The ventral buccal cirri on this segment are usually well-developed and inserted at the

I bases of the parapodia. The paired elytra are flattened, scale-like structures that occur in place of the dorsal cirri, and are

normally attached via the elytrophores to segments 2, 4, 5, 7, 9.... 21, 23; posterior to this point there are a

I number of different attachment patterns, and the scales may be lacking in the posteriormost segments. The elytra may overlap and completely conceal the dorsum, or they may be reduced in size. The surface of the

scales may be smooth, orit may be covered to a greater or lesser extent with papillae, microtubercles (sclerotized

I structures that are nodular, pointed, or multipronged, and that are clearly visible only under high magnification)

i 1RuffSystematics,11719MeridianEast,Suite401,Puyallup,WA98373.

i 105
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Harmothoinae

_c |

cph

nuF I

_ A

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Lepidonotinae

-o |

tC' cer

_c |

,c |

Arctonoinae I

m

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c I}

_c B |

, II

Figure 3,1. Pofolynoid morphology: A, anterior end of typical harmothoid polynoid, dorsal view; B, ventral view I

same; C, prostomia of representatives of three polynoid subfamilies showing insertion pattern of

lateral antennae.

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I or macrotubercles (larger, sclerotized or soft structures that occur irregularly on the surface or near the

posterior edges). The borders of the elytra may be smooth, or they may have sparse or dense fringes of clavate

i or filiformThepapil elongated lae. parapodia are biramous or, in some cases, subbiramous. The notopodia are usually

located along the dorsal margin of the neuropodia; each has an internal supporting acicula which may be

I postsetal distally emergent. lobe and aThelonger, nuropodia narrowerarepresetal usualylobe largerbearing than thethenotopodia, intemal acicula and arewhich distallymaycleftmerge into adistally. rounded

All polynoid setae are simple. Although lacking in a few species, the notosetae range from slender

I orandwithout smoothatoterminal stout with cleft.subdistal The neurosetae transversenormally spinoushave plates.a long Thesmooth tips mayshaft be capillary, and a curved, pointed, subdistal or blunt inflated with

spinous region; the setal tips may be capillary, unidentate, or bidentate with a subequal or small secondary

I tooth. and bidenThe shape of the superior neurosetae is often different from those lower in the fascicle, and both uni- tate tips are sometimes found within the same setal bundle.

Dorsal cirri are inserted along the upper margin of the notopodia on segments not beating elytra; in

I addition, these segments have a more or less developed dorsal tubercle corresponding in position to the etytmphore. Ventral cirri are normally inserted midway along the ventral edge of the neuropodia after segment

2. Small, cylindrical nephridial papillae occur ventrally at the base of the neuropodia, usually from segment

I 6; these structures project posteriorly and upward between the parapodia. The pygidium surrounds a dorsally- directed anus, and has a pair of terminal anal cirri that are similar in shape but often longer than the dorsal

cirri.

!

Taxonomic History

i

For many years, the polynoids were divided into the Iphioninae, Lepidonotinae, and Harmothoinae

I based 20 years, essentially however,on the number and arrangement of the prostomial appendages. Discoveries within the last including a wealth of material obtained from the deep-sea hydrothermal vents, have prompted

the erection of approximately 15additional subfamilies. Dr. Marian H. Pettibone of the Srnithsonian Institution

I has clarifybeentheinstrumental relationshipsin revising within thethefamily. systematics Therefore, o the poly thensubfamilial oids, and isand contingeneric uing her position revisionary of manystudofiestheto

species is currently in a state of flux, a fact reflected in some of the less familiar generic designations listed

I below. The insertion of the lateral antennae is of primary importance indistinguishing some of the subfamilies

within the Polynoidae, including the four subfamilies represented in the Santa Maria Basin material (Fig.

i 3. ventrally 1C). Inbeneath the Harmothoin the anterior ae (Horst, pmstomial 1917), margins the later anda/lorantennae to the large haveceratophore small ceratophores of the median that areantenna. attachedIn

the Lepidonotinae (Willey, 1902) and closely allied Lepidastheniinae (Pettibone, 1989), the lateral antennae

I are Arctonoinae attached terminal (Hanley,ly1989), to anterior the lateral prolong antennae ations have of thelargeprostomium, ceratophoreswithout that aredistinct insertedceratophores. subterminally,In and the

are distinctly separated from the prostornium by a transverse groove.

I The number and arrangement of the elytra are very important in distinguishing the polynoid genera. Even though the scales are often autotomous, their position can be assessed by counting the distinctive

elytrophores along the body. Unfortunately, many species fragment during preservation, and the posterior

I powortion of the body is often lost and not available for examination. Therefore, it is best to narcotize scale rms in a solution of magnesium chloride in seawater immediately after collection. The worms should then

be preserved separately if possible to ensure that the elytra are maintained with the correct specimen.

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Distribution and Biological Notes I

a

Polynoids are found from the intertidal regions to the abyssal depths on a wide variety of sediment II

types, although a few are entirely pelagic. Most species are carnivorous or omnivorous, feeding on a large

u

spectrum of smaller invertebrates, plant fragments, and detritus. These species normally creep along the ,,

bottom, hiding in crevices, under rocks, and in algal holdfasts. The dorsum and the elytra areoften pigmented

with a variety of patterns and colors matching the general background. In addition, the elytral surface is

sometimes covered with detritus and epiphytes, making the specimens difficult to detect.

A number of polynoids are commensal with other organisms, predominantly the echinoderms, molluscs, !

or other polychaetes. In many of these species, the elytm and notopodia are reduced in size, and the notosetae

are fewer in number or absent altogether. Many of these commensals are pigmented similar to the host

organism. I

All polynoids are dioecious, with fertilization taking place externally. The eggs may be discharged

directly into the water column, but in many species they are brooded beneath the elytra and released into the Ill

seawater only after some degree of development. Blake (1975a) illustrates the pelagic larvae of Halosydna II

brevisetosa from Tomales Bay, while other authors (Thorson, 1946; Cazaux, 1968; Daly, 1972) have delineated

the reproductive biology and larval development of the widely distributed species Harmothoe imbricata, l

Typically, a larval nectochaete metamorphoses to a bottom dwelling juvenile when it has developed 5-6 pairs |

of elytm. During metamorphosis the prostomium gains an adult aspect, and the first larval setiger shifts

forward to develop into the tentacular segment. In most free-living polynoids, the number of segments is m

determinant within a small range, and the worms do not grow beyond 30-40 mm in length. A number of the |

commensal species, however, continue to add segments throughout their lives, and much greater body lengths

are attained, i

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I Key to the Species of Polynoidae

I 1A. Lateral antennae attached terminally to anterior prolongations of the prostomium, without distinct

ceratophores ..................................................................................................................................... 2

I lB. Later prostomium al antennae .................................................................................................................................... attached subterminally or ventrally, with ceratophores distinctly demarcated from10

I elytra marginal fringes papillae; most notosetae ending long,

2A. Anterior with surface tubercles and of in

fine tips ............................................................................................................................................ 3

i 2B. Surfaces and margins of anterior elytra smooth; all notosetae (when present) ending in blunt unidentate or notched tips .................................................................................................................................. 7

I 3A. Twelve pairs of elytra with long fringing papillae; numerous notosetae in large mils ..................... 4

3B. Eighteen pairs of elytra with short fringing papillae; relatively few notosetae in small bundles ...... 5

I 4A. Basal plates of larger tubercles on anterior elytra polygonal; largest tubercles on posterior elytra

acutely conical, pointed .................................................................................... Lepidonotus spiculus

I 4A. Basal plates of larger tubercles on anterior elytra rounded; largest tubercles on posterior elytra digitiform, blunt-tipped ............................................................................... Lepidonotus squamatus 2

I 5A. Surfaces of median and posterior elytra smooth; some neurosetae with bidentate tips ......................

.......................................................................................................................... Halosydna johnsom "2

I 5B. Elytral surfaces with chitinized conical tubercles; all neurosetal tips unidentate ............................. 6

6A. Body broad, oval in outline; conical macrotubercles restricted to region behind attachment scar .....

I Halosydna latior 2

6B. Body long, linear; conical macrotubercles dispersed across elytral surface .......................................

i ..................................................................................................................... Halosydna brevisetosa 2

7A. Superior neurosetae thinner than median neurosetae ....................................................................... 8

l 7B. Superior neurosetae thicker than median neurosetae ........................................................................ 9

i 8A. Notopodia elongated; elytra extending across dorsum; proximal ventral margin of neuropodia with row of papillae ........................................................................................ Lepidasthenia longicirrata

8B. Notopodia short; elytra leaving median dorsum uncovered; ventral neuropodial margin smooth ......

I .................................................................................................................. Lepidasthenia berkeleyae

I

I 2Not represented in the Santa Maria Basin material, but reported from southern California.

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9A. Body pigmented dark green or black; neurosetae with large secondary tooth .................................... I

........................................................................................................................ Lepidasthenia virens 2

9B. Body pigmented light yellow; secondary neurosetal tooth very small ............... Lepidasthenia gigas 2 I

10A. Lateral ceratophores subterminal, converging at midline on ventral side of prostomium; parapodia m

subbiramous, notosetae few or absent ............................................................................................ 11 |

10B. Lateral ceratophores ventral, attaching to prostomium and/or median ceratophore; parapodia biramous,

notosetae numerous ........................................................................................................................ 13 I

I

11A. Supraacicular neurosetae blunt, bifid; dorsum often with a pigment band across setiger 8 ...............

.................................................................................................................................. Arctonoe vittata •

11B. Supraacicular neurosetae hooked, similar to subacicular setae; dorsum without pigment band .... 12

12A. Elytra flat; ventral cirri subulate ............................................................................ Arctonoe pulchra I

12B. Elytra with ruffled margins; ventral cirri rudimentary ........................................... Arctonoefragilis 2

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13A. Neumsetae with large semilunar cusp at base of spinous region; notosetae with spinous pockets

extending to the tips ............................................................................................ Subadyte mexicana

13B. Neurosetae without basal cusp ....................................................................................................... 14 II

ending in capillary tips ......................................................................... Tenonia priops I

14A. All notosetae

14B. At least some notosetae ending in blunt or pointed tips .......................................................... _..... 15

m

15A. Upper notosetae ending in pointed tips; lower notosetae thinner, tapering to very fine points ....... 16 I

15B. All notosetae ending in blunt or pointed tips .................................................................................. 19 !

16A. Elytra thick, covered with polygonal cells; upper and lower neurosetae similar in shape ..................

........................................................................................................... Gaudichaudius iphionelloides •

16B. Elytra thin, translucent; upper and lower neurosetae differing in shape ........................................ 17 I

17A. Dorsum without pigment; margins of elytra with sparse fringe of digitate papillae ........................... I

............................................................................................................................. Hesperonoe laevis

17B. Dorsum pigmented; elytral margins smooth ................................................................................... 18 I

18A. Dorsum grayish-green; elytra with smooth surface; upper notosetae with coarse serrations .............

Hesperonoe adventor 2 !

18B. Dorsum salmon-colored; elytral surface with scattered conical tubercles; upper notosetae with indistinct

serrations .................................................................................................... Hesperonoe complanata 2 I

gl

19A. Notosetae with longitudinal striations and minute spinules arranged in two longitudinal rows ..... 20

19B. Notosetae with spinules arranged in transverse rows ..................................................................... 23 !

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20A. Neurosetae less numerous than notosetae, unidentate or with only indistinct indication of secondary

I tooth ................................................................................................................ Malmgreniella baschi 20B. Neurosetae as numerous as notosetae, with well-developed secondary tooth at least in median portion

of fascicle ....................................................................................................................................... 21

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21A. Prostomium with acute cephalic peaks; neurosetae with smooth, bare region between distal spinose

i 21B. bProstomium racts and basewithoftruncate secondaryanterior tooth lobes; ......................................................... distal spinose bracts on neurosetae Malmgreniella nearly reaching macginitiei base of

secondary tooth .............................................................................................................................. 22

I 22A. Supraacicular lobes small, short; dorsal cirri extending beyond tips of neurosetae; elytral surface

without distinct reticulation ....................................................................... Malmgreniella scriptoria

I 22B. Supraacicular lobes large, digitiform; dorsal cirri extending only to tips of neurosetae; elytral surface

distinctly reticulated .................................................................................... Malmgreniella nigralba

I 23A. Superior neurosetae with deeply incised tips; prostomium without eyes ............ Eucranta anoculata

23B. Superior neurosetae with tips bifid or entire; prostomium with eyes present ................................. 24 !

24A. All neurosetae with tips entire ........................................................................................................ 25

I 24B. At least some neurosetae ending in bifid tips ................................................................................. 27

i 25A. Median and lower neurosetae with terminal aristae; elytra thin, appearing smooth ...........................

....................................................................................................................... Bylgides macrolepidus

25B. Neurosetae with smooth, slightly hooked tips; elytra thick, with distinct surface macrotubercles .....

...................................................................................................................................................... 26

II i

26A. Elytra with large, dendritic, acutely-pointed spines; all notosetae with smooth, pointed tips ............

I ....................................................................................................................................... Eunoe senta

26B. Elylra with knob-like macrotubercles; some notosetae with blunt, sculptured, indistinctly splittips .......

................................................................................................................................. Eunoe oerstedt _

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27A. Superior notosetae more slender than most neurosetae; superiormost 1-2 neurosetae greatly enlarged,

I lance shaped ............................................................................................................. gsideria hastata 27B. Superior notosetae thicker than most neurosetae; superiormost 1-2 neurosetae not distinctly enlarged

.... . ........................................ ° ................ ° ............... ° ............ ° .......... ° ............... , ............... , ................ 28

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28A. Elytra with multipronged macrotubercles surrounded by large polygonal cells; neurosetae with

i 28B. Elytra secondary withtooth smoothremote macrotubercles from the tipand..................................................................... without polygonal cells; secondary tooth close Harmothoe to the neurosetal hirsuta

tip°°.. ............ °................ °° ................ °............ °.°° ............................. °........... °o° .......... °° ............... ° ....... 29

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29A. Anterior pair of eyes located on forward margin of the prostomium hidden beneath the prostomial n

peaks; dorsum lightly pigmented or mottled .................................................. Harmothoe imbricata 2

29B. Anteri ....................................................................................................................................................... or eyes located on lateral margins ofprostomium, visible dorsally; dorsum usually dark brown0 I

m

30A. Elytral surface with large thorn-like curved spines but without filiform papillae; fringing papillae R

short .............................................................................................................. Harmothoe multisetosa

30B. Elytral surface with filiform papillae but without large thorn-like curved spines; fringing papillae mum

long ..................................................................................................................... Harmothoe fragilis |

n

Description of Species U

Descriptions and illustrations are presented for a total of 22 species distributed among 13 genera that I

occur in or near the Santa Maria Basin and western Santa Barbara Channel as follows: U

Arctonoefragilis (Baird, 1863) I

Arctonoe pulchra (Johnson, 1897)

Arctonoe vittata (Grube, 1855)

Bylgides macrolepidus (Moore, 1905) n

Eucranta anoculata (Moore, 1910)

I' Eunoe senta Malmgren, 1865

,

Gaudichaudius iphioneIloides (Johnson, 1901)

Harmothoe fragilis Moore, 1910 I

n Harmothoe hirsuta Johnson, 1897

Harmothoe imbricata (Linnaeus, 1767) il

Harmothoe multisetosa (Moore, 1902) I

Hesperonoe laevis Hartman, 1961

Leyidasthenia berkeleyae Pettibone, 1948 I

u

Lepidasthenia longicirrata Berkeley, 1923

Lepidonotus spiculus (Treadwell, 1906), new combination

Malmgreniella baschi Pettibone, 1993 U

Malmgreniella macginitiei Pettibone, 1993 m

Malmgreniella nigralba (Berkeley, 1923) !

Malmg renielta scriptoria (Moore, 1910)

Subadyte mexicana Fauchald, 1972 D

U Tenonia priops (Hartman, 1961)

Ysideria hastata Ruff, new genus, new species I

m

An additional polynoid listed asHerdmanella sp. in the MMS Phase II voucher material was found to

be a 14-segment juvenile and could not be identified to the species level. I

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I Genus Arctonoe Chamberlin, 1920

I Type Species: Arctonoe lia Chamberlin, 1920, by original designation.

I cephalic Diagnosis. peaks, with Body well-defined elongate,lateral flattened, ceratophores with moreinserted than 100 at anterior segments. marginBiloand bed cprostomium onverging ventrally. without

Antennae, tentacular cirri and dorsal cirri smooth. Facial tubercle poorly developed; buccal segment with or

i without nuchal fold. Elytra on segments 2, 4, 5, 7.... 23, 26, 28, 29, 31, 33, continuing on alternate segments to end of body, often leaving dorsum exposed; elytra without tubercles or papillae. Parapodia subbiramous.

Notusetae short, serrated, unidentate or with notched tips; neurosetae stouter, longer, with unidentate hooked

tips.

I Remarks. The genusArctonoe was recently revised and placed into the new subfamily Arctonoinae

by Hardey (1989). A total of three species are known, all of which occur off California.

I

Aretonoe f_gilis (B_I, 1863)

I Figure 3.2 A-E

Lepidonotus fragilis Baird, 1863:108.

I Polynoefragilis: Johnson, 1897:179-181, pl. 7, figs. 36, 45, pl. 8, fig. 52. A rctonoefragilis: Berkeley and Berkeley, 1948:20, fig. 23.--Pettibone, 1953:64-66, pl. 31, figs 281-290.--

Hartman, 1968:45-46, figs. 1-3.--Blake, 1975b:165, pl. 27, fig. 70.--Hanley, 1989:9-12, figs. 4AI F, 5A-F (synonymy).

Material Examined. Washington: San Juan Archipelago (I).---California: off Dillon Beach (1).

I Description. Length to 85 mm for 100 segments; body without pigmentation. Prostomium bilobed, wider than long, with four small eyes on posterior half (Fig. 3.2A). Ceratophore of median antenna large,

stout; smooth style short, clavate, with filiformtip. Lateral antennae shorter and similar in shape. Palps twice

I prostomial length, abruptly tapering to short tips, occasionally with brown subterminal bands.

Basal lobes of tentacular segment achaetous, with two pairs of subequal tentacular cirri similar in

shape but longer than median antenna. Facial tubercle poorly developed. Cirrophores of ventral buccal cirri

I short, clavate, similar to median antenna.

large, cylindrical; styles

Large elytra nearly touching medially, thin, translucent, smooth, lacking tubercles or marginal papillae,

i with yellow, Subbiramous reddish, orparapodia greenish mottling. with smallMedian digitateandnotopodia posteriorandborders large,fothick, lded, convoluted truncate neuropodia (Fig. 3.2B)(Fig. .

3.2C). Notosetae short, slender, with transverse subdistal serrations and unidentate or notched tip (Fig.

I 3.2D sharp).hooked Neurosetae tip (Fig. longer, 3.2E).stout, with faint transverse serrations in swollen subdistal region, narrowing to

Dorsal cirri thick, clavate, often of varying lengths, with filiform tips. Dorsal tubercles large, rounded.

I Ventral 6. Shortcirri pygidial rudimentary cirri clavate after segment with 2. Ventral nephridial papillae low, usually inconspicuous, from segment filiform tips.

Biology. Arctonoefragilis is commonly found in the ambulacral grooves of the asteroid Evasterias

I troscheIii, where the elytra exhibit the general coloration of the host (Morris et aI., 1980). When viewed, the dorsal cirri mimic the appearance of the host's tube feet and spines. It has also been reported as a commensal

I

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Figure 3.2. Arctonoe fragilis: A, anterior end, dorsal view; B, median right elytron; C, median cirrigerous

parapodium, anterior view; D, notoseta; E, neurosetae.

I

with a number of other asteroid species, including Leptasterias aequalis, L. hexactis, Luidia foliolata, •

Orthasterias koehleri, Pisaster ochraceus, P. giganteus, Solaster dawsoni, and StyIasterias forreri. I

Remarks. Arctonoefragilis is characterized by vestigial ventral cirri and the ruffled posterior margins

of the elytra. Because specimens readily fragment and often loose their scales, Baird (1863) provided only a •

minimal description and he did not mention the convoluted elytral borders. However, subsequent authors I

have ascribed specimens to Baird's species because of the rudimentary nature of the venlxal cirri and the

typical association of the worm with asteroids, two features that Baird stressed.

Type Locality and Type Specimens. Esquimalt Harbor, Vancouver Island. Baird (1863) lists the

British Museum as the repository for the type specimen, but Hanley (1989) notes that during a recent search

it could not be located there. I

Distribution. Found in the rocky intertidal and in shelf depths from the Alaskan Peninsula at least as

far south as San Francisco, but uncommon in California. Treadwetl's (1937) report of the species from

Cedros Island offthe Baja Peninsula was based upon a mutilated specimen, and is doubtful (Pettibone, 1953). •

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I Arctonoe pulchra (Johnson, 1897)

Figure 3.3 A-E

I Polynoepulchra Johnson, 1897:177-179, pl. 7, figs. 34, 43, 43a, pl. 8, figs. 50, 50a, 50b.

Arctonoepulchra: Skogsberg, 1942:497-500, fig. D, 6-8; Pettibone, 1953:61-64, pl. 30, figs. 272-280; Hartman,

I 1968:47-48, figs. 1-4.--Blake, 1975b:175, pl. 27, fig. 69.--Hanley, 1989:12-16, figs. 6A-F, 7A-F (synonymy).

I Paci Material Examined. Washington: San Juan Island (3).--California, north of Bodega Bay (2); off fic Grove (2 syntypes); Western Santa Barbara Channel, off Point Conception, Sta. BRC-1, 69-74 m (2).

Description. Length to 70 mm for 70 segments; body colorless or mottled with brownish pigment

I dorsally. Median Prostomium bilobed, wider than long, with four moderate-sized eyes on posterior half (Fig. 3.3A). antenna with large ceratophore and smooth cylindrical style with subdistal swelling and elongated

terminal tip; lateral antennae shorter, pyriform, with long filamentous tips. Palps stout, wrinkled, abruptly

I tapering to short tips, often with brown subterminal pigment bands.

Lateral tentaculophores long, usually achaetous, with two pairs of smooth, subequal tentacular cirri

similar in shape but longer than median antenna. Facial tubercle poorly developed. Ventral buccal cirri on

I segment 2 long, similar to median antenna.

Suboval or reniform elytra covering dorsum. Each elytron thin, flat or slightly undulate, lacking

i tuberclesParapodia and marginal subbiramous, papillae, often withwith small, reddis digitiform h-brown notopodia pigment conce on anterodorsal ntrated over elytrophore side of longer, (Fig.broader 3.3B).

neuropodia; neuropodia with rounded pre- and postsetal lobes (Fig. 3.3C). When present, notosetae short,

I stout, transverse with sspinous triationsedge in swollen and blusubdistally nt, notched region, tip (Fig.narrowing 3.3D). Altol nsharp eurosetae hookedsimilar tip (Fig. in shape, 3.3E).stout, with faint

Ceratophores of dorsal cirri large, cylindrical, tapering; styles long, with subterminal enlargement

I apapillae nd long filamentous tip. Dorsal tubercles conspicuous knobs. Ventral cirri short,subulate. Ventral nephridial tow, inconspicuous, from segment 6. Pygidium rounded with pair of short, subterminally swollen

cirri.

I Biology. Arctonoe pulchra is normally found commensal with the holothurian Parastichopus californicus, using its hooked neurosetae to cling to the epidermis, and crawling into the host's mouth when

disturbed (Johnson, 1897; Morris et al., 1980). It is also often associated with the giant gumboot chiton

I Cryptochiton stelleri, the keyhole limpet Megathura crenulata, the terebellid Loimia medusa, and a number of additional asteroid and gastropod species (Pettibone, 1953).

Remarks. Specimens of Arctonoe pulchra exhibit a wide variety of coloration on the etytra and

I dorsal body that matches the host organisms in cases where the worm is exposed to the environment. These

colors, however, usually fade quickly in preservative. Specimens can often be recognized by the dark pigment

spot occurring on each elytron, but this pigmentation is variable and may be entirely lacking. Notosetae are

I usually present in small individuals, but they tend to be lost, especially in the posterior segments, as the specimens grow.

Type Locality and Type Specimens. California, Pacific Grove, 2 syntypes (LACM-AHF Poly

An additional is in the collections of the Museum in 0044). syntype Zoological of Hamburg Germany (HZM

PE 127b), apparently a gift to Ehlers from either Johnson or the Museum of the University of California

(Hanley, 1989).

I Distribution. Found from the Gulf of Alaska to Baja California, intow intertidal and shelf depths.

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Figure 3.3. Arctonoe pulchra: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous •

parapodium, anterior view; D, notoseta; E, neurosetae. III

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Aretonoe vitCata (Grube, 1855)

Figure 3.4 A-F I

Polynoe lordi: Johnson, 1897:175-177, pl. 7, figs. 35, 44, pl. 8, fig. 51.

Acholoe vittata: Marenzeller, 1902:576-577, pl. 3, fig. 13. l

Arctonoe lia Chamberlin, 1920:6-7, pl. 1, figs. 1-4, pl. 2, figs. 1-3.

Halosydnoides vittata: Okuda, 1936:565-68, figs. 4-5.

Arctonoe vittata: Hartman, 1939:2%30, pl. 3, figs. 33-37; 1948:11-12, fig. 2a-f; 1968:49-50, figs. 1-3.--

Skogsberg, 1942:489-497, figs. B-D.--Berkeley and Berkeley, 1948:20-21, figs. 24-25.--Pettibone,

1953:57-61, pl. 28, figs. 251-258, pl. 29, figs. 259-271.--Uschakov, 1955:132-133, fig. 23E-J;

1982:116-117, pl. 35, figs. 1-7.--Blake, 1975b: 175, pl. 27, figs. 67-68.--Hanley, 1989:4-9, fig. 1A- I

F, fig. 2A-L (synonymy). II

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I Material Examined. Alaska, Boca de Quadra (3).--Washington, San Juan Island (3).--Oregon, off

Tillamook Head (I).---California, off Tomales Point (3); Santa Mafia Basin, off Point Sal, Sta. R-8, 90 m (1).

I Description. Length to 100 mm for 89 segments; body often with solid pigment band across segment 7 or 8. Prostomium bilobed, wider than long, with four small subequal eyes on posterior half (Fig. 3.4A).

Median antenna with thick ceratophore and short, smooth, clavate style with long filiform terminal tip; lateral

I antennae stubby with long filamentous tips. Palps short, abruptly tapering to short tips, often with dark subterminal pigment bands.

Lateral tentaculophores large, achaetous, with two pairs of subequal cirri similar in length and shape

I to median Facial tubercle Ventral buccal cirri of 2 with

antenna. poorly developed. segment large, cylindrical

cirrophore and styles similar to median antenna.

i Small suboval or reniform elytra leaving median dorsum uncovered anteriorly, and overlapping in posterior segments; posterior attachment pattern variable and often asymmetrical. Elytra thin, translucent,

smooth, lacking tubercles or marginal papillae (Fig. 3.413),usually white with some degree of black or brown

i mottling.Subbiramous parapodia (Fig 3.4C) with small, digitiform notopodia; neuropodia broader, longer,

with short, rounded pre- and postsetal lobes. Notosetae short, slender, with subterminal serrations and blunt

I notched segment tips 2 similar (Fig. 3.4D); to notosetae, notosetae lrger, oftenwithlacking fine subterminal in median and spines posterior and notched, segments. bluntSuperior tips (Fig.neurosetae 3.4E); inferior after

neurosetae stouter, with faint transverse serrations in subterminal inflated region, narrowing distally to pointed

I hooked tips Dorsal (Fig.cirri3.4F). with bulbous cirrophores; styles short, cylindrical to clavate, narrowing to tong filiform

tips. Dorsal tubercles well-developed. Ventral cirri short, subulate. Small, terete ventral nephridial papillae

I conspicuous but from segment 6 to end of body. Terminal pair of pygidial cirri resembling dorsal cirri in shape, longer.

Biology. Arctonoe vittata is found both free-living and in commensal association with a wide variety

I of asteroids, gastropods, and polychaetes, where they often tend to match their hosts in body coloration. In California it is most often found with the giant gumboot chiton Cryptochiton stelleri, the asteroidDermasterias

imbricata, and in the pallial grooves of the keyhole limpet Diodora aspera (Morris et al., 1980). Skogsberg

I (1942) and Pettibone (1953) provide a comprehensive list of hosts. Remarks. The dark pigment band across the dorsum of segment 8 is not present in all individuals.

The size of the elytra is also variable, and tends to be related to the particular host. Specimens associated with

I Cryptochiton and Diodora normally have a distinct middorsal gap, while those found with Dermasterias may have larger elytra that touch medially.

The number of notosetae in each segment appears to be a function of specimen size. In small specimens,

I most of the notopodia have setae, although the number per fascicle decreases posteriorly. In larger individuals,

however, the notosetae are usually absent in all but the anteriormost segments.

Type Locality and Type Specimens. Sitka, Alaska. Skogsberg (1942) lists the type specimen in the

I Leningrad Museum.

i Distribution. Found from Japan and the Bering Strait along the Pacific coast to Ecuador, in the

i • rocky intertidal zone and shelf depths. i

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man I') pa _-. ,_-_n \(c _ _ I

ii ,I

I

A o f ,

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Figure 3.4. Arctonoe vittata: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous

parapodium, anterior view; D, notoseta; E, superior neuroseta; F, inferior neuroseta.

I

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I Genus Bylgides Chamberlin, 1919

I Type Species: Bylgia elegans Th6el, 1879, by monotypy.

Diagnosis. Body short, flattened, with about 40 segments. Prostomium with ceratophores of lateral

I antennae inserted beneath weak to well-developed cephalic peaks. Antennae, palps, and cirri with small

papillae. Fifteen pairs of elytra on segments 2, 4, 5, 7... 23, 26, 29, and 32, completely covering dorsum.

Parapodia biramous. Notosetae stout, with transverse rows of spinules and smooth tips; neurosetae thinner,

I long, with subterminal spines and terminal aristae.

Remarks. Th6el (1879) erected the genus Bylgia for B. elegans, a species with slender, capillarytipped neurosetae and a prostomium with lateral antennae but lacking the median antenna. Chamberlin

I (1 9), noting Bylgia was preoccupied crustacea, replaced nameBylgides. Later, Augener

9 1 that in it with the

(1928) proposed the genusAntinoella for species that were similar in most respects but that possessed rather

than lacked a median antenna. More recent work has shown that the holotype of Bylgia elegans has a

I prostomium was misdiagnosed. (1982) placedAntinoella synonymy

defective and Therefore Uschakov into

withBylgides based upon information provided and subsequently published by Pettibone (1993a). The genus

encompasses a number of arctic and boreal species, only one of which is found off California. I

Bylgides macrolepidus (Moore, 1905)

I Figure 3.5 A-E

i Antinoe Antinoellamacrole macrolepida pida Moore, 1905:538-541, pl. 35, figs. 21-23.--Berkeley and Berkeley, 1948:14, fig. 15. : Hartman, 1968:41, figs. 1-3.--Banse and Hobson, 1974:25, fig. 2a..--Lissner et

aI., 1986:D-7.

i BBylgides ylgides macrolep macrolepidus ida: Uschakov, 1982:157-58, pl. 57, figs. 1-6. : Pettibone, 1993a:21-23, fig. 13a-k.

i Material Examined. Alaska, Chatham Strait (holotype, 4 paratypes, and 1 additional specimen); Boca de Quadra (2).--British Columbia, Gulf of Georgia (paratype); off Vancouver Island (paratype).--

California, Santa Maria Basin, off Point Estero, Sta. 3, 291 m (1), off Point Arguello, Sta. 59, 216 m (1).

I intersegmen Description tal transver . Lsene ligthnes.toBilob 45 mmed fporros39tomiusegme m nbroad, ts; dorsantaleriocirri rlyandrounded, body redd withishsmall browncephwitalhiclipeak ghters

(Fig. 3.5A). Anterior eyes large,with well-developed lenses, covering anterior marginof prostomium;posterior

II eyes smaller, sometimes concealed by base of etytrophores of second segment. Median ceratophore large,

II cylindrical; style long, tapering, with slight subterminal swelling and scattered papillae. Ceratophores of

i lateral antennae small, spherical, situated beneath cephalic peaks; subulate styles short,tapering to filamentous

i tips. Palps Tentaculophores about 6 timeslong, prostolateral mial tolength, palps,taperi with nprojecting g to acuteaeicular tips, wlobe ith numerous and severaslmaaeicul ll papillae. ar setae. Subequal

tentacular cirri similar in shape but shorter than median antenna, with scattered papillae. Facial tubercle not

I developedElytra . Ventral buccal cirri long, acuminate, with scattered papillae. strongly imbricated but easily lost, large, thin, translucent, marbled with pale brown pigment.

Each elytron appearing smooth but covered with tiny, curved conical microtuberctes and scattered papillae;

I margin with sparse fringe of minute papillae (Fig. 3.5B).

!

I 119
I

I

Figure 3.5. Bytgides macrolepidus: A, anterior end, dorsal view; B, median left elytron with detail of i

microtuberclesand marginalpapillae;C, median cirrigerousparapodium, posterior view;D, notoseta; i

E, median neuroseta with detail of aristate tip.

!

Parapedia biramous. Notopedia low, cylindrical, extending into long digitate lobe with emergent i

acicula. Neuropodia long, stender, with obliquely truncate postsetal lobe and pointed presetal lobe extending li

into a long digitate cirrus above projecting acicula (Fig. 3.5C). Notosetae numerous, short, thick, nearly

u

straight, with transverse rows of spines and smooth pointed tips (Fig. 3.5D). Neurosetae very numerous, i

slender, long, with subdistal spines; tips of median and inferior neurosetae with fine hairs and long terminal i

aristae (Fig. 3.5E).

a

Dorsal cirri elongate, extending beyond neurosetae, with proximal clavate papillae, subterminal m

swelling, and smooth distal tips. Dorsal tubercles large, well-developed. Ventral cirri slender, tapering, with I

scattered papillae. Ventral nephridial papillae cylindrical, from segment 6. Pygidium elongate, with dorsal

anus; pygidial cirri similar to dorsal cirri. ll

Habitat. Known from scattered records in depths from 36-537 m, on bottom types consisting of I

green mud, fine to coarse sands, and rocks.

!

,20 |
!

I Remarks. Most representatives of the genus Bylgides are found in the arctic and far north Atlantic.

B. elegans has been reported from the Bering Sea, but it differs by lacking the large lensed eyes on the anterior

I prostomial aristate tipsmargin are diagnostic beneath features the cephalic for B.peaks. macrolepidus These bulbous in the Santa eyes alMaria ong with Basintheregion. slender neurosetae with

Type Locality and Type Specimens. Off Freshwater Bay, Chatham Strait, Alaska, 516-536 m,

I holotype 5557, 5559) (USNMfrom 5509) the Gulf andof4 paratypes Georgia and(USNM off Vancouver 5561-5563, Island, ANSPBritish AD2011). Columbia. Additional paratypes (USNM

Distribution. Found in shelf and upper slope depths from the southern Sakhalin Islands around the

I Pacific rim as far down as southern California.

i Genus Eucranta Malmgren, 1865

Type Species: Eucranta viIlosa Malmgren, 1865, by monotypy.

!

Diagnosis. Body short, strongly arched, with about 40 segments. Prostomium with weak to welldeveloped cephalic peaks, with ceratophores of lateral antennae inserted beneath anterior margin. Facial

I tubercle and nuchal fold absent. Fifteen of 2, 4, 5, 7 23, 26, 29, and 32, pairs elytra on segments

completely covering dorsum. Parapodia biramous. Notosetae short, thick, with blunt tips. Most neurosetae

slender, long, with coarse spines in subdistal expanded region and unidentate tips; superior and/or inferior

i ! neurosetae thinner, with deeply incised tips.

I

Remarks. The genus Eucranta is known from a few species found mainly in arctic and subantarctic

waters. The genus is distinguished by having a group of neurosetae in which both forks of the deeply cleft tip

I are about the same size.

I Eucranta anoculaUt (Moore, 1910)

Figure 3.6 A-F

I Moore, pl. 30, figs.

Antinoe anoculala 1910:358-361, 34-40.

Antinoella anoculata: Hartman, 1968:39-40, figs. 1-7.

Eucranta anoculata: Fauchald and Hancock, 1981:23.

I Bylgides'anoculata: Uschakov, 1982: 158-159, pl. 57, figs. 7-11.

Antinoe sp. A: Blake et al., 1988:A-3.

Antinoe sp. C: Steinhauer and Imamura, 1990:F-1 (partita).

I Material Examined. California, off Monterey Bay (holotype and 2 paratypes); Santa Mafia Basin,

off Point Buchon, Sta. 16, 591 m (3), off Point San Luis, Sta. R-3, 409 m (1), Sta. 27, 611 m (2), off Point

I Sal, Sta. R-7, 565 m (8), off Purisima Point, Sta. R-6, 410 m (1); off San Diego (1).

Description. Length to 36 mm for 37 segments; body dusky or colorless, with iridescent cuticle.

Prostomium longer than wide, divided into two lobes, with well-developed cephalic peaks; eyes lacking (Fig.

I 3.6A). Ceratophore of median antenna large, tapering; style slender, acuminate, about twice prostomial

m

length, with numerous minute papillae. Short, tumid ceratophores of lateral antennae attached beneath cephalic

peaks, with short, subulate styles. Longitudinally striated palps to four times prostomial length, tapering to

I slender tips.

!

I 121
I

I

Figure 3.6. Eucranta anoculata: A, anteriorend, dorsal view; B, median rightelytron;C, median cirrigerous I

parapodium, anteriorview; D, tip of notoseta;E,tip ofsuperior neuroseta;F,tip of inferior nearoseta

(D-F from Moore, 1910). I

I

I

Tentaculophores elongated, lateral to palps, each with long acicular lobe and 0-3 stout, curved setae.

Dorsal tentacular cirri with numerous minute papillae, similar in length to median antenna; ventral tentacular I

cirri slightly shorter. Facial tubercle not developed. Ventral buccal cirri on segment two as long as ventral I

tentacular cirri, tapering to slender tips, with minute papillae.

Elytra large, covering dorsum, colorless or with streaks of greenish-yellow pigment, first pair circular •

and following pairs reniform, soft, membranous, translucent, with inconspicuous microtubercles anterior to I

attachment scar; lateral border with very short papillae (Fig. 3.6B).

Notopodia short, stout, extending into long digitate lobe with stout emergent acicula; neuropodia I

larger, broader, tapering to short digitate cirrus above emergent acicula (Fig. 3.6C). Notosetae stout, slightly

curved, with numerous transverse rows of inconspicuous spines, tapering to smooth blunt tips (Fig. 3.6D).

Neurosetae more numerous, long, slender. Upper neurosetae with long spinous region and thin, deeply incised I

tips (Fig. 3.6E); median and inferior neurosetae with prominent serrations onsubdistal expanded region, and

elongated acute smooth tips (Fig. 3.6F).

I

122 I
!

I Dorsal cirrophores long, slender; styles elongate, evenly tapering, extending well beyond neurosetae,

with short clavate papillae. Dorsal tubercles prominent on anterior segments. Short ventral cirri slender,

I tapering, furrows fromwithout segment papillae. 6. Pygidium Ventral nephridial rounded; papillae anal cirrilong, twicecylindrical, length of curving dorsal cirri, backwith into scattered intersegmental small

clavate papillae.

I Habitat. Found in muddy substrates in depths from 565-1600 m. Remarks. Moore (1910: pl. 30,fig. 38d) figured the tipof adeeply incised neuroseta, buthe incorrectly

assigned it to a seta in the ventral part of the fascicle. Hartman (1968) perpetuated this error,but later

I corrected the observation by noting that the species has "deeply incised slender superior neurosetae" (Hartman, 1978).

In some instances the tips of the superior neurosetae appear to be flattened with thickened edges, but

I with no median split. It is probable that the forks when the thin center between the separate appear area

thickened edges has worn away.

i m, Type Locality and Type Specimens. OffPoint Pifios Light, Monterey Bay, California, 1372-1401 holotype (USNM 016882) and 2 paratypes (ANSP AD2900). Although Moore (1910) indicated only one

cotype [=paratype] in his publication, the vial at the Academy of Natural Sciences contains two specimens

with a label in Moore's handwriting stating "cotypes."

I Distribution. Known from bathyal depths off Oregon, California, and northern Mexico. Specimens

reported by Uschakov (1982) from the Kurile Islands and the Sea of Okhotsk may be representatives of

I another genus since incised neurosetae were not observed.

Genus Eunoe Malmgren, 1865

!

Type Species: Polynoe nodosa Sars, 1861, designated by Uschakov, 1955.

I Diagnosis. Body short, strongly arched, with about 40 segments. Prostomium with weak to welldeveloped cephalic peaks; ceratophores of lateral antennae inserted beneath anterior margin. Facial tubercle

I absent; buccal segment with or without nuchal fold. Antennae, palps, and cirriwith filiform papillae. Fifteen pairs of elytra on segments 2, 4, 5, 7.... 23, 26, 29, and 32, completely covering dorsum. Parapodia with

well-developed noto- and neuropodia. Notosetae at least as thick as neurosetae, with blunt or split smooth

I tips; neurosetae unidentate and subterminally serrated.

Remarks. Approximately 40 species are described in this genus, but most are restricted to the colder

waters of the northern and southern hemispheres. Of the two species reported from off California, only one

I was in the Santa Mafia Basin collections.

present

I Eunoe senta (Moore, 1902)

Figure 3.7 A-F

I Gattyana senta Moore, 1902:259-263, pl. 13, figs. 1-13; 1905:525 (correction of type locality).

Eunoe senta: Hartman, 1939:52-53, pl. 10, figs. 128-133; 1968:57-58, figs. 1-6.--Blake, 1975b: 175, pl. 28,

I fig. 78.--Uschakov, 1982:179-180, pl. 67, figs. 5-8.

Material Examined. Alaska, offlcy Cape (holotype and paratype).---Califomia, Santa Mafia Basin,

I off Point Buchon, Sta. 6, 109 m (2), off Purisima Point, Sta. BRC-14, 96-105 m (1).

i 123
I

I

I

Figure 3.7. Eunoe senta: A, anterior end, dorsal view; B, median right elytron; C, median cirrigerous parapodium, •

posterior view; D, superior notoseta; E, median notoseta;F, distal portion of median neuroseta.

I

,24 I
!

I Description. Length to 22 mm for 36 segments; dorsum pale yellow or colorless. Prostomium wider

than long, anteriorly divided into two broadly rounded lobes, with low cephalic peaks (Fig. 3.7A). Two pairs

i of Median black, circular eyes; anterior pair slightly larger, located on lateral margin of prostomium at widest point. ceratophore massive; slender style three times prostomial length, with brown pigment and numerous

long papillae basally, and long bare ffliform tip. Lateral antennae with stubby ceratophores inserted beneath

i cephalic papillae, peaks, as long and as median short, acuminate, antenna, tapering papillateto filiform styles. tip. Palps stout, with longitudinal rows of digitiform

Tentaculophores short, broad, lateral to palps, each with pointed acicular lobe and bundle of stout,

I blunt Facialsetae tubercle on anterior absent. face. BuccalSubequal segmenttentacular with nuchalcirri foldpapillate, coveringsimilar posteriinorlength marginandof pros shapetominm. to medianCirropho antenna. res

of ventral buccal cirri large; styles acuminate, papillate, extending to end of neurosetae.

i pigment Elytra circular to subreniform, soft, thick, easily detached, colorless or with irregular patches of , darkest on lateral margins. Surface covered with numerous hard, chitinous tubercles increasing in

size and complexity toward lateral and posterior borders; smallest spines simple or minutely bifurcate, and

I largest spines dendritic with acutely pointed branches (Fig. 3.7B). Margin with sparse filiform papillae. Notopodia stout, extending into conical lobe with long emergent acicula; neuropodia longer, obliquely

truncate with similar acicular lobe and protruding acicula (Fig. 3.7C). Numerous notosetae arranged in

I whorls; middle upper notosetae short, with close-set transverse rows of spinules and short, smooth tips (Fig. 3.7D); and lower notosetae longer, straight, with subterminal region of widely-spaced transverse rows of

short spinules encircling shaft, and smooth points (Fig. 3.7E). Neurosetae subequal to notosetae in length and

I thickness, coarsely serrated in subterminally expanded region, with slightly hooked, smooth tips (Fig. 3.7F).

Dorsal cirrophores large, cylindrical; styles long, digitiform, narrowing to filamentous tips, with

numerous long papillae proximally. Dorsal tubercles prominent. Slender, smooth ventral cirri extending only

I to base of neuropodia. Short cylindrical nephridial papillae present from segment 6, continuing throughout body. Slender pygidial cirri similar to dorsal cirri, with numerous long papillae.

Habitat. Commonly found in shelf depths from 5-346 m on mixed sand and gravel bottoms.

I Remarks. The spiny elytra, papillose cirri, and dense bundles of notosetae provide this species with

a shaggy aspect. Moore (1902) originally listed the type locality as McCormick Bay in western Greenland.

However, this designation was subsequently attributed to a misplaced vial label, and was corrected by Moore

I (1905) to samples taken in Alaska.

Type Locality and Type Slmeimens. Icy Cape, Alaska, holotype (ANSP AD266 and slide 2.30) and

I 1paratypeDistribution. (ANSP AD5219). Known from the Kurile Islands around the North Pacific rim to southern California.

Also reported by Ditlevsen (1937) from western Greenland.

I Genus Gaudichaudius Pettibone, 1986

I Type Species: lphione cimex Quatrefages, 1866, by original designation.

I Diagnosis. Body short, flattened, with about 38 segments. Prostomium without cephalic peaks but

with ceratophores of lateral antennae inserted beneath anterior margin. Facial tubercle absent; buccal segment

I with dorsum; nuchat surface fold.ofFifteen elytra with pairsprominent of etytra onpolygonal segmentsareas. 2, 4,Parapodia 5, 7.... 23, biramous. 26, 29, andUpper 32, completel notosetae yshort, covering with

transverse spinules and blunt tips; lower notosetae longer, more slender, tapering to fine tips. Neurosetae

I thicker than notosetae, with expanded subdistal spinous region and bare, slightly hooked tips.

i 125
!

Remarks. Pettibone (1986) erected the genusGaudichaudiustoencompasslphionecimexQuatrefages, R

1866 (originally described incorrectly), andHarmothoe iphionelloides Johnson, 1901. Johnson's species had

been placed into the genus Gattyana Mclntosh, 1897, by Berkeley and Berkeley (1945) based essentially on n

the characters of the setae, although it was recognized that it differed from other members of the genus in I

details of the prostomium and elytra (Pettibone, 1953).

ii

Gaudichaudius iphionelloides (Johnson, 1901) I

Figure 3.8 A-F

Harmothoe iphionelloides Johnson, 1901:391-92, pl. 1, figs. 2-7.

Gattyana iphioneIloides: Berkeley and Berkeley, 1945:321; 1948:12, fig. 12.--Pettibone, 1953:44-45, pl. t

22, figs. 194-200.--Uschakov, 1982:155-156, pl. 54, figs. 7-12. I

Gaudichaudius iphionelloides: Pettibone, 1986:37-40, fig. 18a-1(synonymy).

Material Examined. Alaska, Icy Strait (1).--British Columbia, off Valdez Island (1).--Washington: n

Puget Sound (holotype + 1 specimen), Neah Bay (l).---California, Santa Maria Basin, off Point San Luis,

Sta. BRA-25, 65-72 m (1). 1

Description. Length to 35 mm for 36 segments; dorsum generally colorless, ventrum iridescent, n

Prostomium bilobed, wider than long, without cephalic peaks (Fig. 3.8A). Two pairs of large eyes; anterior

pair located on lateral margins of pmstomium slightly anterior to widest part, and smaller posterior pair near m

posterior border. Median ceratophore long, slender, inserted between prostomial lobes; style long, papillate, U

with subterminal swelling, dark pigment ring, and fdiform tip. Ceratophores of lateral antennae short,

cylindrical, inserted subterminally; styles acuminate, with scattered papillae, shorter than median antenna.

Palps stout, tapering to blunt tips, with numerous short papillae, l

Tentaculophores long, lateral to palps, each with short acicular lobe and several long setae on anterior

face. Two pairs of subequal tentacular cirri similar in shape and pigmentation to median antenna. Facial •

tubercle absent. Buccal segment with nuchal fold covering posterior margin ofprostomium. Ventral buccal |

cirri long, acuminate, attached at base of parapodia lateral to mouth.

Etytra strongly imbricated, thick, large, suboval to deeply reniform. Elytral surface light to dark •

brown, covered with variably-sized polygonal cells, each with central pointed or flattened tubercle, occasionally

with black pigment patch (Fig. 3.8B). Lateral and posterior margins with fringe of short papillae. Elytrophores

large, elongate, with additional lobe near inner margin. 1

Pampodia biramous. Notopodia rounded, extending into bluntly pointed acicular lobe; neuropodia

broader, longer, diagonally truncate, projecting into thick presetal lobe with short digitate superior cirrus (Fig.

3.8C). Notosetae numerous; upper notosetae short, stout, curved, pointed, with close-set transverse rows of

fine spinules (Fig. 3.8D); lower notosetae elongated, slender, tapering to fine points (Fig. 3.8E). Neurosetae

thicker than notosetae, numerous, with short subdistal enlarged spinous region tapering to bare hooked tip

(Fig. 3.8F). n Dorsal cirri with elongate, basally bulbous cirrophores; styles long, with subterminal swelling and

long papillae on distal half. Transversely elongated dorsal tubercles present on cirrigerous segments. Ventral

cirri short, acuminate, with sparse papillae. Elongated, cylindrical nephridial papillae projecting between

parapodia from segment 6. Terminal pair of pygidial cirri short, papillate.

Habitat. This species is found in intertidalZostera beds, and subtidally on muddy bottoms with rock

or shell components. •

I

I

126 n
I Figure 3.8. Gaudichaudius iphioneUoides:A, anterior end, dorsal view; B, median right elytron; C, median cirrigerous parapodium, anterior view; D, superior notoseta; E, inferior notoseta; E distal portion

of median neuroseta.

I

I GanyanaRemarks. Gaudichaudius iphionelloides is readily distinguishable from several species ofthe genus found in the north Pacific by the presence of polygonal areas surrounding simple, flattened tubercles

on the elytra.

I Type Locality and Type Specimens. Pleasant Beach near Seattle, Washington. The holotype (MCZ 1897) in the Museum of Comparative Zoology at Harvard has dried out in the past, and now consists of the

posterior 18 setigers along with some free elytra.

I Previously recorded from the Kurile Islands, across the Bering Strait to Alaska, and

Distribution.

down to Puget Sound. The specimen from the Phase I hard-substrate station offPoint San Luis represents an

significant southward extension of the known range. !i

i

|

i 127
!

Genus Harmothoe Kinberg, 1855 I

Type Species: Harmothoe spinosa Kinberg, 1855, designated by Bergstrrm, 1916. I

Diagnosis. Body short, flattened, with up to 45 segments. Bilobed prostomium with or without II

cephalic peaks; lateral ceratophores inserted beneath anterior margin. Facial tubercle present or absent; II

dorsal nuchal fold absent. Fifteen pairs of elytra on segments 2, 4, 5, 7.... 23, 26, 29, and 32, occasionally

m

leaving posteriormost segments exposed; elytral surface with or without microtubercles, macrotubercles, and mini

papillae. Parapodia biramous. Notosetae stout, as thick or thicker than neurosetae, with distinct transverse II

spinous rows and bluntly pointed, glabrous tips. Neurosetae longer, with expanded subdistal spinous region;

m

at least some neurosetae with secondary subterminal tooth. m

Remarks. Harmothoe is a large genus with over 100 valid species. Several genera have been I

proposed to split up this large assemblage (e.g. Lagisca, Evarnella), but the distinctions between these become

obscured as additional species are found. Four species of Harmothoe occurred in the Santa Maria Basin m

material, including one species often listed in the genus Lagisca. •

Harmothoefragilis Moore, 1910 I

roll

Figure 3.9 A-F

Harmothoe (Evarne)fragilis Moore, 1910:353-357, pl. 29, figs. 29-30, pl. 30, figs. 31-33. !

Harmothoe impar: Ditlevsen, 1917:12-13, pl. 2, fig. 16, pl. 3, fig. 11.--Uschakov, 1955:157, fig. 41A-E.--

Pettibone, 1956:550-551. Not Johnston, 1839. m

Harmothoefragilis: Pettibone, 1963:39-41, fig. 9g.--Hartman, 1968:75-76, figs. I-4. R

Antinoe sp. C: Steinhauer and Imamura, 1990:F-1 (partim). U

Material Examined. Washington, Puget Sound (2).--California, Santa Maria Basin, offPoint Buehon, !

Sta. 6, 109 m (1), off Point Sal, Sta. R-7, 565 m (1), off Point Arguello, Sta. BRA-4, 168-237 m (1); off Santa U

Catalina (holotype and paratype); off Santa Barbara Island (paratype and 1 additional specimen); off San

Clemente Island (paratype). I

II

Description. Length to 26 mm for 41 segments; dorsum pale to deep brown with two thin transverse

white stripes converging at elytrophores or dorsal tubercles; ventrum unpigmented. Bilobed prostomium m

slightly wider than long, with acute cephalic peaks (Fig. 3.9A). Two pairs of large eyes; anterior pair located Ii

on lateral margins of prostomium at widest point; posterior pair slightly smaller, at posterior comers of

prostomium. Median ceratophore cylindrical, inserted between prostornial lobes, reaching slightly beyond m

cephalic peaks, pigmented dark brown; cirriform style over twice prostomial length, with numerous long E

papillae, subterminal swelling, and filiform tip. Lateral ceratophores short, cylindrical, inserted beneath m

cephalic peaks; styles less than prostomial length, subulate, with scattered papillae. Thick palps as long as m

median antenna, covered with minute papillae, tapering to short terminal filaments. I

Tentaculophores elongate, lateral to palps, each with digitate acicular lobe and several stout setae on

anterior face. Subequal tentacular cirri similar in length and shape to median antenna. Facial tubercle and m

nuchal fold absent. Ventral buccal cirri slender, reaching to end of neurosetae, with small papillae and •

filiform tips.

Ill

Elytra soft, loosely attached, circular to reniform, completely covering dorsum. Elytral surface pale III

brown, covered with numerous conical or multibranched microtubercles and occasional longer filiform papillae; I

posterior surface with several large, ovoid, blister-like, darker brown macrotubercles (Fig. 3.9B). Lateral

margins with thick fringe of long papillae. I

128 I
I

Figure 3.9. Harmothoefragifis: A, anterior end, dorsal view; B, median left elytron with detail of microtubercles

and marginal papillae; C, median cirrigerous parapodium, anterior view; D, notoseta; E, distal

I portion of superior neuroseta; F, tip of median neuroseta.

I Notopodia large, extending into acutely pointed lobe with emergent acicula; neuropodia longer, slightly

broader, tapering to long digitate terminal cirrus above emergent acicula (Fig. 3.9C). Notosetae stout, curved,

with transverse rows of spinules extending to blunt sculptured tips (Fig. 3.9D). Superior neurosetae slender

I with finely bidentate tip (Fig. 3.9E). Median neurosetae longer, with expanded subdistal spinous region tapering to hooked tip and slender secondary tooth (Fig. 3.9F); inferior neurosetae shorter, often unidentate.

Dorsal cirrophores long, swollen basally; cirri elongate, extending beyond neurosetae, with bare

I filiform tips and numerous subterminal papillae. Dorsal tubercles prominent, often heavily pigmented. Ventral cirri with low cirrophores; styles short, tapering, with occasional minute papillae. Cylindrical nephridial

papillae present at base of parapodia from segment 6. Pygidium with erenulate border surrounding dorsal

I anus; terminal pygidial cirri longer than dorsal cirri, with numerous long papillae.

I! 129
!

Habitat. Harmothoefragilis is found in soft mud, sand and gravel from the shallow subtidal to slope I

depths of nearly 3000 m.

Remarks. H.fragilis is very similar toll. impar (Johnston, 1839) from the Atlantic coast of Europe n

and the Mediterranean, and there is confusion in the literature concerning these two species. Moore's species

differs from H. impar in having shorter lateral antennae, filiform papillae on the elytral surface, and marginal

papillae that are essentially restricted to the lateral border. Moore (1910) illustrated a parapodium studded 1

with small spherical bodies projecting from the surface which are probably parasitic in origin. These spheres

were not observed in the Santa Maria Basin specimens.

Type Locality and Type Specimens. Southern California, off Santa Catalina Island, 278-296 m, n

holotype (USNM 17148) and paratype (USNM 73013); off Santa Barbara Island, 435-567 m, paratype I

(USNM 17147); off San Clemente Island, paratype (ANSP 2877).

1

Distribution. Widely distributed throughout the arctic, and into boreal waters in both the Atlantic 1

and Pacific Oceans. It has been reported in the Chukchi Sea and along the Pacific rim as far south as northern

Japan and southern California. n

Harmothoe hirsuta Johnson, 1897

Figure 3.10 A-E I

Harmothoehirsuta Johnson, 1897:I82-3, pl. 6,figs. 27-29, pl. 7, fig. 38, pl. 8, fig. 53.---Chamberlin, 1919:51-

54, pl. 2, figs. 2-8, pl. 3, fig. 1.--Hartman, 1968:77-78, figs. 1-6.--Blake, 1975b:176, pl. 28, fig. n

75. lm

Material Examined. California, Western Santa Barbara Channel, off Point Conception, Sta. BRC- n

1, 69_74 m (2); off san Pedro (holotype); off San Nicolas Island (1).

Description. Length to 32 mm for 40 segments; dorsum dusky, with brown pigment patches anteriorly,

or unpigmented; ventrum iridescent. Prostomium bilobed, wider than long, with prominent cephalic peaks I

and two pairs of large eyes; anterior pair atlateral margins, and posterior pair at posterior margin of prostomium

(Fig. 3.10A). Median antenna with large, pigmented ceratophore; style long, cirriform, basally pigmented,

with numerous long papillae and filiform tip. Ceratophores of lateral antennae heavily pigmented; styles less

than one half length of median antenna, slender, acuminate, with numerous papillae and filiform tips. Palps_

u

slightly longer than median antenna, terete, minutely papillated and tapering to f'me tips.

Tentaculophores lateral to palps, with 1-3 large curved setae projecting from base; subequal tentacular

cirri similar in length and shape to median antenna, with numerous long papillae. Facial tubercle small,

rounded. Anterior dorsal margin of buccal segment produced forward as small nuchal fold. Ventral buccal im

cirri shorter than tentacular cirri, with small papillae, tapering to ffliform tips. •

Elytra large, strongly imbricated, circular (first pair) or reniform (following pairs); each thin,

translucent, with thick lateral fringe and scattered long surface papillae; median and posterior surface divided •

into large polygonal ceils, many with small to large, bulbous, multipronged macrotubercle in center; some II

cells pigmented dark brown, but most colorless (Fig. 3.10B).

Notopodia short, tapering into acicular lobe with pointed cirrus. Neuropodia longer, broader, extending •

into thick, pointed presetal acicular lobe, and shorter, diagonally truncate postsetal lobe; ventral margin proximal l

to ventral cirrophore rngose (Fig. 3.10C). Notosetae stout, numerous, with long spinous regions tapering to

blunt tips (Fig. 3.10D). Neurosetae slightly more slender than notosetae, with long subdistat spinous region I

tapering to long, bare hooked tip with remote incision forming small secondary tooth (Fig. 3.10E). I

Dorsal cirrophores long, tapering; styles cirriform, extending well beyond neurosetae, with numerous

long papillae, broad basal band of dark pigment, and f'fliform tips. Dorsal tubercles well-developed, knoblike. IZ

130
..i

•. .[I

. ....

• ...._

!

....._

t

I

i

I Figure 3,10. Harmothoe hirsuta: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous

parapodium, anterior view; D, notoseta; E, tip of median neurose_

I

I Ventral cirrophores small, bulbous; styles short, acuminate, with scattered short papillae along dorsal margin.

Nephridial papillae short, bulbous, beginning on segment 6, inconspicuous on posterior segments. Pygidial

cirri twice as long as dorsal cirri, with numerous long papillae. I

i 131
I

Habitat. This is a shallow-water species, normally found in the rocky intertidal and shallow subtidal I

areas.

Remarks. The setae, cirri, and elytra of H. hirsuta are usually coated with sediment and debris, II

giving the organism a very shaggy appearance.

Essenberg (1918) noted that this species had been reported from Icy Strait, Alaska, and from Port

Townsend, Washington. These specimens were located in the collections of the Academy of Natural Sciences 1

in Philadelphia, and were found to be one example of Gaudichaudius iphionelloides (Johnson, 1901) [ANSP

2027], and the missing holotype ofEunoe barbata Moore, 1910 [ANSP 2028] (Spamer & Bogan, 1991).

Type Locality and Type Specimens. California, San Pedro, atthe low water mark, holotype (LACM- 1

AHF Poly 0017). m

Distribution. Known from southem California and Panama, in intertidal regions out to 98 m. • ,

Harmothoe imbricata (Linnaeus, 1767)

im

Figure 3.11 A-E I

Harmothoe imbricata: Johnson, 1897:181, fig. 37.--Izuka, 1912:43-49, pl. 5, figs. 1-4, pl. 6, fig. 1.-- I

Fauvel, 1923:55, fig. 18f-l.--Berkeley and Berkeley, 1948:11, fig. 9.--Pettibone, 1953:32-36, pl. I

13, figs. 114-20, pl. 14, figs. 121-31; pl. 15, figs. 132-37; pl. 16, figs. 138-46 (synonymy); 1963:36-

u

38, fig. 7a-d.--Hartman, 1968:79-80, figs. 1-4.--Blake, 1975a:29; 1975b:176, pl. 28, figs. 76- ,1

77.--Uschakov, 1982:165-166, pl. 59, figs. 1-5 (synonymy).--Tebble and Chambers, 1982:30-31, I[

figs. 7a, 9a-b, 31.

m

Material Examined. Alaska, Boca de Quadra (3).--Washington, Lopez Island (2).--California, 1

Tomales Bay (4). m

Description. Length to 65 mm for 39 segments; dorsum usually mottled, with darker areas around II

cirrophores and elytrophores; ventrum dusky or colorless. Prostomium wider than long, with prominent 1

cephalic peaks (Fig. 3.11A). Two pairs of large eyes; anterior pair located forward beneath cephalic peaks

but usually visible through prostomium; posterior pair slightly smaller, near posterior comers. Median antenna I1

with large, pigmented ceratophore; long style with subterminal swelling, scattered papillae, and filiform tip. 1

Lateral ceratophores short, inserted ventrally on prostomium; styles short, acuminate, with scattered papillae

Im

and filamentous tips. Palps to 5 times length of prostomium, tapering, with longitudinal rows of small m

papillae and short tips. 1

Basal lobes of tentacular cirri elongate, with 1-2 setae projecting from base. Two pairs of subequal

tentacular cirri simitar in length and shape to median antenna. Facial tubercle andnuchal fold absent. Ventral •

buccal cirri with large cirrophores; styles shorter than tentacular cirri, papillate, tapering to f'fliform tips. I

Elytra large, thick, suboval to subreniform, strongly imbricated and covering dorsum, but easily

detached. Elytral surface with numerous blunt microtubercles, scattered surface papillae, and, in larger •

specimens, globular macrotubercles near posterior margin (Fig. 3.11B); lateral and posterior borders with II

fringe of short papillae, or fringe absent.

Notopodia rounded, tapering into pointed acicular lobe; neuropodia longer, broader, extending into I

thick presetal lobe with emergent acicula (Fig. 3.11C). Notosetae stout, moderately thick, with long spinous

regions tapering to blunt, bare tips (Fig. 3.11D). Neurosetae slightly more slender than notosetae, with short,

expanded, subdistal spinous region tapering to long, bare hooked tip with or without small secondary tooth I

(Fig. 3.11E).

!

132 I
I

I

I

I

I e ":__'i.... "''_............ "_.i_-'i;' cph

,,

|' _" E

I

Figure 3.11. Harmothoe imbricata: A, anterior end, dorsal view; B, median right elytron with detail of

microtubereles and marginal papillae; C, median cirrigerous parapodium, anterior view; D, distal

I portion of median notoseta; E, distal portion of median neuroseta.

I

i 133
!

Dorsal cirri with long, cylindrical cirrophores; styles basally pigmented, papillate, with subdistal l

swelling and filiform tips extending well beyond neurosetae. Dorsal tubercles prominent, bluntly conical.

Ventral cirri short, acuminate, with numerous small papillae. Cylindrical nephridiat papillae long, projecting

between parapodia from segment 6. Terminal pair of pygidial cirri up to twice dorsal cirri length. 1

li

Biology. Harmothoe imbricata is a successful cosmopolitan found throughout the arctic and in

boreal seas. The species utilizes a wide variety of habitats, and is found intertidally under rocks and in am

eelgrass beds, and subtidalty on rocky, muddy, or sandy substrates, in kelp holdfasts, mussel beds and old 1

Sabellaria reefs. It is an active swimmer and can tolerate a wide range of salinities. It is found both free- II

living and in commensaI association with echinoderms and other polychaetes. Blake (1975a) recorded egg Ill

diameters of 120-123 _ in specimens from Tomales Bay. The eggs are brooded under the elytra throughout •

much of the year, and after hatching, the California larvae of this species have a prolonged pelagic life before B

settling and metamorphosis. In other oceans, however, H. imbricata has been found to exhibit a full range of

developmental types from direct development under the elytra to direct spawning into seawater (Blake, 1975a). 1

This plasticity in mode of reproduction, coupled with the apparently diverse habitats in which the species is 1

able to live, undoubtedly contribute toward making H. imbricata one of the most widely-distributed species of

polynoids. I

Remarks. The elytra ofH. imbricata exhibit great variations both in pigment pattern and in coloration.

The first pair are often colorless, and following pairs have various mottled or solid patterns in shades ranging Ill

from white, light tan, red, green, brown, grey, and black, l

II

The secondary tooth on the neurosetae is usually well-developed in the middle portion of the fascicle,

and is often broken, worn, or not developed on the upper and lower neurosetae. Very few bidentate setae are •

present injuvenile or subadult specimens, and most of the neurosetae appear unidentate. I

Type Locality and Type Specimens. The species currently accepted as Harmothoe imbricata was

described by Linnaeus in 1767 based upon material sent from Iceland, although it is unclear whether this •

material was an actual specimen or merely a drawing (Chambers and Heppell, 1989). No type specimen is I

known to exist.

Distribution. Widespread throughout the northern hemisphere, extending down to the Mediterranean n

and to New Jersey in the Atlantic, and from the Yellow Sea around the Pacific rim to southern California. It

is abundant in the intertidal and shallow subtidal, but is also found out to abyssal depths.

Harmothoe multisetosa (Moore, 1902) I!

Figure 3.12 A-F l

l

Lagisca multisetosa Moore, 1902:267-69, pl. 14, figs. 29-36.--Berke|ey and Berkeley, 1945"322; 1948:15-

16, fig. 17.--Hartman 1968"105-106, figs. 1-6. •

Harmothoe multisetosa: Moore, 1910:340-41 .--Pettibone, 1953"29-31, pl. 12, figs. 104-113 (synonymy). l

Material Examined. Alaska,Icy Cape(holotype).--Washington, off SanJuanIsland (9).--California, •

Santa MariaBasin, off Point Buchon, Sta. 6, 109 m (1), off Purisima Point, Sta. BRA-13, 89-101 m (1). I

Description. Length to 35 mm for 43 segments; dorsum dark brown with two thin transverse white

bands per segment converging at elytrophores or dorsal tubercles; ventral surface colorless, iridescent. •

Prostomium wider than long, with prominent cephalic peaks (Fig 3.12A). Two pairs of large eyes; anterior

pair slightly larger, located near lateral borders at widest point of prostomium. Median antenna with long,

tapering, pigmented ceratophore; style more than twice prostomium length, with numerous scattered papillae, II

subterminal swelling, and long filiform tip. Lateral ceratophores short, inserted beneath cephalic peaks; I

styles about length of prostomium, acuminate, with scattered papillae and filiform tips. Palps longer than

median antenna, tapering to filiforrn tips, with minute papillae. •

I

,34 I
Figure 3.12. Harmothoe muItisetosa:A, anteriorend,dorsalview; B, median right elytron;C, mediancirrigerous

parapodium, posterior view; D, notoseta; E, tip of median neuroseta; F, tip of inferior neuroseta.

!

I Upper pair Basalof lobes tentacular of tentacu cirrilarsimilar cirri with in length digitateandacicular shape tolobemedian and 1-3antenna; stout, curved lower pair setaeabout on anterior 3⁄4 as long. face.

Facial tubercle absent. Ventral buccal cirri with long, cylindrical cirrophores; styles similar to lower tentacular

cirri.

Large suboval to subreniform elytra covering dorsum except for posteriormost segments. Elytra thin,

translucent, with blunt or bifid microtubercles, numerous larger thorn-like curved spines with acutely pointed

I tips, fringeandof occasional short papillaelarge, (Fig.blister-like 3.12B). Elytral macrotubercles surface uniformly near posterior tan orborder; gray, orlateral mottledandwithposterior brownishborders pigment; with

macrotubercles darker brown.

I Notopodia obliquely truncate, with inferior acutely pointed acicular lobe; neuropodia longer, extending into pointed presetal lobe with protruding acicula and long digitiform superior cirrus (Fig. 3.12C). Notosetae

stout, curved, with transverse rows of spinules and blunt to pointed smooth tips (Fig. 3.12D). Neurosetae

about as thick as notosetae, enlarged subdistally; superior neurosetae with long subdistal spinous region

i 135
!

tapering to smooth points; median neurosetae with shorter spinous region, hooked smooth tip, and small I

secondary tooth (Fig. 3.12E); inferior neurosetae shorter, unidentate (Fig. 3.12F).

Dorsal cirri extending beyond neurosetae, with long, tapering cirrophores, and styles similar to median _l

antenna. Dorsal tubercles knoblike, often with brown pigment. Ventral cirri short, basally inflated, tapering

to fdiform tip, with occasional small papillae. Bulbous nephridial papillae short, projecting between parapodia

from segment 6. Terminal pair of pygidial cirri similar in shape and longer than dorsal cirri. 1

Habitat. This species is common in intertidal areas on pilings, and across the continental shelf out to

250 m in regions with rocks, algae, barnacles, or mollusc shells.

Remarks. Although the dark brown dorsal pigmentation may fade over time in preservative, the thin !

transverse white stripes on each segment remain visible.

A number of authors (Ditlevsen, 1917; Augener, 1928; Annenkova, 1937; Wesenberg-Lund, 1950) II

have synonymized Harmothoe multisetosa with H. aspera (Hansen, 1878) from arctic and boreal waters. I

However, the elytra of the latter species lack both the bifid microtubercles and the large, soft macrotubercles

found in H. multisetosa. These structures were observed in the California material, and Moore's species is •

regarded as distinct pending further investigation. II

Type Locality and Type Specimens. The original assignment of the type locality to McCormick

Bay in western Greenland was subsequently corrected to Icy Cape, Alaska (Moore, 1905), holotype (ANSP _

AD267 and slide 3.7).

Distribution. Known along the Pacific coast of North America from Alaska to Mexico.

Genus Hesperonoe Chambeflin, 1919 •

Type Species: Harmothoe complanata Johnson, 1901, by monotypy. I

Diagnosis. Body short, flattened, with about 40 segments. Bilobed prostomium with cephalic peaks; I

ceratophores of lateral antennae inserted beneath anterior margin. Facial tuberele and nuchal fold absent. I

Fifteen pairs of elytra on segments 2, 4, 5, 7.... 23, 26, 29, and 32, completely covering dorsum. Parapodia

birathous. Upper notosetae thicker than neurosetae, with transverse striations and blunt tips; lower notosetae 1

longer, more slender, tapering to capillary tips. Superior neurosetae slender, with long spinous region tapering

to fine tips; inferior neurosetae thicker, with expanded subdistal spinous region and smooth, slightly hooked

tips. I

Remarks. Hesperonoe is similar to the genus Harmothoe, but it differs in having two different kinds

of setae in both the notopodia and neuropodia. Three species, Hesperonoe laevis Hartman, 1961, H. complanata

(Johnson, 1901), and H. adventor (Skogsberg, 1928), occur off California. 1

Hesperonoe Iaevis Hartman, 1961 I

Figure 3.13 A-F

Hesperonoe Iaevis Hartman, 1961:47-48, pl. 1, figs. 1-6; 1968:95-96, figs. 1-5. I

Material Examined. Washington, Puget Sound (1).--Oregon, off Coos Bay (2).---California, Santa

Mafia Basin, off Point Sal, Sta. 38, 197 m (1), Sta. 40, 392 m (1); off Santa Barbara (syntype). _l

Description. Length to 40 mm for 38 segments; dorsum pale. Bilobed prostomium about as long as

wide, dusky gray, produced anteriorly into cephalic peaks (Figs. 3.13A). Anterior pair of eyes located near

widest point of prostomium; posterior pair smaller, closer together. Ceratophore of median antenna large, I
I Figure 3.13. Hesp C, superior eronoe notoseta; Iaevis: AD, ,anterio inferior r endnotoseta; , dorsalE,viedistal w; B,portion medianofcirrigerous superior neuroseta; parapodiumF,,distal anterior portionviewof;

inferior neuroseta.

I

extending beyond cephalic peaks; style about twice prostomium length, tapering to filiform tip, with tiny

I scattered papillae. Lateral cemtophores small, inserted beneath cephalic peaks; styles tapering, half as long

as median antenna. Palps three times prostomial length, tapering to f'fliform tips, with minute papillae.

Basal lobes of tentacular cirri large, with digitiform acicular lobe. Dorsal pair of tentacular cirri

I evenly tapering, reaching to palps, papillae; pair slightly

end of with small scattered ventral shorter. Facial

tubercle absent. Cylindrical cirrophores of ventral buccal cirri inserted at base of first parapodia; styles long,

tapering, similar to ventral tentacular cirri.

I Elytra large, subcircular to subreniform, completely covering dorsum. Each elytron thin, smooth

except for inconspicuous microtubercles anterior to attachment scar, pale or with white, gray, or reddishi brown mNotopodial ottling; margins lobe nearly with occasi as large onal asdigitate neuropodiappillinae.first segment; thereafter notopodia small, tapering to

digitate lobe with emergent acicula. Neuropodia larger, longer, extending into short digitate lobe above

I protruding acicula (Fig. 3.13B). Superior and median notosetae moderately thick, tapering to blunt points,

i 137
!

with fine, scarcely discemable transverse striations (Fig. 3.13C); inferior notosetae thinner, longer, tapering •

to fine points (Fig. 3.13D). Upper neurosetae thin, with long, coarsely serrated region, tapering to smooth,

sharp tips (Fig. 3.13E); neurosetae in median and lower portion of fascicle thicker, with subterminal swelling,

shorter serrated region, and pointed or sharp tips (Fig. 3.13F). i

II Dorsal cirri with large, elongated cirrophores; long tapering styles extending beyond neurosetae, with

scattered clavate papillae. Dorsal tubercles welI-devetoped, projecting laterally. Acuminate ventral cirri

smooth. Digitiform nephridial papillae present from segment 6, continuing to end of body. Pygidium low,

broad; pygidial cirri nearly twice as long as dorsal cirri. I

Habitat. Found in muddy sediments from 20-410 m in association with the echiuran Listriolobus

pelodes. I

Remarks. Two additional species of Hesperonoe occur in California, but both are restricted to

intertidal areas. H. complanata is bright yellowish orange in life and is commensal with the ghost shrimp I

Callianassa californiensis. H. adventor is grayish-green and is found in the burrows of the echiuran Urechis !

caupo.

Type Locality and Type Specimens. California, off Santa Barbara, 42 m, holotype (LACM-AHF •

Poly 0026) and numerous paratypes (LACM-AHF Poly 0027). i

Distribution. Known from Puget Sound to southern California.

ll

Genus Lepidasthenia Malmgren, 1867

Type Species: Polynoe elegans Grube, 1840, by monotypy. I

i

Diagnosis. Body elongate, flattened, with more than 100 segments. Bilobed prostomium extending _ i

anteriorly into ceratophores of terminally-attached lateral antennae; antennae and cirri smooth. Facial tubercle • i

present; buccal segment with or without nuchal fold. Elytra on segments 2, 4, 5, 7 .... 23, 26, 29, 32 (or 31),

34, 37, normally continuing on every third segment to end of body. Parapodia subbiramous; well-developed

neuropodium with dorsal cleft between pre- and postsetal lobes. When present, notosetae short, slender,

blunt; neurosetae stouter, longer, with subdistal spinous region and uni- or bidentate tips.

Remarks. The genus Lepidasthenia includes approximately 40 valid species, most of which are !

tropical or subtropical in distribution. Members of the genus apparently continue to add segments throughout i

their lives, and individuals with nearly 200 setigers have been reported. Four species have been reported from

California, two of which were found in the Santa Maria Basin material. !

Lepidasthenia berkeleyae Pettibone, 1948

Figure 3.14 A-E

Lepidasthenia berkeleyae Pettibone, 1948:413-416, fig. 2a-f; 1953:53-54, pl. 26, figs. 237-239.--Lie,

1968:368-369.

Lepidasthenia interrupta: Hartman, 1968:l15-116.---Blake, 1975b:176, pl. 27, fig. 71. Not Marenzeller,

1902. !

Material Examined. British Columbia, Deep Bay, topotype ofL. longicirrata.--Washington, Puget

Sound (holotype, 2 paratypes, and 3 additional specimens).---California, off Point Arena (1); Tomales Bay

(3); Santa Maria Basin, off Point San Luis, Sta. 21, 49 m (1), Sta. 23, 99 m (1), off Purisima Point, Sta. R- e

5, 154 m (1).

!
I

!

B

I D E

I _L _)h __----__/dC

'I A -__..< -_:,_

rlo__. . •

I

,

!

i Figure 3.14. parapodiuLePm,idastheniaanteri D,anteri Csube,pedc oroenrrkeleyae r dn,euro dorsal s:viAeta;view; ,ew; E, distal B, median portioleft n ofelytron; median neurosetm ediana.cirrigerous

I

Description. Length to 75 mm for 100 segments; dorsum colorless or with wide transverse bands of

I Prostomiu_ mPigment' '_"_'" ven"antrumunpigmentedth long, projecting anteriorly into ceratophores cf lateral antenna (Fig. 3.14A). Four

and darker pigmentat bases of parapodia, cirropbores, and elytrophores;

large eyes; antericr pair rear widest part of prostomium; posterior pair closer together and near posterior

i sm,aoothm rgin. withMcemtoph ;ligh edant iantesunnaobrdis ewith s tal sho swelling rt and fihformStylupfiptolengthL ,athre terale tantennae imes prostomial less than twiceslen cylindricalp 'mstomial der' length,

similar in shape to median antenna. Palps long, subulate, without pigment or papillae.

I Two pairTsenSsub tacul°ph° equaltreneoftsimilarl acular arge' laciterrrial to palps,lengaC in thhaband etousUtshapWi IOtobdigitiform tehmedianantenna acicula . Fracial tubercle bulbous°n; anteri°r face.

nuchal fold lacking. Ventral buccal cirri long, slender, with filiform tip and slight subdistal swelling.

I minute sensory papillaSU b°val elytra eleavinal gongouter edgemed; ian d°rssUmrface smoU oth,_ c°vered'tacking tu_ercleE ach elytron thinor s papillaetran ,slucenwt,ithwithdarkOccasiOnpi algment

concentrated around elytrophore and extending in band toward middorsum (Fig. 3.1413).

!

139
!

Parapodia subbiramous. Notopodia short, digitiform, not reaching to base of neuropodial lobes; !

neuropodia long, slender, with rounded pre- and postsetal lobes separated by deep dorsal cleft (Fig. 3.14C).

Superior neurosetae slender, with long subdistal spinous region tapering to fine knobbed tip (Fig. 3.14D). •

Median and inferior neurosetae stouter, with short spinous region tapering to blunt bifid tip (Fig. 3.14E).

Dorsal cirrophores basally bulbous; styles smooth, slender, similar to median antenna in anterior

segments, and tapering evenly to filiform tips in median and posterior segments. Dorsal tubercles poorly 1

developed. Ventral cirri short, smooth, subulate. Ventral nephridial papillae small, short, from segment 6.

Pygidium with terminal pair of acuminate cirri similar to dorsal cirri.

Biology. Lepidasthenia berkeleyae has been reported as a commensal with Praxillella pacifica and i

Maldanella robusta, living in the thick mud tubes of these maldanids. Generally, this species is found in areas

me

with muddy bottoms, in water depths from 49-480 m. ,HaL

Remarks. Hartman (1968:115) recognized Lepidasthenia interrupta, originally described by I

Marenzeller (1902) from off Japan, as the California species with thin superior neurosetae and no ventral

i

neuropodial papillae. However, she incorrectly inserted Marenzeller's drawing of the anterior end of I

Lepidonotus elongatus rather than his sketch of the parapodium ofLepidasthenia interrupta as her figure 2. •

Although Marenzeller does not mention it in his text, this drawing of the 23rd parapodium clearly shows a

u

papilla on the ventral margin of the neuropodium. Therefore, the California specimens that lack these papillae am

cannot be the same as Marenzeller's species. H

The secondary tooth on the median and inferior neurosetae is often screened by the subterminal spines

which extend nearly to the tip of each seta. •

Type Locality and Type Specimens. Washington, Puget Sound, off Green Point, 92 m, holotype I

(USNM 21099, specimen dried); near Gertrude Island, 126 m, paratype (USNM 21100); off Heron Island, 63

m, paratype (USNM 32293, gravid female). [I

Di el stribution. Reported from British Columbia to southern California.

Lepidasthenia longicirrata Berkeley, 1923 1

Figure 3.15 A-E

Illl

Lepidasthenia longicirrata Berkeley, 1923:214-15, pl. 1, figs. 8-13.--Pettibone, 1953:51-53, pl. 26, figs. 1

229-236.--Hartman, 1968:117-118, figs. 1-5.--Uschakov, 1982:89-90, pl. 22, figs. 4-6.

Lepidametria longicirrata: Berkeley and Berkeley, 1948:18-19, fig. 22. u!

Material Examined. British Columbia, off Jesse Island (holotype); off Port Albion (topotype).--

Washington: Puget Sound (l).----California, Santa Maria Basin, off Purisima Point, Sta. R-4, 92 m (1), Sta. •

BRA-21, 75-90 m (1), off Point Arguello, Sta. 70, 200 m (1). II

Description. Length to 120 mm for 118 segments; dorsum with wide bands of light brown pigment,

and darker pigment at bases of parapodia, cirrophores, and elytrophores; ventrum unpigmented, iridescent, i

Prostomium wider than long, bilobed, extending into ceratophores of lateral antennae, with four moderatesized eyes (Fig. 3.15A). Ceratophore of median antenna small; style long, smooth, cylindrical, with fitiform

tip and pigmented subterminal swelling; lateral antennae shorter, similar in shape. Palps long, thick, tapering •

to filamentous tips, without pigment or papillae.

Tentaculophores large, lateral to palps, achaetous but with digitiform acicular lobes. Two pairs of

smooth, slender tentacular cirri; upper pair similar in shape and slightly longer than median antenna. Small •

papilla present beneath each palp and lateral to bulbous facial tubercle. Nuchal fold absent. Ventral buccal

cirri long, slender, tapering.

!

14o I
I Figure 3.15. Lepidasthenia view; C, superior longicirrata neuroseta; :A, D, anterior distalend, portion dorsalofview; median B, anterior neurseta; cirrigerous E, distal parapodium, portion ofposterior inferior

neuroseta.

!

Large suboval ely_a nearly covering dorsum; each elytmn thin, translucent, smooth, lacking tubercles

I or marginal papillae, with brown pigment especially around elytrophore.

Parapodia subbiramous. Notopodia digitiform, elongated and reaching nearly to base of neuropodial

lobes; neuropodia long, slender, with dorsal cleft between elongate, rounded presetal lobe and slightly shorter

I postsetal lobe (Fig. 3.15B). Proximal ventral with of short,

margin of neuropodia fringe squat papillae having

terminal cilia; similar fringe also present dorsally between notopodia and elytrophore or cirrophore on anterior

segments. Superior neurosetae long, slender, with long subdistal spinous region tapering to fine knobbed tip

(Fig. 3.15C); median neurosetae stouter, with short spinous region tapering to bare bifid tip with distinct secondary tooth (Fig. 3.15D); inferior neurosetae shorter, slender, curved, minutely bidentate (Fig. 3.15E).

Dorsal cirri slender, long in anterior region with slight subdistal swelling, and shorter, acuminate in

i posterior setigers. Dorsal tubercles not developed. Ventral cirri short, subulate, smooth. Ventral nephridial

papillae small, short, from segment 6. Terminal pair of pygidial cirri slender, resembling posterior dorsal

cirri.

!

I 141
i

|

Biology. Lepidasthenia Iongicirrata is normally found free-riving in muds, among rocks, or in shelly •

substrata, from the intertidal to 330 m. It is also sometimes found associated with parchment-like, sand and

shell-covered tubes which the worm apparently constructs, i

Remarks. The fringe of papillae on the proximal ventral margin of the neuropodia is easily abraded,

and is not always present on all segments. The similar row on the dorsal margin has not been previously

reported in the literature; this feature is apparently restricted to the anterior region, and was not observed I

posterior to segment 21. A single short, blunt notoseta was observed on an anterior parapodium in the small

specimen from Puget Sound.

The holotype consists of a median and posterior fragment, but the anterior end is missing. Therefore, i

Edith Berkeley selected two topotypes from the Berkeley private polychaete collection, and these were transferred w

to the Smithsonian in 1964 (Pettibone, 1967). One of these (USNM 32851) was considered by the Berkeleys

to be the commensal form of L. longicirrata, but it is now generally accepted asL. berkefeyae, i

Ill

Type Locality and Type Specimens. British Columbia, dredged from a sponge bed off Jesse Island

near Nanaimo, 27 m, holotype (USNM 32853); off Port Albion, topotype (USNM 32852). III

Distribution. Known along the North American coast from Vancouver Island to southern California, I

and also reported from the Gulf of Tonkin in the western Pacific (Uschakov, 1982).

Genus Lepidonotus Leach, 1816 I

Type Species: Aphrodita squamata Linnaeus, 1767, designated by Malmgren, 1867. I

Diagnosis. Body short, arched, with 26 segments. Bilobed prostomium extending anteriorly into i

ceratophores of terminally-attached lateral antennae. Antennae and cirri smooth. Facial tubercle present;

buccal segment with or without nuchal fold. Twelve pairs of elytra on segments 2, 4, 5, 7.... 21 and 23;

elytra with or without tubercles and papillae. Notopodia small or vestigial; unidentate notosetae short, slender, i

spinose, or notosetae lacking. Neuropodia large, with or without acicular lobe; neurosetae stout, long, with i

subdistal spines and unidentate or occasionally bidentate tips.

Remarks. The genusLepidonotuscontaJnsover70 species. As currently defined, the generic diagnosis l

is very broad, and several groups will undoubtedly be split away from the genus after revision. Only one

species was seen in the Santa Maria Basin material.

!

Lepidonoms spicul_ (_eadwell, 1906) new combination

Figure 3.16 A-H I

i

Polynoe spicula Treadwell, 1906:1151-1152, fig. I1. i

Lepidonotus caeloris [sic]: Moore, 1908:331 (partita); 1910:333-334 (partim).--Berkeley, 1923:213 (partim). i

Not Moore, 1903. i

Lepidonotus caelorus: Hartman, 1938:108-9, fig 35b-d (partim).--Berkeley and Berkeley, 1948:9-10, fig. 6-

7 (partim).--Pettibone, 1953:15-16, pl. 1, figs 1-8, pl. 2, figs. 9-19 (partim). Not Moore, t903. I

Lepidonotus squamatus: Berkeley and Berkeley, 1954:455-456 (partim).--Khlebovich, 1961:165.--Imajima

and Hartman, 1964:26-27, pl. 1, figs. g-j (partita). Not Linnaeus, 1767.

Material Examined. Washington, Puget Sound (10).--California, Monterey Bay (2); Santa Mafia i

Basin, off Point Buchon, Sta. BRA-27, 96-126 m (19), off Point Arguello, Sta. BRA-6, 54-63 m (44).

I

142 |
I

I

I pa man tC

IAn

I //

_=".FJ_

| '°" F ._i:i

elph :_ \ :ii

._i

. .'.t

G :i!

cph

I

I ...... '_"" '_ _°

.. ::........:::...:.,-,,-.:... _)_J_

I E D

!

Figure 3,1@ LepiC._noms spiculus, new combination: A, anterior end, dorsal view; B, first fight e]y_on; E,

i median left elytron; D, detail of elytral surface from anterior etytron (top group), median elytron,

and posterior elytron; E, median cirrigerous parapodium, anterior view; F, superior notoseta; G,

inferior notoseta; I-I,distal half of median neuroseta.

I

I 143
!

Description. Length to 20 mm for 26 segments; most body surfaces colorless except for dark middorsal i

pigment patch on segment 23. Prostomium slightly wider than long, bilobed with anterior portions projecting

forward as ceratophores of lateral antenna (Fig. 3.16A). Two pairs of dark eyes; anterior pair located at Hi

widest part of prostomium, and slightly smaller posterior pair near posterior margin. Median antenna with II

large, cylindrical, basally pigmented ceratophore; style smooth, with ring of brown pigment around subterminal

enlargement and with long filiform tip; styles of lateral antennae similar but shorter. Palps smooth, colorless, i

longer than median antenna, tapering to short filiform tips. l

Long tentaculophores with 1-2 spinose setae; two pairs of subequal tentacular cirri similar in shape

and pigmentation to median antenna. Facial tubercle small, bulbous. Buccal segment with pair of smooth _l

ventral buccal cirri similar in shape to tentacular cirri; dorsal anterior margin produced forward as small

nuchal fold.

Twelve pairs of tough, leathery, firmly attached elytra, strongly imbricated and completely covering i

dorsum; surface studded with numerous light orange to dark tan-colored tubercles, becoming small and sparse i

under areas of overlap. Anterior elytra suboval, with small to large tubercles centered on polygonal, closely

packed basal plates (Fig. 3.16B); largest tubercles conical with sculptured sides and spinose tips; smaller i

tubercles near anterior and lateral borders columnar with spinose tips. Median elytra subreniform to pyriform, II

with smaller, lower tubercles on more rounded basal plates (Fig. 3.16C). Posterior elytra pyriform to

subtriangular, with numerous tall, acutely conical tubercles on round basal plates (Fig. 3.16D). Lateral and 1

posterior margins with thick fringe of long ffliform papillae. i

Parapodia biramous; notopodial lobe short, rounded, located on anterodorsal surface of neuropodia; I

neuropodia larger, truncate, distally cleft with presetal lobe slightly longer than postsetal lobe (Fig. 3.16E). i

Notosetae spinous; superior row slender, short, curved, with blunt tips (Fig, 3.16F); median and inferior rows

II

slender, longer, tapering to capillary tips extending beyond end of neuropodium (Fig. 3.16G). Neurosetae i

thicker than notosetae, amber colored, with slight subterminal enlargement tapering to long, smooth, slightly !

hooked tips; enlarged region with two rows of bracts becoming progressively larger distally (Fig. 3.16H). i

Dorsal cirri with large, basally inflated cirrophores; styles slender, smooth, extending well beyond i

neurosetae, with pigmented subterminal swelling and long flliform tips. Dorsal tubercles large, well developed, i

Ventral cirri short, subulate, reaching to base of lower neurosetae. Nephridial papillae beginning on segment

8, prominent from segment 10, continuing through next to last segment. Anus dorsal, centered on longitudinally

striated median ridge extending through last three segments. Pygidium with two very long, smooth, slender i

cirri with pigmented subterminal swellings and filiform tips.

1R

Remarks. This species was originally described by Treadwell (1906) in the genus Polynoe, but it I

was subsequently referred to Lepidonotus caelorus Moore, 1903 by Hartman (1938), and most subsequent |

authors have accepted this synonymy. However, as part of her extensive work of the family Polynoidae, Dr.

Marian H. Pettibone at the Smithsonian Institution has reexamined Treadwell's material and noted that the

ornamentation of the elytra is distinctive. The largest anterior tubercles are conical with spiny tips rather than i

bluntly digitiform, and they are situated on polygonal rather than rounded basal plates. In the posterior

I

region, the largest tubercles are acutely conical rather than bluntly digitiform. Dr. Pettibone has graciously Ill

provided of her extensive notes and sketches on this genus, permitting the resurrection of Treadwell's i

some

i

species in this volume.

Type Locality and Type Specimens. California, Monterey Bay, 84-126 m, 3 syntypes (USNM i 5203). I

Distribution. Found from Japan north around the Pacific rim and down the west coast of North

America as far as southern California. I

1

!
!

I Genus Mahngreniella Hartman, 1967

I Type Species: MalmgrenieIla dicirra Hartman, 1967, by original designation.

Diagnosis. Body flattened, with up to 46 segments. Bilobed prostomium with or without cephalic

I lateral ceratophores inserted subterminally, joining beneath median ceratophore. Facial tubercle peaks; present

or absent; dorsal nuchal fold absent. Fifteen pairs of elytra on segments 2, 4, 5, 7.... 23, 26, 29, and 32,

smooth or with microtubercles confined to anterior region. Parapodia biramous. Notosetae stout, as thick or

I thicker than neurosetae, with transverse or longitudinal spinous rows, and blunt or pointed tips. Neurosetae with expanded subdistal spinous region and bare, hooked, bidentate or unidentate tips.

Remarks. Malmgreniella was originally established by Hartman (1967) for a polynoid found living

I commensally with stylasterine hydrocorals in the Antarctic. Recently, Pettibone (1993b) has emended the

genus and expanded it to include many of the species previously referred to Malmgrenia Mclntosh, 1874,

which is now considered indeterminable. The genus is close toHarmothoe Kinberg, 1856, and encompasses

I number of commensal with other invertebrates. It is off southern species a having a

relationship represented

California by a group of closely-related species that are all characterized by having nearly smooth elytra with

microtubercles confined to a small anterior patch, and notosetae with fine spinutes arranged in two longitudinal

I rows. These species have two distinct of body pigmentation. A dark brownish or nearly black pigment

types

occurs on the epithelial surface of the posterior dorsal and ventral body, on the cirrophores and basal portions

of the cirri, and in broad arcs across the elytra. Beneath the surface are reddish-brown pigment granules that

I the posterior portion of the prostomium, scattered the elytra, and occasionally in the parapodia,

occur on across

dorsal and ventral cirri, and around the nephridia. The expression of these two pigments varies widely

between individual specimens, and one or the other of the pigments may attimes be virtually absent. Therefore,

I individuals of the same species may superficially appear to be quite different from one another. The details of

the prostomium, parapodia, and the setae must be assessed in order to correctly identify specimens within this

genus.

I

Malmgreniella baschi Pettibone, 1993

I Figure 3.17 A-E

Eunoe sp. C: Lissner et al., 1986:D-7.

I Harmothoe Harmothoe sp. nr. A: Lissner et al., 1986:D-7. Iunulata: Steinhauer and Imamura, 1990:F-1 (partita). Not delle Chiaje, 1830.

MaImgreniella baschi Pettibone, 1993b:72-75, fig. 47a-i.

I Material Examined. California, Santa Maria Basin, off Point San Luis, Sta. 22, 99 m (1), off Point

Sal, Sta. PJ-1,145 m (2), Sta. PJ-6, 148 m (3), PJ-9, 169 m (2), PJ-13, 144 m (2), off Purisima Point, Sta. RI 4,Santa 92 mCatalina (2); WesIsland tern S(holotype anta Barbara and Channe 2 paratypes); l, off Point off SanConception, Diego (3). Sta. 79, 98 m (1), Sta. 86, 197 (1); off

Description. Length to 14 mm for 38 segments; posterior dorsum and ventrum often with black

I transverse pigment bands. Bilobed prostomium about aslong as wide, with indistinct cephalic peaks and two pairs of black, lensed eyes; anterior pair larger, situated ventrolaterally in front of widest point of prostomium;

posterior pair near posterior margin (Fig. 3.17A). Short, broad median ceratophore inserted in anterior

I prostomi filiform atip. l nCeratophores otch; style about of lateral twice antennae prostomialshort, length, thick,tapering, joining with beneathoccasional median cceratophore; lavate papillaestylesandshort, long

basally stout, tapering to f'lliform tips. Palps smooth, acuminate, as long or longer than median antenna,

I tapering to short f'diform tips.

I 145
I

I

_, IL_°_ ,,i__o._ I

I

.-_. .... _.,o_ ;oo_ _!f I

D E I

, I

PiN 3.17. Malmgreniella baschi: A, anterior end, dorsal view; B, median right elytron; C, median cirrigemus I

I

parapodium, posterior view; D, superior neuroseta; E, distal portion of median neuroseta.

I

Tentaculophores terete, lateral to pmstomium, occasionally pigmented, with 0-2 stout, curved setae

on anterior face. Tentacular cirri similar to median antenna, with basal pigment and scattered papillae; lower I

pair slightly shorter. Facial tubercle bulbous, pigmented. Cirrophores of ventral buccal cirri short, broad; I

styles similar to inferior tentacular cirri.

Fifteen pairs of thin, oval to reniform elytra with occasional micropapillae on lateral and posterior I

I

borders (Fig. 3.17B). Surface smooth except for small area of short rounded microtuberctes anterior to

I

attachment scar on some anterior elytra; surface with or without mottled dark pigment over attachment point

and in irregular C-shaped band along median, posterior, and lateral areas; additional reddish-brown pigment I

granules sometimes present. I

Notopodia short, rounded, tapering into distally pointed acicular lobe (Fig. 3.17C). Neuropodia I

elongated, extending into short, rounded postsetal lobe, and triangular presetal lobe with tapering extension I

above emergent acicula. Curved notosetae more numerous and about as stout as neurosetae, with longitudinal

striations, two faint longitudinal rows of minute spinules, and bluntly pointed tips. Superior neurosetae with

prominent spinules on long spinose region, tapering to pointed tips (Fig. 3.17D); median and inferior neurosetae I

with shorter enlarged spinous region and tess prominent spinules, tapering to bare hooked tips with only

I

indistinct indication of secondary tooth (Fig. 3.17E). I

146 i
I

I Dorsal cirrophores large, cylindrical, with cilia on upper margin and often with pigment on ventral

side; tapering styles extending to tips of neurosetae, with scattered clavate papillae. Prominent dorsal tubercles

I knpairob-like. of long,Ventral taperingcirricirshort, ri. acuminate, with scattered papillae. Pygidium with dorsally-directed anus and

Biology. MaImgreniella baschi is commensal with the ophiuroid Ophiopsila californica, and has

I been collected in sands and in the tubes of Chaetopterus variopedatus (Pettibone, 1993b). Remarks. The shape of the anterior prostomium is somewhat variable, but the lobes are not produced

into acute peaks. This species differs from its congeners mainly in possessing relatively few neurosetae, the

I tips of which are unidentate with only the suggestion of a secondary tooth. On a few specimens, the longest median neuroseta could occasionally be found with a thin secondary tooth.

Type Locality and Type Specimens. California, Santa Catalina Island, holotype (USNM 133581)

I and 7 (USNM 133582-133585). paratypes

Distribution. Found in shelf depths in southern California.

I Malmgreniella macginitiei Pettibone, 1993

i Figure 3.18 A-D

Harmothoe lunulata: Berkeley and Berkeley, 1941:21 (parfim).--Blake, 1975b: 176, figs. 72-74. Not delle

Chiaje, 1830.

Malmgrenia lunulata: Pettibone, 1953:25 (partim), pl. 9, fig. 77, pl. 10, figs. 80-82, 85, 87, pl. 11,figs. 92-

95. Not delle Chiaje, 1830.

Harmothoe crassicirrata: Lissner et al., 1986:D-7. Not Johnson, 1897.

I Harmothoe cf. lunulata: Kudenov, 1975:42, Lissner al., 1986:D-7 Not delle fig. Ia-g; et (partim). Chiaje,

1830.

Malmgreniella macginitiei Pettibone, 1993b:70, fig. 45a-j.

I Material Examined. California, Tomales Bay (holotype and 1 paratype); Santa Maria Basin, off

Point San Luis, Sta. 21, 49 m (1); off Newport Beach (1); off San Diego (7).

I Description. Length to 13mm for 35 segments; posterior dorsum and ventrum usually with transverse

bands of dark pigment. Bilobed prostomium slightly longer than wide, with reddish-brown pigment often

concentrated in median furrow; cephalic peaks acute (Fig. 3.18A). Two pairs of moderate-sized eyes; anterior

I pair point prostomium; slightly posterior pair closer together, located

situated in front of widest of smaller

near posterior margin. Median ceratophore short, broad, pigmented; style less than twice prostomial length,

basally pigmented, with occasional clavate papillae and long filiform tip. Ceratophores of lateral antennae

I short, thick, beneath median short, with scattered and filiform

joining ceratophore; styles tapering, papillae

tips. Stout palps about as long as median antenna, smooth, tapering to short filiform tips.

i Cylindrical tentaculophores lateral to prostomium, pigmented, with 1-2 stout, curved setae on anterior

face. Tentacular cirri basally pigmented, with scattered papillae; upper pair slightly longer than median

antenna, and lower pair somewhat shorter. Facial tubercle as prominent pigmented ridge. Cirrophores of

ventral buccal cirri bulbous, pigmented; styles long, tapering, with scattered clavate papillae.

I Fifteen pairs of thin, strongly imbricated, subcircular to reniform elytra with occasional tiny bordering

papillae (Fig. 3.18B). Elytral surface smooth except for small area of short rounded microtubercles anterior

i tmedian o attachmen and tposte scarrionor sareas ome anandterioover r scal attachment es; surfacepoint; usualladditional y with mottled granules darkofpigment reddish-brown in irregular pigment band aloften ong

present throughout.

I

I 147
I

I

jpa

• )

I

!

' t

microT ''... ".. • ... I

c I

f I

mL

Figure 3.18. Malmgreniellamacginitiei: A, anterior end, dorsal view; B, median left elytron; C, median cirrigerous •

parapodium, anterior view,withdetail of neuropodial supraacicular lobe; D, distal portion of median

neuroseta with detail of tip.

!

Nompodia short, broad, tapering into pointed inferior lobe with emergent acicula (Fig. 3.18C).

Neuropodia elongated, extending into short digitate lobe above emergent acicula. Notosetae numerous, smut, B

curved, tongitudinaIly striated, with two rows of minute spinules and bluntly pointed tips. Neurosetae numerous

I

and slightly more slender; superior neurosetae with pointed or minutely bifid tips; median neurosetae with

shorter enlarged spinous region and bare hooked tips, each with moderate secondary tooth tapering to slender I

tip (Fig. 3.18D); inferiormost neurosetae with slightly hooked tips and reduced or absent secondary teeth.

Distal spinous bracts not approaching base of secondary tooth.

Cirrophores of dorsal cirri large, cylindrical; styles tapering, basally pigmented, with occasional

small ctavate papillae, extending nearly to tips of neurosetae. Dorsal tubercles prominent, knob-like. Ventral

m

cirri short, acuminate, with occasional papillae. Pygidium small, with low lobes surrounding the dorsalIydirected anus; two pygidial cirri similar in size and shape to dorsal cirri. I

Biology. MalmgrenielIa macginitiei is a commensal polynoid that has been collected in the burrows

of the ghost shrimp Callianassa californiensis, in the tubes of the maldanid Axiothella rubrocincta, and on I

the arms of the ophiuroid Amphiodia urtica (Pettibone, 1993b). •

,48 |
!

I Remarks. This species differs from its congeners mainly in possessing acutely pointed cephalic

peaks on the prostomium, and in details of the neurosetae. The tips of the secondary teeth are drawn into long,

I f'me barepanoints, d devaloidthough o these are often broken off. Inaddition, the region just beneath each secondary tooth is f the transverse spinous bracts.

Type Locality and Type Specimens. California, Tomales Bay, intertidal, holotype (USNM 24933),

i 1 paratype (LACM-AHF Poly 1119); Newport Bay, 1 paratype (USNM 55059). Distribution. Washington to southern California, in intertidal and shelf depths.

I Malmgreniella nigralba (Berkeley, 1923)

Figure 3.19 A-E

I Malmgrenia nigralba Berkeley, 1923:213, pl. 1, figs. 5-7; 1924:193.--Hartman, 1968:133, figs. 1-3.

Malmgrenia lunulata: Pettibone, 1953:25 (partita), pl. 11, figs. 96-103. Not delle Chiaje, 1830.

I Harmothoe Harmothoe nr. nigralba lunulata: Lissner et al., 1986:D-7 (partim). Not delle Chiaje, 1830. : Hanley, 1987:153, figs. 3f, h.

MalmgrenieIla nigralba: Pettibone, 1993b:60, figs. 39, 40.

I Material Examined. Alaska, Prince William Sound (1).--British Columbia, near Nanaimo (3

syntypes).---Califomia, Western Santa Barbara Channel, off Point Conception, Sta. 79, 98 m (6); off San

I Diego (5). Description. Length to 24 mm for 40 segments; dorsum and ventrum colorless or with light or dark

pigment bands especially in posterior segments. Prostomium bilobed, about as long as wide, with median

I furrow and truncate anterior lobes (Fig. 3.19A). Two pairs of moderate eyes; slightly larger anterior pair on lateral margin anterior to widest portion of prostomium. Ceratophore of median antenna short, bulbous; style

slightly longer than prostomium, with short clavate papillae, tapering to filiform tip. Ceratophores of lateral

antennae short, broad, inserted subterminally and converging beneath median ceratophore; styles half prostomial length, pyriform, with clavate papillae. Palps about length of median antenna, tapering, wrinkled, with short

tips and minute papillae.

I Tentaculophores large, lateral to prostomium, with or without stout, curved seta on anterior face; dorsal and ventral tentacular cirri subequal, similar to median antenna, with short clavate papillae. Facial

tubercle prominent, often with brownish pigment; nuchal fold absent. Ventral buccal cirrilong, tapering, with

I scattered clavate papillae.

Fifteen pairs of thin, subcircular to subreniform elytra coveting dorsum. First pair and posterior

pairs essentially smooth; median pairs with small area of rounded microtubercles anterior to attachment scar

I and occasional minute marginal papillae (Fig 3.19B). First pair with circle of dark pigment on reticular

background; following pairs with pigment and reticulation generally confined to posterior half of elytral

surface.

I Notopodia small, rounded, tapering into digitate inferior lobe with emergent acicula; neuropodia

larger, tapering to subconical presetal acicular lobe with short, broad, distally rounded supraacicular cirrus

(Fig. 3.19C). Notosetae slightly curved, with longitudinal striations and faint subdistal spines arranged in

i two longitudinal rows, tapering to blunt tips (Fig. 3.19D). Neurosetae similar to notosetae in number and

thickness; superior neurosetae with long subdistal spinous region and rounded or bidentate tips (Fig. 3.19E);

: median neumsetae with shorter subdistal spinousregion, taperingto strongly hooked tip and prominent secondary

I tooth from axis of distal bracts close to base of tooth diverging slightly seta; spinous reaching very secondary

(Fig. 3.19F); inferior neurosetae smaller, bidentate or ending in knobbed tip (Fig. 3.19G).

I

I 149
!

Figure 3.19. Malmgreniella nigralba: A, anterior end, dorsal view; B, posterior fight elytron with detail of I

pigmented reticular cell; C, median cirrigerous parapodium, anterior view; D, notoseta; E, distal

portion of superior neuroseta; E distal portion of median bifid neuroseta; G, distal portion of inferior •

knob-tipped neuroseta. I

,50 I
!

I Dorsal cirrophores large, cylindrical; styles tapering, extending about to tips of neurosetae, with

scattered small clavate papillae. Dorsal tubercles prominent, nodular. Ventral cirri short, with scattered

I papillae, with low tape loberisurrounding ng to filiformdorsally tips. Nephridial directed anus, papilwith lae short, pair ofcyllong, indrictapering al, evidanal ent bycirri. segment 11. P;ygidium

Biology. Malmgreniella nigralba has been found in California, Washington, and British Columbia

I living sands from in thethevertical low intertidal burrowsout of the to approximately holothuroid Leptosynapta 100 m. clarki. It appears to prefer coarse gravelly

Remarks. The elytra of this species are distinctly reticulated although this feature is not always

I obvious if the black pigmentation has faded. The transverse spinous bracts on the neurosetae are present distally to the base of the secondary tooth. This tooth was seen to be much longer in the California specimens

than in material from colder waters further to the north.

I Type Locality and Type Specimens. British Columbia, Vancouver Island, 3

Piper's Lagoon, syntypes

(USNM 32875-32876).

i Distribution. Southern Alaska to southem California, in intertidal and shelf depths.

Malmgreniella scriptoria (Moore, 1910)

I Figure 3.20 A-E

Harmothoe scriptoria Moore, 19t0:344-346. pl. 28, figs. 13-17.--Hartman, 196t:49; 1968:85-86, figs. 1-

|

Malmgrenia lunulata: Pettibone, 1953:25-28 (partita), pl. 9, figs. 73-76, pl. 10, figs. 78-79, 83-84, 86, pl.

11, figs. 88-91. Not delle Chiaje, 1830.

I Harmothoe lunulata: Lissner et al., 1986:D-7 and 1990:F-1

nr. (partim).--Steinhauer Imamura, (partim).

Not delle Chiaje, 1830.

Malmgreniella scriptoria: Pettibone, 1993b:68-69, fig. 44A-K.

I Material Examined. California, Monterey Bay (holotype and 1 additional specimen); Santa Maria

Basin, off Point Sal, Sta. 33, 396 m (2), off Purisima Point, Sta. R-2, 161 m (3), Sta. R-3, 409 m (1), Sta. RI 6, 410 m (1), off Point Arguello, Sta. 74, 201 m (3); Western Santa Barbara Channel, off Point Conception, Sta. 80, 196 m (1).

Description. Length to 22 mm for 40 segments; dorsum dusky brown or colorless. Prostomium

I slightly longer wide, truncate (Fig. 3.20A). pairs of small to

than with median furrow and anterior lobes Two

moderate eyes; slightly larger anterior pair on lateral margin of prostomium near widest point. Reddishbrown pigment granules sometimes present near anterior margin of prostomium and between posterior pair of

I Ceratophore of median antenna short, thick; style acuminate, than

eyes. longer prostomium, basally pigmented,

with minute papillae, tapering to filiform tip. Ceratophores of lateral antennae inserted subterminally, joining

beneath median ceratophore; styles short, tapering, with occasional minute papillae. Palps about length of

I median antenna, tapering, wrinkled, with short tips and minute papillae.

Tentaculophores large, projecting forward, with or without stout, curved seta on anterior face. Upper

pair of tentacular cirri similar to median antenna, with basal pigment and minute papillae; lower pair slightly

I shorter. Facial tubercle prominent, often with brownish pigment. Nuchal fold absent. Cirrophores of ventral

buccal cirri large, directed forward; styles long, tapering, with scattered papillae.

i minute marginal Elytra covpapillae ering dorsurn, easily detached, subcircular to reniform. Elytra thin, delicate, with occasional (Fig. 3.20B); surface smooth except for small area of rounded microtubercles anterior

to attachment scar, with crescent of brownish pigment near posterior margin and around attachment scar, or

i pigment absent.

i 151
!

I

./ i I

.v

A _ E

_:,_:,/_ dc\ sopAc, D I

/ I

• ,_-_:_-:b:.£i -.=_,_.:.,.

•-':'.""'i_.i' i!:-_: I

\ ' !

I

Figure 3,20. Mafmgreniella scriptoria: A, anterior end, dorsal view; B, median right etytron with detail of

surface; lobe; D, Cnotoseta; , medianE,cirrigerous distal portion parapodi of median um, anneuroseta terior view,withwithdetail detailofoftip. neuropodial supraacicular I

II

Notopodia low, rounded, tapering into digitate inferior lobe with long emergent acicula; neuropodia I

prominent, triangular, tapering to presetal lobe with short cirrus above emergent acicula; pre- and postsetal

II

margins often pigmented (Fig. 3.20C). Notosetae stout, slightly curved, tapering to blunt tips, with longitudinal

striations and faint subdistal spines arranged in two longitudinal rows (Fig. 3.20D). Neurosetae more numerous I

I

and slightly more slender than notosetae; upper neurosetae with long subdistal spinous region and knobbed or

I

bidentate tips; median and lower neurosetae with shorter subdistal spinous region, tapering to strongly hooked

tip and prominent, short secondary tooth (Fig. 3.20E). Distal spinous bracts reaching very close to base of I

secondary tooth.

Dorsal cirrophores large, tapering; styles acuminate, extending to or beyond tips of neurosetae, with II

scattered small papillae and granules of brownish pigment. Dorsal tubercles prominent, pigmented. Ventral I

cirri short, with pigment granules basally, tapering to filiform tip. Nephridial papillae short, cylindrical,

m

present by segment 9. Pygidium with low lobe surrounding dorsal anus; pygidial cirri similar to and slightly I

longer than dorsal cirri. •
!

I Biology. Found in the ambulacral grooves of the heart urchin Brisaster latifrons, in middle shelf to

upper slope depths in areas of green muds having a sand or gravel component.

I rather thanRemarks smooth. antennae Examination and cirri of theastype illustrated material by confn'ms Moore (19that t0:Malmgreniella figs. 13-14). Thescrspecimen iptoria hasinpapill the U.S. ate

National Museum (USNM 17156) is a gravid female in which the numerous eggs have distorted the parapodia,

I cawhite usingmarkings" the dorsal(Moore tubercles1910) to appear are clearly less conspicuo visible inusthethanelytra normal. of theThespecimen "close and at thecuriAcademy ous fine pencil-like of Natural

Sciences of Philadelphia, and appear to be regions devoid of any subsurface pigment granules.

I Type Locality and Type Specimens. Offthe Point Pifios Light, Monterey Bay, 101-305 m, holotype (USNM 17156 and ANSP AD4602X slide 6.7). Moore (1910) examined two specimens from Monterey Bay,

and he made an error in his manuscript when he designated the specimen fromAlbatross Station 4452 as the

I type Moore's(p. 346). handwriti Thatng specimen stating it toisbeinathe"cotype." Philadelphia The specimen collectionfrom(ANSP Albatross AD2893) Stationwith4460an ininternal the collections label in

of the U.S. National Museum is a gravid female and was indicated by Moore to be the type on page 344 of his

I manuscript. Loi (1980) and Pettibone (1993b) have recognized this specimen (USNM 17156) as the holotype. Distribution. Known from Puget Sound to southern California.

I Genus Subadyte Pettibone, 1969

I Type Species: Polynoe pellucida Ehlers, 1864, by original designation.

Diagnosis. Body elongate, flattened, with about 40 segments. Bilobed prostomium with or without

I cephalic peaks; lateral ceratophores inserted ventrally. Antennae, palps, and cirri with papillae. Facial

tubercle present or absent; buccal segment with small nuchal fold. Fifteen or sixteen pairs of elytra on

segments 2, 4, 5, 7.... 23, 26, 29, 32, and occasionally 34; elytra without microtubercles but with vesicular

I papillae. Parapodia biramous. Notosetae as thick or thicker than neurosetae, curved, with spinous pockets and blunt unidentate or bidentate tips. Neurosetae longer, with distinct basal cusp, spinose subdistal region,

and unidentate or bidentate tips.

I Remarks. Pettibone (1969) erected the genusSubadyte for a group of short-bodied species previously

included inScalisetosus Mclntosh, 1885. Members of this genus are distinguished by the presence of spinous

pockets along the notosetae, and neurosetae with a large cusp at the base of the spinous region. Currently

I there about 7 known, several described from Australian and

are species including recently waters (Hanley

Burke, 1990; 1991). A single species was recognized in the Santa Maria Basin material.

I Subadyte mexicana Fauchald, 1972

Figure 3.21 A-G

I Subadyte mexicana Fauchald, 1972:27-29, pl. 1figs. a-e.--Steinhauer and Imamura, 1990:F-1 .--Pettibone,

1993c:685-687, figs. 4-5.

I Subadyte sp. A: Lissner et al., 1986:D-7.--Blake et al., 1988:A-3.--Steinhauer and Imamura, 1990:F-1. Subadyte sp. B: Lissner et al., 1986:D-7.

I 91 m Material Examined. California, off Point Buchon, Sta. 10, 690 m (1), off Point San Luis, Sta. R- 1, (3), Sta. R-2, 161 m (1), Sta. R-3, 409 m (2), off Point Sal, St,-:.41,495 m (1), Sta. R-8, 90 m (7), Sta.

R-9, 410 m (24), Sta. PJ-1,145 m (5), Sta. PJ-7, 123 m (4), Sta. PJ-8, 142 m (3), Sta. PJ-10, 147 m (7), Sta.

I PJ-14, 134 m (1), Sta. PJ-15, 155 m (2), Sta. PJ-16, 130 m (2), Sta. PJ-23, 143 m (3), off Purisima Point, Sta.

i 153
I

Figure 3.21. Subadyte mexicana: A, anteriorend, dorsal view; B, median right elytron with detail of marginal "

papillae; C, median cimgerous parapodium, anterior view; D, notoseta; E, superiorneuroseta; F,

median neuroseta; G, inferior neuroseta. I

II

46, 597 m (1), Sta. R-4, 92 m (14), Sta. R-5,154 m (8), Sta. R-6, 410 m (57), off Point Arguello, Sta. 65, 107

m (1), Sta. 72, 401 m (3); off San Diego, Sta. E (1). I

Description. Length to 6.8 mm long for 30 segments, width including parapodia 2.0 mm; dorsum

with dusky pigment tending to concentrate in two longitudinal bands above etytrophores and cirrophores; Ill

ventrum colorless. Prostomium strongly bilobed, wider than long, with distinct cephalic peaks; two pairs of I1

moderate reddish eyes arranged in trapezoidal pattern on posterior half (Fig. 3.21A). Median ceratophore Ill

large, cylindrical, inserted between anterior prostomial lobes; style three times prostomial length, tapering,

with scattered long papillae and long filiform tip. Ceratophores of lateral antennae bulbous, inserted beneath I

cephalic peaks, nearly meeting beneath median ceratophore; styles slender, about length of prostomium,

acuminate, with numerous papillae. Palps long, tapering, with longitudinal rows of minute papillae. Pharynx

with border of soft, distally pointed papillae, 11 in dorsal arc and 11 in ventral; lateral two pairs minute. II

E

154 I
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Tentaculophores lateral to palps; upper lobe long, cylindrical, with emergent acicula; lower lobe

I shorter; 1-2 curved setae with spinous pockets occasionally present on anterior face. Tentacular cirri similar

in shape to median antenna, with scattered papillae; upper cirri distinctly longer than lower pair. Facial

tubercle present, with upper lip produced into prominent longitudinal ridge. Buccal segment with small

I nuchal fold of Ventral buccal cirri with filiform

extending over posterior margin prostomium. tapering, long

tips and scattered papillae.

i Suboval to subreniform elytra on segments 2, 4, 5, 7.... continuing on alternate segments to end of body, strongly imbricated and covering dorsum. Etytra thin, translucent, with scattered papillae on surface

and along lateral and posterior margin (fig 3.21B). Some surface papillae vesicular, enlarged proximally, and

i all with cParapodia lavate tipsbiramous; . notopodia low,rounded, extending into long, slender presetal lobe with emergent

acicula (Fig. 3.21C). Neuropodia longer, broader, tapering into long pointed lobe with terminal emergent

I abecoming cicula. Notosetae progressively thick, csmaller urved; eand ach closer seta witogether th proximal toward smoothblunt, shaftnotandchedmedian tip (Fig. region3.21D). of spinose Neurosetae pockets

numerous, slender, longer than notosetae, with basal cusp proximal to spinous region. Superior neurosetae

i coarsel and unidentate y serrated, tapering to notched tips (Fig. 3.21E); median neurosetae longer, with indistinct serrations tips (Fig. 3.21F); inferior neurosetae with small distinct serrations on short distal region,

tapering to unidentate tips (Fig. 3.21G).

I extendingCeratophores beyond tipsofofdorsal neuroset cirriae.cylindri Dosal cal,tubercles enlarged not basally; developed. styles long, Ventralapering, cirri short, with scattered slender, papill smooth. ae,

Nephridial papillae short, cylindrical, from segment 6. Pygidium rounded, with terminal anus; terminal cirri

I not observed. Biology. Found in sandy and muddy sediments on the continental shelf and slope, and reported from

a whale-fall community in the Santa Catalina Basin (Pettibone, 1993c).

I Remarks. Subadyte mexicana is extremely fragile, and unbroken adult specimens are rarely found.

The eyes are often indistinct, and at times the visual pigment is barely discernable. This is especially true of

the specimens from the deeper stations, indicating that the visual pigments may not fully develop at depth.

I Some specimens exhibit distinct longitudinal bands of pigment along the dorsum, while others appear colorless. In the smaller specimens, a relatively greater proportion of the neurosetae exhibit distal serrations and notched

tips. Presumably the nearly smooth neurosetae with unidentate tips are acquired later as the worm grows

I larger.

Type Locality and Type Specimens. Baja California, 15.5 miles from Natividad Island light, 844

m, holotype (LACM-AHF Poly 1008).

I Distribution. Southern Califomia and Mexico as far south as Cedros Island, in mid-shelf to midslope depths.

I Genus Tenonia Nichols, 1969

I Type Species: Tenonia kitsapensis Nichols, 1969, by monotypy.

Diagnosis. Body short, flattened, widest anteriorly and tapering posteriorly, with up to about 39

I segments. margin. Antennae, Bilobed prostomium palps, and wcirr ithi cwithout ephalic peaks; papillae. ceratoFapchial orestubercle of lateralabsent; antennaebuccinserted al segment beneathwithantesmall rior

nuchal fold. Fifteen pairs of elytra on segments 2, 4, 5, 7.... 23, 26, 29, and 32, completely covering dorsum.

I Par tapering apodiatobiramo capillaryus. tips; All setae inferioorf similar neurosetae thickness; with expanded notosetae subdi and ssutalperior spinous neurosetae region long, and bifid slender, tips. serrated,

Remarks. Tenonia is similar to the genus Harmothoe in which the sole species was originally

I described. pointed. It differs, however, in having notosetae that are slender and capillary-tipped rather than thick and

I 155
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Tenoniapriops (Hartman, 1961)

Figure 3.22 A-E !

Harmothoe priops Hartmart, 1961:50-51; 1966:t93-194, pl. 1,figs. 4-6; 1968:83-84, fig. 1.

Tenonia kitsapensis Nichols, 1969:205-208, fig. 1 A-J. 1

Tenonia priops: Barreca, 1984:801-803.

Material Examined. Washington, off Port Madison (holotype and 15 paratypes of Tenonia n

kitsapensis).--Oregon, off Coos Bay (2).---California, Monterey Bay (1); Santa Maria Basin, off Point Sal,

Sta. R-8, 90 m (2).

Description. Length to 12 mm for 39 segments; dorsum with distinctive wide transverse bands and 1

narrower intersegmental bands of dark reddish brown pigment. Prostomium trapezoidal, bilobed, with weakly- n

developed cephalic peaks and pigment speckles on wide posterior portion (Fig. 3.22A). Two pairs of large

eyes; anterior pair located beneath cephalic peaks and visible through transparent prostomium. Median 1

antenna with large ceratophore inserted in notch between prostomial lobes; smooth, acuminate style pigmented !

basally, about twice prostomial length. Ceratophores of lateral antennae inserted beneath median ceratophore;

smooth pigmented styles short, basally bulbous, tapering to filiform tips. Palps longer than median antenna, 1

thick, tapering abruptly to filiform tips, without papillae.

Tentaculophores short, lateral to palps, without setae or acicular lobe. Two pairs of subequal tentacular m

cirri similar in length and shape to median antenna, with dark pigment basally. Facial tubercle absent. Buecal •

m

segment with small dorsal nuchal fold. Ventral buccal cirri smooth, pigmented, tapering, inserted on short

cirrophores at bases of parapodia. m

Fifteen pairs of circular elytra not covering dorsum in anterior setigers. Elytra thin, translucent, 1

easily detached, nearly smooth except for occasional inconspicuous microtubereles, with brown pigment

around attachment scar; elytral margins smooth (Fig. 3.22B). •

Notopodia short, tapering into pointed lobes with emergent acicula; neuropodia much longer, extending I

into slender triangular lobe with digitate cirrus above emergent acicula (Fig. 3.22C). Notosetae uniformly

thin, with fine serrations, tapering to capillary tips (Fig. 3.22D); superior notosetae bent, and inferior notosetae •

longer and straight. Superior neurosetae resembling lower notosetae; inferior neurosetae slightly thicker, I

coarsely serrated in subdistal expanded region, with bifid tips (Fig. 3.22E).

Dorsal cirri with large, cylindrical cirrophores and long, acuminate styles without papillae or pigment, n

Dorsal tubercles not developed. Ventral cirri short, tapering, inserted at midlength of parapodia. Short

nephridial papillae present by segment 8, and well-developed in posterior segments. Pygidium blunt, with

dark pigment terminally, and pair of long, smooth, basally inflated cirri, l

Habitat. Found in soft sediments ranging from muds to clean, f'me sands with shell fragments or

gravel, in 2-170 m.

Remarks. This small species readily fragments and is rarely encountered whole, making it difficult 1

1

to accurately determine the number of pairs of scales. In addition, Nichols (1969) reports that nearly mature

m

females may have only 13 or 14 elytral pairs. The distinctive barred dorsal pigment pattern may fade and 1

disappear with prolonged storage in alcohol. 1

m

Type Locality and Type Specimens. California, off Santa Barbara Point Light, 25 m, holotype

(LACM-AHF Poly 0019) and paratype (LACM-AHF Poly 0020).

Distribution. Recorded along the Pacific coast of the U.S. from Puget Sound to southern California_ I

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156 I
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I

Figure 3.22. Tenonia priops: A, anterior end, dorsal view, with pharynx everted; B, median left elytron; C,

i median cirrigerous parapodium, anterior view; D, notoseta; E, distal portion of inferior neuroseta.

I 157
I

Genus Ysideria Ruff, new genus I

Type Species: Ysideria hastata Ruff, new species. Gender: feminine.

!

Diagnosis. Body dorsally flattened, with up to 37 segments. Bilobed prostomium with cephalic

peaks, with 3 antennae, 2 palps, and 2 pairs of eyes; ceratophore of median antenna in anterior notch; im

ceratophores of lateral antennae inserted beneath anterior margin. Tentaculophores of f'n-stsegment lateral to I_

prostomium, with pair of dorsal and ventral tentacular cirri, and with aeicular lobe on inner face with 0-2

U

setae. Second segment with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri. II

Fifteen pairs of elytra on segments 2, 4, 5, 7.... 23, 26, 29, and 32. Parapodia biramous, with small 1

notopodia inserted on superior margin of targe neuropodia, each with projecting acicular lobe. Notosetae

B

more slender than neurosetae, with transverse spinous rows, tapering to blunt glabrous tips. Superior neurosetae 1

thick, lancelike, with enlarged subterminal spinous region and entire, blunt tips; median and inferior neurosetae 1

thinner, with enlarged subterminal spinous region and hooked bifid or entire tips. Dorsal cirri on nonelytrigerous Ill

segments, with basally enlarged cirrophores and cylindrical styles; dorsal tubercles nodular; ventral cirri llll

short. Pygidium with pair of cylindrical anal cirri. I[

Etymology. The name is of arbitrary derivation.

Remarks. Ysideria is distinguished from most of the genera within the Harmothoinae by having •

notosetae that are more slender than the neurosetae. It is similar in many characters to the genusArcteobia, 1

but it differs in having notosetae that are all pointed rather than having some that taper to capillary tips, and

in having thick, stout, uniramous superior neurosetae. 1

Ysideria hastata Ruff, new species

Figures 3.23 A-B, 3.24 A-G I

Material Examined. California, Monterey Bay, off Santa Cruz, Sta. 1 36°56.1YN, 122°04.13'W, I

31 m, 18 Oct 1992; -----offPoint Loma, San Diego, Sta. A- 8, 32°39.84' N, 117°16.84' W, 62 m, 22 May 1980 •

(holotype, USNM 171057, 1 paratype, LACM-AHF Poly 1720); Sta. A-10, 32039.50 ' N, 117°16.13' W, 48 II

m, 14 Jan 1986 (1 paratype, SBMNH 142933); Sta. A-11, 32039.98 `N, 117016.27 ' W, 48 m, 19 Jul 1988 (1

paratype, USNM 171059), 3 Apr 1989 (1 paratype, USNM 171058); Sta. A-13, 32°40.97 ' N, 117016.57 ' W, I

43 m, 3 Apt 1989 (1 paratype, LACM-AHF Poly 1721); Sta. B-3, 32045.42 ' N, 117°18.38' W, 59 m, 4 Jan

1993 (1 paratype, MCZ 4014).

1

Description. Holotype complete, in two pieces, 22 mm long for 37 segments, width excluding setae 1

5 mm. Body elongate, flattened dorsally, arched ventrally, of nearly constant width through median region.

Dorsum with transverse triangular area of brown pigment on each setiger, becoming darker posteriorly; Ill

ventrum with similar triangular pattern plus patch of pigment at base of nephridia, or pigment in broad •

transverse band spreading onto parapodia. Prostomium trapezoidal, bilobed, with moderate to distinct cephalic m

peaks (Fig. 3.23A). Two pairs of dark eyes; anterior pair larger, anterior to widest part of prostomium; 1

posterior pair adjacent to first elytrophores. Median antenna with large, cylindrical, slightly tapering ceratophore 1

in anterior prostomial notch; style tapering, about as long as prostomium, with filiform tip and scattered short, lJ

clavate papillae. Lateral ceratophores short, inserted ventrally on prostomium; styles short, subulate, with

fitiform tips and scattered papillae. Palps stout, tapering, about prostomial length, with minute papillae and 1

short tips.

Tentaculophores elongate, lateral to prostomium and adjacent to palps, with acicular lobe on inner

face with 0-2 stout, curved setae. Two pairs of subequal tentacular cirri similar in shape and slightly longer II

1

than median antenna, with scattered clavate papillae. Second (buccal) segment with pair of large elytrophores

and first biramous parapodia; ventral buccal cirri similar to tentacular cirri, situated on basally enlarged im

cirrophores lateral to ventral mouth. Antennae, tentacular cirri, and buccal cirri with brown pigment proximally. I
Figure 3.23. Ysideriahastata, new species, holotype: A, anteriorend,dorsal view; B, posterior end,dorsalview,

left posteriormost parapodium underdeveloped.

I Fifteen pairs of large etytra completely coveting dorsum; f'trst pair suboval, slightly thicker, with

shiny surface; subsequent pairs subreniform (anterior) to subcircular (posterior), thin, dull, often wrinkled.

I Elytra smooth except for occasional micropapillae, without tubercles or fringing papillae; entire surface mottled with brown pigment, or pigmentation restricted to median and posterior regions and area around

attachment point (Fig. 3.24A). Notopodia small, much shorter than neuropodia, tapering into acutely conical

I acicular lobe on lower side (Fig 3.24B). Neuropodia large, broad, extending into presetal lobe with emergent aeicula and small, short, blunt supraacicular process; postacicular lobe shorter, subtriangular. Notosetae

moderate in number, arranged in several whorls, slightly curved, uppermost short and lowermost longer,

I reaching nearly to tip of neuropodia; each notoseta pale yellow, slender, with distinct transverse spinous rows,

i 159
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II

Figure 3.24. Ysideria hastata, new species, holotype: A, right eighth elytron from segment 15; B, left cirrigerous i

parapodium from segment 18, anterior view; C, inferior and superior notosetae, segment 18; D,

median neuroseta from 2nd segment; E, superior neuroseta, segment 18; E median neuroseta; G, •

inferiorneuroseta. []

160 1
!

I tapering to blunt, glabrous tip (Fig. 3.24C). Neumsetae less numerous than notosetae, arranged in vertical

series; those on first two parapodia slender, with long, basally expanded spinous region, tapering to thin

•I knobbed short expanded tips (Fig. region 3.24D). of spinous Thereafter bracts, superior tapering 1-2 neurosetae to glabrous,massive blunt,tips harpoon (Fig. shaped, 3.24E); amber mediancolored, neurosetae with

thicker than notosetae, light amber, with short, slightly curved spinous region tapering to glabrous, slightly

I hooked, Cirrophores bifid tips (Fig.of 3.24F); dorsal cirri inferior cylindrical, neurosetaewithmore pigmented slender,basal taperingenlargement to entire blunt on posterior tips (Fig, 3.24G). side; styles

cylindrical, heavily pigmented, sparsely papillate, tapering to filiform tips not extending beyond neurosetae.

I Dsmall orsalpapillae; tuberclesventral low tcirrophores o bluntly colow, nical,withwithdarklittle brownpigment. pigmentVentral patch oncirrimedian short,side. acumina Cylindrical te, withnephridi scatteredal

papillae projecting between parapodia from segment 6. Pygidium with anus medial to last parapodia (Fig.

I 3.23B), with terminal pair of short, sparsely papillate pygidial cirri tapering to filiform tips. Etymology. Latin hastatus, armed with a spear, in reference to the large spear-shaped superior

neurosetae.

I Remarks. The elytra of Ysideria hastata exhibit a large variation in the distribution of pigments.

This elytral mottling along with the presence of pigment on the posterior ventrum and the heavy specialized

neurosetae are suggestive of a commensal mode of living. To date, however, no potential host organisms have

I been reported.

Most of the specimens examined have the notosetae encased in a golden yellowish substance of

i unknown origin and composition. This material does not appear on the neurosetae. A similar substance occasionally occurs on the notosetae ofMalmgreniella scriptoria.

This species isrecorded asHarmothoe sp. A in the SCAMIT listing of soft bottom macroinvertebrates

I from infaunal D monitoring programs in the Southern California Bight. istribution. Reported from central and southern California from Monterey Bay southward to Point

Loma in shelf depths between 29-62 m.

!

Acknowledgements

i I wish to thank Dr. Marian H. Pettibone at the Smithsonian Institution for her conversations, notes,

and insights on some of the species in the collections of the United States National Museum of Natural History

(USNM). In addition, I would like to extend gratitude to Dr. Earle E. Spamer, Collection Manager at The

my

Academy of Natural Sciences ofPhiladelphia (ANSP), Dr. Ardis Johnston, Harvard Museum of Comparative

Zoology (MCZ), and Ms. Leslie Harris, Los Angeles County Museum (LACM) for their assistance in obtaining

i type material. A special thanks to Mr. Ron Velarde and the City of San Diego Ocean Monitoring Program for providing the specimens of the new genus and species described below.

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Hartman, O. 1978. Polychaeta from the Weddeli Sea quadrant, Antarctica. Antarctic Research Series 26:125- I

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Horst, R. 1917. Polychaeta Errantia of the Siboga-Expedition. Pt. 2. Aphroditidae and Chrysopetalidae. !

Siboga-Expeditie 24b: 1-140.

n

Imajima, M. and O. Hartman. 1964. The polychaetous annelids of Japan. Allan Hancock Foundation Occasional !

Paper 26:1-452.

Izuka, A. 1912. The errantiate polychaeta of Japan. The Journal of the College of Science, Imperial University

of Tokyo, Japan 30(2):1-262. i

Johnson, H.P. 1897. A preliminary account of the marine annelids of the Pacific coast, with descriptions of

new species. Part I. The Euphrosynidae, Amphinomidae, Palmyridae, Polynoidae, and Sigalionidae. •

Proceedings of the California Academy of Sciences 1:153-190.

Johnson, H.P. 1901. The polychaeta of the Puget Sound region. Proceedings of the Boston Society of Natural mini

History 29:381-437. !

Khlebovich, V.V. 1961. Polychaeta of the littoral zone of the Kurile Islands. Issledovaniia dal'nevostochnykh

morei SSSR 7:151-260

Kudenov, J.D. 1975. The occurrence of the polynoid Harmothoe cf. lunulata from the tube of the maldanid

Axiothella rubrocincta (Polychaeta). Bulletin of the Southern California Academy ofSciences 1975:42-

43. I

Lie, U. 1968. A quandtative study of benthic infauna in Puget Sound, Washington, USA, in 1963-1964.

Fiskeridirektoratets Skfifter, Serie Havundersokelser 14:229-556.

ml

Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bemstein, R. Cimberg, T. Kauwling, and W. Anikouchine.

1986. Assessment of long-term changes in biological communities in the Santa Maria Basin and m

western Santa Barbara Channel. Phase I. Final Report submitted forthe Minerals Management Service,

U.S. Department of the Interior, Pacific OCS Office, under Contract No. 14-12-0001-30032. III

Loi, T. 1980. Catalogue of the types of polychaete species erected by J. Percy Moore. Proceedings of the m

Academy of Natural Sciences of Philadelphia 132:121-149. i

Marenzeller, E. 1902. Stidjapanische Anneliden. 3. Aphroditea, Eunicea. Denkschriften der Akademie der m

Wissenscl'taften Wien 72:563-582. !

m

Moore, J.P. 1902. Descriptions of some new polychaeta with a list of other polychaeta from north Greenland

waters. Proceedings of the Academy of Natural Sciences of Philadelphia 54:258-278.

Moore, J.P. 1905. New species of polychaeta from the north Pacific, chiefly from Alaskan waters. Proceedings

of the Academy of Natural Sciences of Philadelphia 57:525-554.

m

Moore, J.P. 1908. Some polychaetous annetids of the northern Pacific coast of North America. Proceedings I

of the Academy of Natural Sciences of Philadelphia 60:321-364.

Moore, J.P. 1910. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern I

California in 1904. Polynoidae, Aphroditidae and Segalionidae. Proceedings of the Academy of II

Natural Sciences of Philadelphia 62:328-402.

Morris, R.H., D.P. Abbott, and E.C. Haderlie. 1980. Intertidal invertebrates of California. Stanford University l

Press, Stanford, California. 690 pp.

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!

I Nichols, F.H. 1969. Tenonia kitsapensis, a new genus and species of the family Polynoidae (Polychaeta) from

Puget Sound (Washington). Proceedings of the Biological Society of Washington 82:205-208.

I Okuda, S. 1936. Japanese commensal polynoids. Annotationes Zoologicae Japonenses 15:561-571.

Pettibone M.H. 1948. Two new species of polychaete worms of the family Polynoidae from Puget Sound and

i San Juan Archipelago. Journal of the Washington Academy of Science 38(12) :412-416.

Pettibone M.H. 1953. Some scale-bearing polychaetes of Puget Sound and adjacent waters. University of

Washington Press, Seattle, Washington. 89 pp.

I Pettibone M.H. 1956. Marine polychaete worms from Labrador. Proceedings of the U.S. National Museum

105(3361 ):531-584.

I Pettibone M.H. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae. Bulletin of the U.S. National Museum 227(1): 1-356.

i PettiboneProceedings M.H. 1967.ofType-specimens of polychaetes described by Edith and Cyril Berkeley (1923-1964). the U.S. National Museum 119:1-23.

Pettibone M.H. 1969. Review of some species referred to Scalisetosus Mclntosh (Polychaeta, Polynoidae).

I Proceedings of the Biological Society of Washington 82:1-30.

Pettibone, M.H. 1986. Review of the Iphioninae (Polychaeta: Polynoidae) and revision of lphione cimex

I Quatrefages, Smithsonian Contributions Gattyana deludens to Zoology Fauvel,428:1-43. and Harmothoe iphioneUoides Johnson (Harmothoinae).

Pettibone, M.H. 1989. A new species of Benhamipolynoe (Polychaeta: Polynoidae: Lepidastheniinae) from

I Australia, associated with the unattached stylasterid coral Conopora adeta. Proceedings of the Biological Society of Washington 102(2):300-304.

i Pettibone,(Polychaeta: M.H. 1993a.Polynoidae: Revision ofHarmothoinae). some species referred Smithsonian to Antinoe Contributions , AntinoelIato ,Zoology Bylgides545:1-41., and Harmothoe

Pettibone, M.H. 1993b. Scaled polychaetes (Polynoidae) associated with ophiuroids and other invertebrates

I and reviews of species referred to MaImgrenia Mclntosh and replaced by Malmgreniella Hartman, with descriptions of new taxa. Smithsonian Contributions to Zoology 538:1-92.

i Pettibone,Basin. M.H.Proceedings 1993c. Polynoid of thepolychaetes Biological Society associatedof Washington with a whale106(4):678-688. skeleton in the bathyal Santa Catalina

Skogsberg, T. 1942. Redescription of three species of the polychaetous family Polynoidae from California.

I Proceedings of the California Academy of Sciences 23:481-502.

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I Philadelphia. Part 2: Annotated catalogue of recent type specimens. Tryonia 23:1-303. Steinhauer, M. and E. Imamura (eds.). 1990. California OCS Phase II Monitoring Program. Year-Three

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I Pacific OCS Region, under contract No. 14-12-0001-30262.

Tebble, N. and S. Chambers. 1982. Polychaetes from Scottish waters. A guide to identification, Part 1,

Family Polynoidae. Royal Scottish Museum Studies, Edinburgh. 73pp.

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reference to the planktonic larvae inthe Sound (Oresund). Meddeletser fra Kommissionen for Danmarks

Fiskeri- og Havunders0gelser, serie Plankton 4(I): 1-523. I

Treadwell, A.L. 1906. Polychaetous annelids of the Hawaiian Islands collected by the steamer Albatross in

1902. Bulletin of the U.S. Fisheries Commission 23(3): 1145-1181.

Treadwell, A.L. 1937. The Templeton Crocker Expedition. VIII: Potychaetous annelids from the west coast I

of Lower California, the Gulf of California and Clarion Island. Zoologica 22:139-160.

Uschakov, RV. 1955. Polychaetous annelids of the far eastern seas of the USSR. Akademiya Nauk SSSR, i

Keys to the fauna of the SSSR 56:1-433. [translated 1965 from Russian by the Israel Program for

Scientific Translations, Jerusalem]

Uschakov, P.V.1982. Polychaetes of the suborder Aphroditiformia of the Arctic Ocean and the northwestern i

part of the Pacific, Families Aphroditidae and Polynoidae. Fauna of the USSR, Polychaeta 2(1): 1-

272.Canada][lranslated 1987 from Russian by the Multilingual Services Division, Translation Bureau, Ottawa, I

Wesenberg-Lund, E. 1950. Polychaeta. Danish Ingolf-Expedition 4(14): 1-92.

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the voyage of the Southern Cross 12:262-283.

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166 I
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I 4. FAMILYACOETIDAEKINBERG, 1865

| by James A. Blake 1

I

I Introduction

I The Acoetidae (=Polyodontidae) are scaleworms with large bodies that are elongated and dorsoventrally flattened. They typically have 300 or more segments, and some may reach gigantic proportions with more

than 600 segments and lengths of up to a meter or more and widths of up to 40 mm. These worms live in long

I tubes constructed of thin interwoven threads that are combined with adhering particles of mud or sand. They are carnivorous and extend out of their tubes to catch prey. Acoetids are generally tropical in distribution and

include eight genera and about 45 species.

!

i Morphology

Acoetids have bodies that are partially covered with numerous dorsal elytra. The dorsal surface of

I the body is usually transversely grooved, often obscuring segmental boundaries. The ventral surface is smooth. The head region is well developed. The prostomium and tentacular segment are more or less fused

and bear long ventral palps, antennae, tentacular cirri, and eyes (Fig. 4.1A, 4.2A). The prostomium is oval or

I slightly bilobed, and bears a pair of lateral antennae and sometimes a median antenna. In most genera, includingAcoetes and Polyodontes, the anterior pair of eyes is large and located on a stalked structure called

the ommatophore (Fig. 4.1A, 4.2A). These eyes are very characteristic and readily distinguish acoetids from

I other scaleworms. The tentacular segment bears one or two aciculae and a pair of dorsal and ventral tentacular • cirri; setae may be present or absent. Elytra are first present on the second segment which is biramous and

forms the lateral lips of the mouth. The pharynx is eversible, large, muscular, and bears two pairs of jaws.

I There is usually a pair of papillae around the oral opening. The elytra overlap along the sides of the body, leaving the mid-dorsum uncovered. They are attached

to large elytrophores on segments 2, 4, 5, and 7, continuing on alternating segments down the body (Fig. 4.1E,

I 4.21). Individual elytra are relatively small, rounded and smooth (Fig. 4.1B). Dorsal cirri are present on

those segments lacking elytra (Fig. 4.2E). Branchiae are reported for some species of Polyodontes. When

present, the branchiae have a thickened cuticle and represent non-vascular coelomic extensions on the parapodia

I and the bases of the elytra. They may be papilliform, filamentous, or arborescent.

Setae are all simple. Notopodia are reduced anteriorly, with a bundle of notosetae present or absent.

Notopodia of middle body segments form enlarged flattened lobes on the anterodorsal sides of the neuropodia

I (Fig. Again, may present or spinning glands (=web glands) are present,

4. 1E). notosetae be absent; coiled

producing soft silky threads that are used in tube building. The silken fibers are actually modified setae which

emerge from the notopodium. Neuropodia are well developed and accessory bracts may be present. Neurosetae

I include stout acicular spines with spinelets, thin hirsute pointed setae, and thin, spirally coiled setae with

i 1ENSRConsultingandEngineering8, 9WaterStreet,WoodsHole,Massachusetts02543

I 167
!

various types of serrations (Fig. 4.1G-J, 4.2D, F-H); some neurosetae have terminal hairs (Fig. 4.1K). From m

segment 9, the upper group of neurosetae form a low anterodorsal bract on the neuropodium opposite the

place where the spinning fibers arise. According to Pettibone (1989), these setae may function in the separation I

of the spinning fibers. Two types of neurosetae are present: (a), stout and long (Fig. 4.1Ka; 4.2Ja); and (b), U

shorter, more slender (Fig. 4.1Kb; 4.2Jb). The structure of these setae differ among genera and species, and

Pettibone (1989) has greatly emphasized them as taxonomic characters. Ventral cirri are short and pointed m

(Fig. 4.1C-E; 4.2B, E, I). The pygidium bears 2 long anal cirri. il

Biology n

The Acoetidae are tube dwellers. They line the wails of their tubes with fine collagenous threads. U

These threads are secreted by the spinning glands and are woven into a criss-cross, spiralled pattern. They

are usually combined with mud, clay, or sand particles, forming very strong, thickened tubes that may extend !i

up to a meter or more in length and be up to 4 cm in width. The tubes of acoetids are unique among the l

polychaetes for the complete absence of a membranous lining. According to Pettibone (1989), the felted

appearance formed from the threads are homologous with the fibers comprising the dorsal felt of Aphrodita. 1

The l tube and body of acoetids provides habitat for a variety of commensals. Entoprocts are attached

to the wails of the tubes (Franzrn, 1962; Nielsen, 1966; Pettibone, 1989), gastropods live under the elytra

(Moore, 1972), montacutid bivalves live between parapodia (Rullier, 1965; Intes and Le Loeuff, 1975), and []

other polychaetes live within the tubes (Fox and Ruppert, 1985). The entoprocts use the respiratory currents U

generated in the tubes to filter food particles.

Acoetids are carnivorous, using their tubes as retreats from which they extend, grab their prey, and i

pull back. Pettibone (1989) describes reports from the Mediterranean where large specimens of Polyodontes U

macillosus have been captured on fishing lines. The worms were attached to the bait (hermit crabs, worms,

and small fish) by the everted proboscis. B

HI

Sexes are separate. Nothing is known about the reproduction and development of species of Acoetidae.

Pettibone ( !989) describes juveniles of two species, i=i

Taxonomic History I

The most important review and revision of the Acoetidae is by Pettibone (1989). In this work the m

entire taxonomic history of the family is reviewed along with keys, descriptions, and illustrations of all of the U

known species. All available information on the biology and natural history of these scateworms is also

reviewed. IBm

The species of the Acoetidae have been included in virtually every scale worm family at one time or B

another. In most recent keys or monographs, including Hartman (1968), species of this group have been

included in the family Polyodontidae. Pettibone (1989), however, points out that the Acoetidae Kinberg, I

1856, based upon Acoetes Audouin and Milne-Edwards, 1832 has priority because the family name II

Polyodontidae Augener, 1918, based upon Polyodontes Renieri in BlainviUe, 1828, is preoccupied in Fishes

by Polyodontidae Bonaparte, 1837, that is based on the genus Polyodon Schneider, 1801. I

Most Acoetidae are tropical or subtropical. However, Hartman (1968) records three species of II

acoetids (as Polyodontidae) from southern California. Peisidice aspera Johnson, also included with this

group by Hartman (1968), belongs to the genus Pholoides in the family Pholoidae (Blake, 1995, this volume). I

168 I
!

I One species,Acoetes pacifica has been collected from the Santa Mafia Basin. Two additional species,

Polyodontes lupinus and P. panamensis, have been reported from southern California (Hartman, 1968;

I Pettibone, 1989).

i Key to the Genera and Species From California

i 1A. Upper neurosetae of type a from segment 9 long, tapering to fine tips, thickly spinous distally (Fig. 4.1Ka); type b neurosetae shorter than type a (Fig. 4.1Kb) but not completely hidden by notopodia

................................................................................................................ Genus Acoetes: A. pacifica

I lB. Uplumose pper neurosetae of type a from segment 9 long, abruptly tapering to slender tips (Fig. 4.2Ja), subdistally and short spinous rows basally; type b neurosetae very short (Fig. 4.2Jb), hidden

by notopodia ............................................................................................... Genus Polyodontes ... 2

I 2A. With numerous elongate oval branchiae on elytrophores and dorsal cirrophores from about segment

10, replaced by few digitiform branchiae from about segment 34 .................... Polyodontes lupinus

I or single digitiform on some segments segment ..............

2B. Branchiae absent, with branchiae from 30

.................................................................................................................. Polyodontes panamensis

!

I Descriptions of Species

I Genus Acoetes Audouin and Milne-Edwards, 1832

I Type Species: Acoetes pleei Audouin and Milne-Edwards, 1832, by monotypy.

Diagnosis. Prostomium bilobed, with pair of bulbous stalked ommatophores directed anteriorly and

i pair of small eyes; median antenna with ceratophore in middle ofprostomium. First or tentacular segment distinct dorsally; tentaculophores lateral to prostomium, each with 2 aciculae, 2-4 groups of capillary setae

and pair of tentacular cirri. Second segment with first pair of etytra and long ventral buccal cirri; biramous

i parapodia with bundle of notosetae; neurosetae slender, spinous, lanceolate. Acicular neurosetae from segment 3. Dorsal cirri with short cirrophores and short styles on segments lacking elytra; ventral cirri short, subulate.

Notopodium from segment 9 wide, flattened, anterodorsal to neuropodium, with notoacicula, internal spinning

glands, and small row of delicate notosetae; neuropodium with lower group of neurosetae within anteroventral

I bract, neurosetae curved, spinous, lanceolate; middle row of stout acicular neurosetae, or afistae;

with without

upper group of neurosetae, emanating from low anterodorsal bract (hidden by notopodium), of 2 types: a)

longer, abruptly tapered to slender spinous tips, long spines subdistally and short spinous rows basally, b)

I short (hidden notopodium), Distal border muscular pharynx with 13-15 pairs of papillae, by bipinnate. of large

middorsal and midventral ones on wide lobulated bases, middorsal one longer than others; 2 pairs of hooked

i -- jaws, each with 5-9 lateral teeth.

i 1 Remarks. The genus Acoetes contains 12 species, one of which is known from California and has

been found in the Santa Mafia Basin. All of the known species were described by Pettibone (1989) and a key

i provided.
!

Acoetespacifica (Treadwell, 1914) I

Figure 4.1

I

Panthalis pacifica Treadwell, 1914:184, pl. 11, figs. 1-7.--Hartman, I939:87, pl. 26, figs. 309-312.-

Berkeley and Berkeley, 1941:24.--Emerson, 1971 :137.--Fauchald, 1972:29; 1977:9. |

Acoetes pacifica: Pettibone, 1989:87-91, figs. 63-64 (synonymy).

I

Material Examined. California, western Santa Barbara Channel, Sta. 73, 98 m (1); Santa Mafia d,.

Basin, off Point Sal, Sta. R-8, 90 m (1). I_

u

Description. A relatively small species, recorded up to 40+ mm long and 10 mm wide for 53+

segments (Pettibone, 1989); Santa Maria Basin specimens smaller, 31 mm long, 7 mm wide, for 62 segments II

on complete specimen from Sta. R-8. Body broad, dorsoventrally flattened. Anterior few pairs of elytra •

large, oval, coveting dorsum, following ones leaving middorsum uncovered; elytra delicate, thin, more or less

i

pigmented with brown on medial part, with lateral pouch from about segment 9 (Fig. 4.1B). Color in alcohol:

flesh. I

Prostomium bilobed with pair of rounded ommatophores, without distinct neck (Fig. 4.1A); median

antennae with or without papillae on sides of cemtophore; lateral antennae arising ventral to ommatophores, I

with tips extending beyond ommatophores; posterior pair of eyespots lateral to ceratophore of median antenna; I

ventral palps long, smooth, tapered, 2-3 times longer than prostomium (Fig. 4.1A). Tentacular segment with

bundle of slender setae on inner side, row of globular papillae, and pair of dorsal and ventral tentacular cirri, Ill

each longer than median antenna. Distal border of extended pharynx with 13-15 pairs of papillae, middorsal |

one much longer than others, midventral one longer than others but shorter than middorsal one; 2 pairs of

hooked jaws, each with 6-8 lateral teeth. an

Second segment with first pair of elytrophores (Fig. 4.1A), ventral buccal cirri longer than following I_

ventral cirri, and biramous parapodia; notopodium small digitiform lobe (Fig. 4.1C) with bundle of long,

slender, finely spinous notosetae (Fig. 4.1F); neuropodium with longer presetal acicular lobe, shorter, rounded It

postsetal lobe, and ventral bract; neurosetae slender spinous capillaries (Fig. 4.1G). 1

Third segment with first pair of dorsal cirri, each with short cirrophore and style extending beyond

setae (Fig. 4.1D); parapodia similar to segment 2 except presetal neuropodial lobe shorter and middle neurosetae

stout, acicular, aristate (Fig. 4.11). Parapodia of segments 4-8 with fewer, shorter notosetae; upper neurosetae m

similar to anterior parapodia; lower neurosetae slender, with larger spines more basally and close-set spinous

rows distally on tapered capillary tips (Fig. 4.1J); middle stout acicular neurosetae with distal hairs on bases 1

of aristae. 1

Notopodia becoming larger,rounded, flattened from segment 9 with appearance of notoacicula, spinning

glands, andfew short notosetae (Fig. 4.1E); neuropodium with slightly bilobed presetal acicular lobe, truncate •

postsetal lobe and distinct anteroventral bract; lower neurosetae numerous, similar to anterior parapodia; i

middle row of stout acicular neurosetae with distal hairs on bases aristate, some in more posterior parapodia

with additional subdistal spines along one side; upper group of neurosetae, emerging from low anteroventral 1

bract, of 2 types: a) long, slender, abruptly tapered to slender tip, with subdistal long hairs and rows of shorter II

spines more basally (Fig. 4.1Ka); b), short, slender, aristate (Fig. 4.1Kb), hidden by notopodium. Branchiae

absent. Pygidium with two long, tapering anal cirri. II

!1 Biology. Constructs tubes in fine, green muds; recorded from continental shelf depths in California,

90-163m.

I

170 i
I

I

I _c

_VtC _ I

°_q...' X._.(]i .@ _:: J'

__ A spinninoQland _i"!!::: : ,!,_,

, . ,, "\ :-, ::'.- "_, \ t _ 'i.::-,'.:" -, " "

I

Figure 4.1. Acoetes pacifica: Anterior end, dorsal view, left paralx_lium of segment 2 omitted; B, elytmn

from segment 25; C, elytragerous parapodium of segment 2, anterior view; D, cirrigerous

I parapodium of segment 3, anterior view; E, elytragerous parapodium of segment 25, anterior view, acieula and spinning gland dotted; F, notoseta from segment 2; G, neuroseta from segment

2; H, upper neuroseta from segment 3; I, middle neuroseta from segment 3; J, lower neuroseta

I . fr1989). om segment 3; K, upper neurosetae from segment 9 (types a and b indicated). (after Pettibone,

I

I Type Locality. California, off San Diego, 91-163 m (holotype, LACM-AHF Poly 52; 2 paratypes

AMNH 769, 771).

I Remarks. Acoetes pacifica differs from A. mortenseni from off Panama in having cylindrical ommatophores on the prostomium without distinct necks instead of ones with necks and smooth palps instead

of ones which are papillated.

I Distribution. Central California to Lower California; Panama; Gulf of Mexico; 10-640 m.

I

I 171
I

Genus Polyodontes Renieri, 1828 I

II

Type Species: Phyllodoce maxilfosa Ranzani, 1817, by monotypy.

Diagnosis. Prostomium bilobed, with pair of large ommatophores on anterior part of prostomium i

and pair of small eyespots; median antenna with ceratophores in middle of prostomium. First or tentacular i

segment distinct dorsally; tentaculophores lateral to prostomium, each with 2 aciculae, 2 groups of capillary

setae and pair of tentacular cirri. Second segment with first pair of elytra and long ventral buccal cirri; I

biramous parapodia with bundle of capillary notosetae; neurosetae both stout and slender, spinous, tapering II

to slender tips. Stout acicular neurosetae, with or without aristae, beginning on segment 3 or more posteriorly.

Dorsal cirri on non elytra-bearing segments, with short, subulate cirrophores and short styles; ventral cirri i

short, subulate. Notopodium from segment 9 wide, flattened, anterodorsal to neuropodium, with notoacicula, i

internal spinning glands, and small bundle of short capillary notosetae on inner side of notopodium; neuropodium

truncate, with lower group of neurosetae within more or less distinct anteroventral bract; middle row of stout i

acicular neurosetae; upper group of neurosetae, emanating from low anterodorsal bract (hidden by notopodium), Ii

of 2 types: a) long, stout, thickly spinous, tapering to rme tips and b) short, slender, tapering to sharp tips,

bipinnate. With or without parapodial branchiae. Distal border of large muscular pharynx with 13-19 pairs II

of papillae, middorsal and midventral ones on wide lobulated bases, longer than others; 2 pairs of hooked l

jaws, each with 2-10 lateral teeth.

..... i

eve tAn .-_:?".:-. . - f a iI

'

DtC .(_ elph

f./; _ommatophore "',.'.. II

" I

? '"°'°''°2°

........... .. ,., ':,-"2 K !

vc _

!

Figure 4.2. Polyodontes panamensis: A, anterior end, dorsal view; B, elytragemus parapodium of segment 2,

anterior view; C, notoseta from segment 2; D, middle and lower neurosetae from segment 2; E, II

cirrigerous parapodium of segment 3, posterior view; F-H, upper (F), middle (G), and lower (H) I

neursetae from segment 3; I, elytragerous parapodium of segment 9, anterior view, acicutae and

spinning gland dotted; J-K, upper (J, types a and b indicated) and lower (K) neurosetae from II

segment 9. (after Pettibone, 1989). I

172 i
!

I Remarks. The genus Polyodontes contains 12 species, two of which have been recorded from

California (Pettibone, 1989). Polyodontes lupinus (Stimpson), and P. panamensis (Chamberlin) have been

I reported (see above)fromandsodetailed uthern Cdescriptions alifornia (Hamayrtman, be found 1968;inPettibone, Pettibone (1989). 1989). Figure Both s4.2pecies depicts are ssome eparamorphological ted in the key

features of a typical species of Polyodontes.

I Literature Cited

!

Berkeley, E. and C. Berkeley. 1941. On a collection of Polychaeta from Southern California. Bulletin of the

Southern California Academy of Sciences 40:16-60.

i Blake, J.A. 1995. Chapter 5. Family Pholoidae Kinberg, 1858. In: Blake, J.A., B. Hilbig and RH. Scott

(eds.), Taxonomic Atlas of the Santa Maria Basin and Western Santa Barbara Channel. Volume

I 5:175-188. Santa Barbara Museum of Natural History, California.

Emerson, R. 1971. Chapter 6: Some polychaetous annelids from the Santa Barbara Shelf Area. In Dale

Straughan (compiler), Biological and Oceanographical Survey of the Santa Barbara Oil Spill 1969-

I 1 1 and 117-147. of Southern California.

970, (Biology Bacteriology): University

Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adjacent areas in

I the eastern Pacific Ocean. Allan Hancock Monographs in Marine Biology 7:1-575.

Fauchald, K. 1977. Polychaetes from intertidal areas in Panama, with a review of previous shallow-water

records. Smithsonian Contributions to Zoology No. 221:1-81.

I Fox, R.S. and E.E. Ruppert. 1985. Shallow-water marine benthic macroinvertebrates of South Carolina,

i species identification, community composition and symbiotic associations. The Belle W. Baruch

I Library in Marine Science 14:1-330.

Franz6n, A. 1962. Studies on Entoprocta from the west coast of Sweden. Zoologiska Bidrag frS_nUppsala

33:311-326.

I Hartman, O. 1939. Polychaetous annelids, Part I: Aphroditidae to Pisionidae. Allan Hancock Pacific

Expeditions 7:1-156, 28 pls.

I Hartman, O. 1968. Atlas of the Errantiate Annelids from California. Allan Hancock

Polychaetous Foundation,

University of Southern California, Los Angeles, 828 pp.

I Intes, A.rendu a D.syst6matique. Le Loeuff. 1975 Cahiers . Les ORSTROM, Ann61ides Polych_tes Oc6anographie de C6te13:267-321. d'Ivoire I: Polych&es errantes: Compte

Moore, D.R. 1972. Cochliolepis parasitica, a nonparasitic marine gastropod, and its place in the Vitrinellidae.

I Bulletin of Marine Science 22:100-112.

Nielsen, C. 1966. Studies on Danish Entoprocta. Ophelia 1:1-76.

I Pettibone, M.H. 1989. Revision of the aphroditoid polychaetes of the Family Acoetidae Kinberg (=Polyodontidae Augener) and reestablishment ofAcoetes Audouin and Milne-Edwards, 1832, and

Euarche Ehlers, 1887. Smithsonian Contributions to Zoology, Number 464, 138 pp.

I Rullier, E 1965. Contribution _ la faune des Ann6lides polych&es du Dahomey et du Togo. Cahiers

ORSTROM, Oc6anographique 3(3):5-66.

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Treadwell, A.L. 1914. Polychaetous annetids of the Pacific coast in the collection of the Zoological Museum i

of the University of California. University of California Publications in Zoology 13:175-234, pls.

11-12. I

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I 5. FAMILY PHOLOIDAE KINBERG, 1858

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James A. uYBlake 1

l Introduction

I The Pholoidae includes small crawling scaleworms that are found in algal holdfasts, under rocks and

in crevices among oysters and mussels, and creeping over the surface of muddy sediments littered with shell

hash. depths.Some species are interstitial. They are widely distributed and occur from the intertidal zone to great

I Morphology

I The bodies of pholoids are small, flattened, with 19 to 90 segments. The dorsum may have adhesive

tubercles and the ventrum may bear papillae. The prostomium is fused to the fLrStor tentacular segment, and

I bears are present a singleor median absent. antenna, A facial atubercle pair of palps is usually and ofpresent ten two onpairstheofdorsal sessilefaceeyes;of twtheo shmouth ort lateral openinantennae g. The

muscular pharynx is eversible, and has two pairs of interlocking jaws and a circlet of papillae around the

i opening. The tentaculophores represent the In'st segment, are lateral and anterior to the prostomium and bear 1-2 pairs of tentacular cirri; setae may be present or absent on the tentacular segment. The elytra are present

from segments 2, 4, 5, 7, continuing on alternate segments to segment 23, after which they are present on

i every segment, or on alternating segments. Elytra are soft and delicate, with surface and marginal papillae; or

stiff and rigid with concentric rings, and fringes of marginal papillae; they cover the dorsum or leave the middorsum uncovered. Parapodia are biramous and supported by acicula. Bracts are present, but basal ones are

i absent in the neuropodia. Notopodia are smaller than neuropodia. Notosetae are finely spinous capillaries,

and neurosetae are compound falcigers with short blades. Dorsal cirri and branchiae are completely absent,

but prominent, knoblike dorsal tubercles are present on segments lacking elytra. Ventral cirri are short and

tapered. The pygidium is small, and bears two anal cirri. The aberrant interstitial genus Metaxypsamma

I has uniramous with and nodular lobes instead of Woff, 1986, parapodia a single neuroacicula, papillated

elytra (Pettibone, 1992).

I Taxonomic History

I The species of Pholoidae have been included in the Sigalionidae, Aceetidae, and Polyodontidae by

various authors. Hartman (1959; 1968) placed Peisidice Johnson, 1895 in the Polyodontidae and Pholoe

I Johnston, family Peisidicidae 1839 in thewasSigali established onidae. bySheDarboux also inclu(1900) ded Pholoides for PeisidicePruvbut ot,was 1895largely as a synonym ignored until of Phol resurrected oe. The

by Hartman and Fauchald (1971). Fauchald (1977) established the family Pholoididae, based upon Pholoides

I Pru family, vot asbutaplaced new name PhoIoeforinPeisidicidae. the Sigalionidae. Fauchald included Parapholoe Hartmann-Schr6der, 1965 in this

I 1ENSR Consulting and Engineering, 89 Water Street, Woods Hole, Massachusetts 02543

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Pettibone (1982) resurrected the family Pholoidae Kinberg, 18582in which she included Pholoe and I

Pholoides (including Peisidice). In a recent monograph, Pettibone (1992) synonymized Paraeupholoe

Hartmann-Schr6der, 1962 and Parapholoe Hartmann-Schr6der, 1965 with Pholoides Pruvot and added

Metaxypsamma Wolf, 1986, and three new genera to the Pholoidae. In this same work, she provided a g

definition of the family Pholoidae, keys to the genera, and complete accounts of the species in all genera

except Pholoe within which, two species were treated. No review of the species of Pholoe has been published. •

The Photoidae are separated from the Sigalionidae by the characters listed in Table 5.1. The key II

separates the two genera Pholoe and Pholoides which occur in California and Santa Maria Basin. A review

and partial revision of the genus Pholoe is in preparation by M. E. Petersen, personal communition). I

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Table 5.1. Comparative Characteristics of the Families Pholoidae and Sigalionidae (after Pettibone,

1992). i

Characteristic/Family Pholoidae Sigalionidae I

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Body Short, subrectangular, with Long, narrow, slender, with u

segments few to moderate segments numerous (>300);

(<90); crawling forms burrowing forms 1

m

Branchiae Absent Present, attached to lateral sides

of elytrophores and dorsal /

tubercles |

Tentaculophores of segment I Medial to palps Dorsal to palps

I

Neuropodia Without basal bracts and distal With basal bracts (absent in 16

stylodes Sigalion) and/or distal stylodes •

im

Compound neurosetae Blades short, falcate Blades short and long, •

multi-articulated, falcigers or !1

spinigers

I

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Family Pholoidae Kinberg, 1858. Type genus: Photoe Johnston, 1839. Original spelling: Pholoidea, corrected by •

Pettibone 1982. I
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I Biology

I a proboscis equipped four jaws, suggesting they are (Fauchald

Pholoids have with that carnivorous

and Jumars, 1979). Direct evidence for a carnivorous feeding mode has been provided by Pleijel (1983), who

described feeding habits of Pholoe minuta (Fabricius) from the west coast of Sweden. Pleijel was able to

I make observations on living animals in an aquarium andalso examined contents and feces. He found the

gut

most common prey organism to be sedentary polychaetes and small crustaceans. P.minuta encounters prey

items as they move through the sediment. The worm then attacks by rapidly evening its armed proboscis. An

I important prey species was the spionid Prionospio, which was either eaten whole or in part. Heffeman (1988) described the fme structure of the digestive system ofP. minuta, including histochemistry of secretory

cells associated with the proboscis and jaws. The digestive tract of P. minuta is composed of an anterior

i foregut or oesophagus (mouth to segment 6), a muscular pharynx (6 to 16th segment), and the intestine which continues to the anus. During feeding, the pharynx is everted and overlain with the oesophagus. Mucoid

secretions produced by the oesophagus probably serve to lubricate the pharynx during eversion and inversion

I and livesits(Heffernan, tough cuticular 1988).surface Secretions is aprointectio the pharynx n againstassist abrasi inomovement n by the sediment of the preyparticalong les inthewhich digestive the worm tract.

Aspects of the life history of Pholoe spp. are provided by Christie (1982), Heffernan et al. (1983),

I Heffeman (1985), and Heffernan and Keegan (1988a-b). Christie (1982) dealt with two species: P.minuta and P.pallida Chambers, 1985 (as P.cf. anoculata Hartman) from England, while the various studies conducted

by Heffernan dealt with P.minuta from Galway Bay, Ireland. Pholoe minuta has been found to be a polytelic

I species, and Keegan, that reaches 1988a). sexual The worms maturitylivebyatthe leastthirdfouryear, years.thenChristie annually(1982) spawni found ng twboth o oPr three minutatimesand(Heffernan P.pallida

to retain their gametocytes in the gonads until maturity. Gametogenesis was well synchronized and followed

I byspawning, arestrictedand breeding the type ofseaslarval on. Christie development. found that Smalltheeggs two(120-150 species differed prn) werein the produced size of bythePgametes, . minuta intime Apriofl

that developed into planktotrophic larvae. In contrast, P.pallida produced large, yolky eggs (180-200 pm in

I diamete primitiver) insperm, November wheasthatmales develoofpedP. paintUoidalecithproduced otrophic long-headed larvae. Malesaber ofPrant . minuta sperm. produced Primitive small-headed, sperm are

associated with a dispersal type of spawning mechanism and normally planktolxophic larvae, while aberrant

i sperm 1956). areSpawning associatedtimeswithforsome P. minuta form inofIreland brood protecti (March-Ap on riandl) nreported on-planktotrophic by Heffemandevel andopment Keegan(Franzrn, (1988a)

agreed with the observations by Christie (1982) in England. Lacalli (1980, 1981) found pelagic larvae of P.

I minuta in March-April in May-June in west in New Greenland Brunswick, and suggested Canada. that Curtisthe(1977 larvae) repwereortedlecithotrophic. that spawning ofP. minutaoccurred

Pholoe anoculata Hartman, 1965 was determined to be a dominant infaunal species in Atlantic

I cfound ontinental to reproduce slope depths year round, from thebutCan sexually adian border mature tospecimens off Northwere Carolin onlya encountered (Blake, 1993). in upper The species slope (600- was

1000 m) sediments, while specimens from depths of 2000 m or greater were not gravid. These results suggested

I that recruitment to the lower slope depths was by larvae that came from shallower depths. Actual observation on the morphology of embryos and larvae of pholoids is limited to Pholoe minuta

(._esson, 1963; Sveshnikov,1967; Lacalli, 1980; Heffernan and Keegan, 1988b), Pholoe synophthalmica

I (Cazaux, 1968), Pholoides asperus (Blake, 1975a as Pholoe minuta), and for several of the smaller interstitial species (Laubier, 1975; Wolf, 1986; Pettibone, 1992). Four patterns are apparent: (1) species with small eggs

and planktotrophic larvae; (2) species with large eggs and leeithotrophic larvae; (3) species with large eggs,

I viviparity. special formsPelagic of broolarvae ding ofunder photoids the epass lytra through with direct trochophore, development; metatrochophore, and (4) speciesandwithnectolarge chaeteeggsstages and

similar to those of the Polynoidae and Sigalionidae. Species with direct development or viviparity do not

I exhibit any of the traditional morphology associated with pelagic larvae.

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Key to the Genera and Species of Pholoidae

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1A. Prostomium oval, bilobed, with median antenna on ceratophore in anterior notch (Fig. 5.1A, 5.2A);

with or without lateral antennae; tentaculophores achaetous, with dorsal and ventral tentacular cirri; i

elytra on segments 2, 4, 5, 7, and continuing on alternate segments to segment 23, then on every li

segment to end of body; elytra delicate, with border and surface papillae (Fig. 5.1C-D, 5.2D-E) ....

.................................................................................................................... Genus Pholoe ............. 2

tB. Prostomium subrectangular, with median antenna on ceratophore on anterior border; without lateral II

antennae (Fig. 5.3B); tentaculophores with notosetae and single tentacular cirrus; elytra on segments

2, 4, 5, 7, and continuing on alternate segments to end of body; elytra thick, with concentric rings and 1

numerous long border papillae (Fig. 5.3B, G) ........................................................ Genus Pholoides

2A. Eyes present; median antenna and dorsal and ventral tentacular cirri short, with pseudoarticulated D

tips (Fig. 5.1A); papillae on elytra with weak folds or joints, not expanded attip, more or less of the

same length on elytra throughout body (Fig. 5.1D,E) ................................................. Pholoe glabra i

2B. Eyes absent; median antenna and dorsal and ventral tentacular cirri long, with thin tapering tips (Fig. i

5.2A); papillae on elytra simple, with weakly expanded tip, short on anterior elytra, becoming very

long on middle and posterior elytra (Fig. 5.2E) ................................................... Pholoe courmeyae I

Descriptions of Species i

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Genus Pholoe Johnston, 1839

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Type Species: Pholoe inornata Johnston, 1839. i

Diagnosis. Body small, linear, upto 90 segments. Elylra on segments 2, 4, 5, 7, andcontinuing on l

alternate segments to 23, then on every segment to endof body. Elytra delicate, with papillae on border and i

surface. Prostomium andfirsttentacular segment fused, ventrallyforming anterior lip of mouth, with projecting

facial tubercle; or facial tubercle lacking. Prostomium rounded, bilobed, with median antenna on ceratophore i_

in anterior notch, with or without lateral antennae; eyes present or absent, sometimes fused. Parapodia

biramous; notopodial lobe conical, acicular, with subdistal bract on dorsal face; neuropodial lobe larger,

conical, acicular, sometimes with distal papillae. Notosetae simple, slender, capillary, finely spinose, of 2 i

types: shorter, strongly bent or geniculate, and longer, slightly curved or straight. Neurosetae stouter than U

notosetae, compound, with shafts spinose subdistally and blades short, spinose, falcate, and unidentate.

Pygidium with pair of anal cirri, i

Remarks. Pholoe gIabra appears to be the most common species in California (see below). Pholoe

minuta (Fabricius) is a widespread species and may be present in California estuaries and other nearshore

habitats, but it has not been encountered in this study. Several species appear to have been confused with P. i

minuta in the North American literature and a review of these records is needed. Pettibone's (1953) description i

ofP. minuta from the Puget Sound appears to be ofP. glabra.

i

Specimens of a deep-water species of Pholoe have been encountered in the MMS collections as well i

as in samples taken from the continental slope off northern California. The species is identified as Pholoe

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courtneyae, new species.

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I Pholoe glabra Hartman, 1961

Figure 5.1

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Pholoe glabra Hartman, 1961:51-54, pI. 2, figs. 1-8.--1968:157, figs. 1-6.--Blake, 1975b:172, 176-177,

i fig. 82.

Material Examined. California, Santa Maria Basin, off Port San Luis, Sta. 21, 49 m (41); offPoint

I 6,Sal,148Stas. m,PJ-1,145 m, (21); PJ-2, 142 m, (25+); PJ-3,138 m, (25+); PJ-4, 150 m (25+); PJ-5, 152 m (8); PJ- (40+); PJ-7, 123 m, (100+); R-8, 90 m (40+); Sta. 103, 197 m (1).

Description. Body small, linear, depressed, with about 38 segments; 10-15 mm long, 2 mm wide.

I Mid posteriorly dorsum (Fig. completely 5.1D); covered each elytron by overl withapping shortelytra. marginal Elytra papillae; oval anteriorl papillaey (Fsimple, ig. 5.1C), slender becoming with narrow subquadrate folds

or articles, several small sensory hairs on tip. Color: white with rust-colored dorsum.

I Prostomium with 2 anterior triangular-shaped lobes, separated by a deep notch (Fig. 5.1A); with 2

pairs of smaU, closely set, black eyes; median antenna short, thick, tapering distally, with 3-4 pseudoarticulated

joints; palps massive, tapering. Facial tubercle short, triangular. Proboscis with 18 terminal papillae and 2

I pairs of dark jaws. Two pairs of thick tentacular cirri, similar in appearance to median antenna, with pseudoarticulations.

Notopodium small, distally rounded; neuropodium large, distally pointed, with small surficial papillae;

I ventral cirrus thin, fmgerlike (Fig. 5.1B). Both podial lobes bearing stout aciculae and dense fascicles of

setae; notoseize simple, with 6-12 serrated, geniculate seize in supm-acicular position (Fig. 5.1E) and 5-12

long, smooth capillaries in sub-acicular position. Neurosetae all compound falcigers, with short blade and

I f'me serrations on base of blade and tip of shaft (Fig. 5.1F). Pygidium with pair of long, cirriform anal cirri.

Biology. This small carnivorous species is widespread and abundant from central California to Mexico

i in continental shelf and slope depths in sediments of olive green silt. In the Phase II monitoring program the species was one of more abundant polychaete species, with densities of 154and 259 individuals m-2at stations

R-5 and PJ-1, respectively.

i Basin As part of a life history study that was based on specimens from the MMS Phase II Santa Maria samples, Kropp and Carroll (1990) found gravid females during all seasons except July and October

1987. Males exhibited a similar pattern. No data were presented on egg diameter, precluding an assessment

i of reproductive maturity in the population. One specimen collected at PJ-1 during May 1987 had both eggs and sperm. Recruitment apparently occurred at station PJ-1 during October of 1986 and 1987 and again in

January 1988.

I papillaeRtheatmarks have.sePholoe veral folds glaborraarticl is cheasr,acte pserizudoarticulat ed by haviengdjoints a shoront, ttriang he tipsularoffaci thealmedi tubeanrcante le, sho nnartaneldytrtheal

dorsal andventral tentacular cirri, andvery few papillae on the surface of the body. Further,the serrations on

i the blades of the compound setae are very distinct; in related species these are either absent or very difficult to discern.

Distribution. California, intertidal beaches; subtidal on the continental sheff and upper slope, to

I approximately 300 m.

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Figure 5.I. Photoe gIabra: A, anterior end, dorsal view; B, middle parapodium, anterior view; C,first elytron; I

D,middlelytronwi,thinsetsshowingdetaiolfsmarginalpapillae; E,superinorotopodial seta;F,

neuropodial compound falciger. (all except insets after Hartman, 1961) i

Pholoe courtneyae Blake, new species I

I

Figure 5.2

Material Examined. California, off San Francisco, Gulf of the Farallones, continental slope stations I

sampledas part of U.S. EPA 102-Site Disposal Designation Survey, R/V Point Sur, J.A. Blake, Chief Scientist:

u

Sta. 3-1, 37°23.82'N, 123°15.55'W, 1338 m, 12Sep 1991,7 paratypes CUSNM 169972); Sta. 3-4, 37°27.83'N,

123°11.60'W, 1040 m, 14 Sep 1991, 2 juveniles; Sta. 3-5, 37°27.39'N, 123°14.42'W, 1195 m, 14 Sep 1991, I

3 paratypes (USNM 169973); Sta. 3-11, 37°25.50'N, 123°15.02'W, 1225 m, 15 Sep 1991, 1juvenile; Sta. 3- II

14, 37°23.51'N, 123°21.02Tf, 1880 m, 15 Sep 1991, holotype (CASIZ 103568); Sta. 3-15, 37°24.71'N,

123°12.16'W, 1010 m, 15 Sep 1991, 4 paratypes (CASIZ 103569) on slides Sta. 3-16, 37°23.20'N, I

123°14.44"W, 1220 m, 15 Sep 1991, 5 small paratypes (LACM-AHF Poly 1718) Sta. 4-2, 37°15.29'N,

123°07.1YW, 995 m, 18Sep 1991, 1juvenile; Sta. 4-3, 37°15.13'N, 123°I0.83T¢, 1222 m, 18 Sep 1991, 1

specimen (JAB); Sta. 4-4, 37°16.13'N, 123°14.18"W, 1427 m, 18 Sep 1991 paratype (LACM-AHF Poly I

1719); Sta. 4-8, 37°13.38'N, 123°10.46'W, 1235 m, 18 Sep 1991, 2 juveniles; Sta. C-2, 37°20.81'N, I

123°02.32'W, 1025 m, 19Sep 1991, 1juvenile.---Santa Mafia Basin, offPoint Arguello, Sta. 56, 900 m (3);

Sta. 63, 930 m (2). •

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Description. Body small, linear, depressed; holotype 7.2 mm long, 1 mm wide, with 52 setigers,

I other coveredspecimens by overlapping up to 6 mmelytralong,(Fig. 1.2 5.2A); mm wide, elytra42 setigers; translucent, mostoval specimens anteriorlyjuveniles. (Fig. 5.2A, DorsumB),completely becoming

subquadrate posteriorly (Fig. 5.2C); each elytron with elongate thin, papillae, both marginal and submarginal

i onsometimes anterior upelytra, to one-fourth marginallength on posterior of elytron; elytra; eachpapillae elytral papiUa of middle simple, and slender, posteriorweakly elytrajointed, becomingwithlonger, blunt,

expanded tip bearing delicate sensory cilia (Fig. 5.2B). Additional papillae on surface of body, surrounding

i mouth, with on ventral surface, and on parapodia; parapodial papillae of several types: most short, nipplelike, but some parapodial papillae longer, f'mgerlike; body papillae with terminal sensory cilium (Fig. 5.2D).

Color white to fight tan.

I short, withProstomium thick, pear-shaped a broadly ceratophore, rounded lobecfrom ontinuing whichdistally medianasantenna taperingarises; filament; eyes minute absent; lateral medianantennae antenna

present on either side of median ceratophore (Fig. 5.2A). Palps thickened basally, directed anteriorly, tapering,

I more terminal or less papillae, smooth; 2-4 distal lateralends papillae, with sensory and 2 pairs cilia.of Facial dark jaws. tubercleTentaculophores thin, filamentous. with Proboscis dorsal and with ventral16

tentacular cirri; dorsal tentacular cirri expanded basally, tapering to long, narrow, blunt-tipped filament,

sometimes with 1 or 2 weakly developed joints; ventral tentacular cirri shorter than dorsal ones, similar in

I appearance, with 1-2 notches or joints.

Segment 2 (or buccal segment) first setigerous, with large ventral cirri (buccal cirri) displaced medially,

I broadl segments y rounsmall, ded basally, elongate,tapering basallyapicall enlarged, y bearing distally first pointed pair of elytra. (Fig. Notopodium 5.2D). Neuropodium of segmentelongate, 2 and subseque distallynt

pointed, with small surficial papillae continuing onto mid-ventrum of body; ventral cirrus from segment 3

I fascicles long, thin,oftapering setae; notosetae with 1-2simple, joints orof notches 3 types: along (1) anterior length.rowBothof about podial20lobes short,bearing serrated,thickgeniculate aciculaesetae and

beginning on posterior side of notopodium, continuing to anterior side (Fig. 5.2E); (2) about 5-8 long, thin

I alternating serrated setaewithonserrated either sideones. of shorter Neurosetae ons 5-6 (Fig.compound 5.2F); andfalcigers, (3) aboutwith 15 longer, iconspicuous thin, non-serrated knobby denticles capillarieson

blades and few f'me hairlike spinelets on tip of shaft, only visible with 1000x optics, otherwise blades and

I shaft ending Remarks. appearingThe smooth anoculate(Fig. Pholoe 5.2G).courtneyae Pygidium was withatpair of long, filamentous anal cirri. first confused with P. anocuIata Hartman, 1965

from the U.S. Atlantic continental slope. This latter species is poorly known and both the original description

I byTypes Hartman were never (1965)designated and the supplementary and the only figures one byofHartman the species andwere Fauchald of an (1971) elytron were and twobrieffalcigers and incomplete. from one

of the two specimens from Station S1-4(Hartrnan and Fauchald, 1971:plate 2, figs. f-h) that were cited among

I the Programoriginal (ACSAR) material(1983-1987) examined bynumerous Hartmanspecimens (1965). were As part identified of the that Atlantic were Continental referred toP.Slope anoculataand. Rise Th

species was one of the most abundant encountered from the Canadian Boundary to off the Carolinas and

proved to be an important component of the slope assemblages (Blake, t993, 1994; Blake and Grassle, 1994). Initial comparisons of the California Pholoe specimens demonstrated that they were very similar to

the Atlantic specimens. However, because of the inadequate nature of the published descriptions ofP. anocu/ata,

I compare it was necessary them withto esthetabCalifornia lish a morphspecimens. ological baseline The extensive for theACSAR Atlantic collections specimens were in order available to moreas cwell arefulaslya

syntype specimen of P. minuta anoculata obtained from the LACM (LACM-AHF 0069) 3. When these

I Atlantic materials were carefully studied and then compared with California specimens from the Santa Maria

] Basin and offthe Farallones, it became obvious that two separate, yet closely related species were involved.

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Figure 5.2. Pholoe courtneyae (Paratypes, Sta. 3-15, CASIZ 103569): A, anterior end, dorsal view, with only I

second elytron included for clarity; B, first and second elytra from right side of another paratype, I

with insets showing details of elytral papillae [not to scale]; C, elytron from a posterior segment; D,

parapodium from setiger 10 in posterior view, with insets of different parapodial papillae [not to •

scale]; E, short geniculate serrated notoseta; F, long, thin serrated notoseta; G, neuropodial falciger I

from middle of series.

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I A full redescription of Pholoe anoculata will be published elsewhere. Pholoe courtneyae differs from P. anoculata inbeing larger, 7+ mm long instead of 2-3 mm long, in having relatively smooth tentacular

cirriinstead of ones that are knobby (especially the ventral tentacular cirri), in having elytra with papillae that

' _ are relatively smooth along their edges instead of with minute constrictions, in having smooth, forwardly

_ directed palps instead of palps with constrictions and that are directed laterally. Furthermore, the blades and

shaft endings of the neuropodial falcigers ofP. courtneyae bear inconspicuous denticles and spinelets visible

I only undeEvr oil immersion, instead of larger, more conspicuous denticles and spines. aluation of the collections from the Faraliones has revealed the existence of two additional anoculate

species of Pholoe. One comes from middle and lower slope depths of 1800 - 3060 m. This species is small,

I less than 4 nun long and very similar to Atlantic specimens of P. anoculata. The second species is from middle slope depths (1338-2005 m). It is large, up to 12 mm long, and differs from both P. courtneyae and the

small California P.anoculata-like species in having elytral papillae that are consistently short along the entire

I bmentioned ody insteareview d of becofoPminganoculata longer. in posterior setigers. Both of these species will be treated in the above

Etymology. PhoIoe courtneyae is named for Dr. Catherine A. (Kitty) Courtney, Project Manager

I and oceanographer for several of the Farallones site selection Dr. Courtney was instrumental in

surveys.

assuring my participation in these deep-sea surveys of important benthic communities. She provided constant

encouragement and assistance throughout the course of the program. at

i Biology. Pholoe courtneyae has only been rarely encountered in slope sediments of the eastern

Pacific, probably owing to its small size and the fact that so few samples have been processed through 300 grn

i sieves. The species is likely an omnivore and probably similar in habitat preferences to its close relative P.

anoculata from the Atlantic slope. Blake (1994) determined that more than 80% of the specimens of P.

anoculata from off Cape Lookout, North Carolina occur in the upper 2 cm of the sediment.

i Distribution. California continental slope, in middle slope depths, 900 - 1880 m.

Genus PhoIoides Pruvot, 1895

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PhoIoides Pruvot, 1895. Type species: Pholoe dorsipapillata Marenzeller, 1893.

Pe.isidice Johnson, 1897. Type speeies: Peisidice aspera Johnson, 1897. Fide Pettibone, 1982.

I Paraeupholoe Hartmann-Schr6der, 1962. Type species: Johnson, 1897.Fide Pettibone, Pareupholoe fimbriata

1992.

Parapholoe Hartmann-Schrtider, 1965. Type species: Parapholoe tuberculata Hartmann-Schr6der, 1965.

I Fide Pettibone, 1992.

Diagnosis. Body small, linear, up to 48 segments. Middorsum not covered by elytra. Elytra on

I segments 2, 4, 5, 7, continuing on alternate segments to end of body; thick, subrectangular, with concentric rings and numerous, long border papillae and scattered tubercles. Prostomium and first or tentacular segment

fused, ventrally forming anterior lip of mouth, without facial tubercle. Prostomium subrectangular, median

I

3 The type specimen of Photoe min_a anocufata Hartrnan, 1965 (LACM-AttF Poly 0069) that was sent m me for

I examination came from Sta. S 1-4 (Hartman, 1965), and was labeled a paratype. I am informed that the second specimen from that station is labeled a holotype (LACM-AHF Poly 0068). However, a holotype was not

designated by Hartman (1965), nor indicated subsequently by Hartmun and Fauchald (1971). According to

the International Code of Zoological Nomenclature (ICZN, 1985), these specimens are syntypes until a lectotype

I ises ndet smignusated. t be termed A holoatype syntycapen ononltyabepardeatype sign.ateIdambyunthewilling original to design authoar.te Cthis onsespecime quentlyn, the as aspecime lectotypen I uwas ntil

the second specimen has been examined and confirmed as the same species. There is evidence that a second

cryptic species is present among the samples examined by Hartman (1965) (M.E. Petersen, unpublished),

I lending further reason to be cautious in designating a lectotype.

i 183
!

antenna with ceratophore on anterior border, with style enlarged and papillate subdistally; lateral antennae i

absent; 2 pairs of eyes. Tentacutophores with notosetae and single pair of tentacular cirri similar to median

antenna. Parapodia biramous, notopodium rounded, subc0nical; neuropodium larger, subconical. Notosetae i

simple, slender, capillary, timely spinose. Neurosetae stouter than notosetae, compound, with shafts smooth I

or spinose subdistally and blades short, falcate, smooth or faintly spinose. Ventral cirri short, subulate,

papillate. Pygidium with pair of anal cirri. •

Remarks. The genus Pholoides Pruvot, 1895 has priority over Peisidice Johnson, 1897. Pettibone I

(1992) referred the monotypic genera Paraeupholoe Hartmann-Schr6der and Parapholoe Hartmann-Schr6der

to synonymy with Pholoides after examining the hototypes of their type species, i

Pholoides asperus (Johnson, 1897)

Figure 5.3 i

Peisidice aspera Johnson, 1897:184, pl. 9, figs. 56-59, pl. 10, fig. 63.--Moore, 1908: 338; 1910: 386.-- 1

Berkeley and Berkeley, 1941:24; 1942:189; 1948:23, fig. 28.--Hartman, 1939:7; 1948:14; 1968:147,

figs. 1-3.--Lie, 1968:286.--Hartmann-Schr6der, 1977:81, figs. 17-18.

Paraeupholoefimbriata Hartmann-Schr6der, 1962:110, pl. 1, figs. 5, 11, pl. 2, figs. 6-7, 10, pl. 3 (figs. 8-9, I

12. Fide Pettibone, 1992.

Parapholoe tuberculata Hartmann-Schr6der, 1965, figs. 41-47. Fide Pettibone, 1992.

Peisidice tuberculata: Hartman and Fauchald, 1971:29.--Hartmann-Schr6der, 1977:81 .Fide Pettibone, 1992. i

Pholoe minuta: Blake, 1975a:30, figs. 3A-B (descriptions of larvae). Not P minuta Fabricius, 1780.

Pholoides aspera: Fauchald, 1977:67, fig. 16A-B.

Pholoides tuberculata: Carrasco and Gallardo, 1983:832. I

Pholoides asperus: Pettibone, 1992:18-21, figs. 10-11 (Synonymy). i

Material Examined. California, Tomales Bay, with barnacles, 15 m, (4).---Santa Maria Basin, off •

Point Buchon, Sta. BRA-27, 96-126 m, in rocks (31); off Purisima Point, Sta. R-4, 92 m (3,juveniles); Sta.

BRC-13, 92-100 m, in rocks (24).

Description. A small species, recorded up to 11 mm long, 4 mm wide, and 48 segments; more I

typically, 7-8 mm long, 2-3 mm wide, for about 35 segments. Body elongate, subrectangular in outline,

dorsoventrally flattened, elytra not covering dorsum (Fig. 5.3A); dorsum with globular tubercles and sand

grains (Fig. 5.3B), ventrum covered with small papillae. Color in alcohoI: light brown; sometimes with 1

brown spots on elytra. II

Prostomium subrectangular, fused with f'trst tentacular segment; bearing 2 pairs of eyes, posterior

pair smallest; median antenna with long ceratophore, with subdistal bulge bearing numerous thin papillae i

(Fig. 5.3B-C). Tentacular segment with tentaculophores bearing internal acicula, acicular lobe, 2 fascicles of I1

long capillary notosetae, long dorsolateral tentacular cirrus similar in form to median antenna, and additional

short tentacular cirrus (or long accessory papilla) present ventrally. Segment 2 with long, ventral cirrus (Fir. I

5.3D). Long thick patps present ventrolateral to tentaculophores. Pharynx with 9 pairs of terminal papillae Q

and 4 jaws.

Elytrophores subtriangular elytra first present on segment 2, continuing on segments 1

1

bearing oval to

4, 5, 7, and on alternate segments to end of body. Each elytron relatively stiff, with concentric rings, thickest

w

on medial side, with humerus long papillae on margins (Fig. 5.3G). Segments without elytra with dorsal

tubercles bearing small distal papilla (Fig. 5.3E). i

II

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184 i
!

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mAn dT

I

I

I

I vc F

I Figure 5.3. Pholoides aspera: A, entire animal, dorsal view; B, anterior end, dorsal view; C, dorsolateral view

of prostomium and left tentaculophore, acicula dotted; D, parapodium from segment 2, anterior

I view; posterior E, middle view; parapodium G, with dorsal tubercle, anterior view; F, middle elytral bearing parapodlum, elytron; H, neuropodial compound falciger. (A, from Pettibone, 1953; B, from

Johnston, 1897; C-H, after Pettibone, 1992)

l Parapodia with neuropodium longer than notopodium; notopodium rounded; neuropodium with

preacicular lobe bearing 2 long papillae (Fig. 5.3E-F); ventral cirri short, tapering, with short papillae. Notosetae

I include spinous numerous blades (Fig.curved 5.3H).spinous Pygidiumcapillaries; with 2 neurosetae anal cirri similar stout, incompound form to median falcigers,antenna. with smooth or weakly

Biology. Pholoides asperus is widely distributed in the eastern Pacific, occurring from the low intertidal

l to about 350 m on rocks, in crevices, and on mixed sediment types. Pelagic larvae were collected and described by Blake (1975a: identified as Pholoe minuta). The nectochaete or post-larval stage clearly exhibits

the papillated median antennae and tentacular cirri that are characteristic for this species. The rather sluggish

I nature of these larvae and the yolky appearance of the gut suggested that they were lecithotrophic.

Remarks. Pholoides asperus is the only species of this genus in the eastern Pacific. Because of its

characteristic shape, size, papillated antenna and tentacular cirri, and form of the etytra, the species is not

I likely to be confused with any other scaleworm.

Distribution. Eastern Pacific: Alaska to California, Galapagos Islands, Chile; low intertidal to 350 m.

I

i 185
I

Literature Cited I

Akesson, B. 1963. The comparative morphology and embryology of the head in scale worms (Aphroditidae, 1

Polychaeta). Arkiv f6r Zoologi 16(7): 125-163. J

Berkeley, E. and C. Berkeley. 1941. On a collection of Polychaeta from Southern California. Bulletin of the •

Southern California Academy of Sciences 40:16-40. 1

Berkeley, E. and C. Berkeley. 1942. North Pacific Polychaeta, chiefly from the west coast of Vancouver

Island, Alaska, and Bering Sea. Canadian Journal of Research Sec. D, 20:183-208. I

ml

Berkeley, E. and C. Berkeley. 1948. Annelida, Polychaeta, Errantia. Canadian Pacific Fauna. Fisheries

Research Board of Canada, Toronto 9b:(1): 1-100. •

[] Blake, J.A. 1975a. The larval development of Polychaeta from the northern California coast. IH. Eighteen

species of Ermntia. Ophelia 14:23-84.

Blake, J.A. 1975b. Phylum Annelida: Class Polychaeta. Pp. 151-243, In, Smith, R.I. and J.T. Carlton (eds.), I

Light's Manual: Intertidal Invertebrates of the Central California Coast. University of California

Press, Berkeley. I Blake, J.A. 1993. Life history analysis of five dominant infaunal polychaete species from the continental

slope off North Carolina. Journal of the Marine Biological Association of the United Kingdom 73:123-

141. I

Blake, J.A. 1994. Vertical distribution of benthic infauna in continentaI slope sediments off Cape Lookout,

North Carolina. Deep-Sea Research 1I, 41:919-927. []

Blake, [] J.A. and J.E Grassle. 1994. Benthic community structure on the U.S. South Atlantic slope off the

Carolinas: Spatial heterogeneity in a current-dominated system. Deep-Sea Research II, 41:835-874.

I

Cazaux, C. 1968. l_tude morphologique du d6veloppement larvaire d'Ann61ides Polych_tes (Bassin I

d'Arcachon). I. Aphroditidae, Chrysopetalidae. Archives de Zoologie Exp6rimentale et G-6n6rale 109;

477-543. II

Carrasco, F.D. and V.A. Gallardo. 1983. Abundance and distribution of the macrobenthic infauna of the Gulf 1

of Aranco, Chile. Internationale Revue der Gesamten Hydrobiologie 68:825-838.

1

Chambers, S. 1985. Polychaetes from Scottish waters. Part 2. Families Aphroditidae, Sigalionidae and I

Polyodontidae. Royal Scottish Museum, Edinburgh. 38 pp.

Christie, G. 1982. The reproductive cycles of two species of Pholoe (Polychaeta: Sigalionidae) off the 1

Northumberland coast. Sarsia 67:283-292.

Curtis, M.A. i977. Life cycles and population dynamics of marine benthic polychaetes from the Disko Bay m

area of west Greenland. Ophelia 16:9-58.

|

Darboux, J.-C. 1900. Recherches sur les Aphroditiens. Bulletin Scientifique de la France et de la Belgique

33:!-274. 1

U

Fabricius, O. 1780. Fauna Groenlandica. Hafniae et Lipsiae. 452 pp.

Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families and genera. Natural _l

History Museum of Los Angeles County, Science Series 28:1-190.

Fauchald, K. and P.A. Jumars. 1979. The diet of worms: A study ofpolychaete feeding guilds. Oceanography aim

and Marine Biology Annual Review 17:193-284. •

186 |
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I Franzen, A. 1956. On spermiogenesis, morphology of the spermatozoon, and biology of fertilization among

invertebrates. Zoologiska Bidrag frhn Uppsala 31:355-480, plates 1-6.

I Hartman, O. 1939. Polychaetous annelids. Part I. Aphroditidae to Pisionidae. Allan Hancock Pacific Expeditions 7(1): 1-156.

i Hartman, O. 1948. The polychaetous armelids of Alaska. Pacific Science 2:3-58. Hartman, O. 1959. Catalogue of the polychaetous annelids of the world. Part I. Allan Hancock Foundation

Publications Occasional Paper 23:1-353.

I Hartman, O. 1961. Polychaetous annelids of California. Allan Hancock Pacific Expeditions 25:1-226.

Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other North

I Atlantic areas. Allan Hancock Foundation Occasional Paper No. 28:1-378.

Hartman, O. 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation,

I University of California, Los Angeles. 828 pp. Hartman, O. and K. Fauchald. 197t. Deep-water benthic polychaetous annelids off New England to Bermuda,

and other North Atlantic areas. Part II. Allan Hancock Monographs in Marine Biology 6:1-327.

I Hartmann-Schr6der, G. 1962. Zweiter Beitrag zur Polychaetenfauna von Peru. Kieler Meeresforschungen

18(1):109-147.

I Hartmann-Schr6der, G. 1965. Zur Kenntnis des Sublitomls der chilenischen Kiiste unter besonderer Berfieksichtigung der Polychaeten und Ostracoden, Part II. Die Polychaeten des Sublitorals.

Mitteilungen aus dem Hamburgischen Zoologiscben Museum und Institut 62(Erganzungsband):59-

I 305.

Hartmarm-Schrtider, G. t977. Polychaeten aus dem Sublitoral und Bathyal vor der portugiesischen und

marokkanischen K_ste Auswertung der Fahrt (1967) von KS. "Meteor. .... Meteor" ForschungsI ergebnisse, Reihe D, 31:63-90.

Heffernan, E 1985. Demography of Pholoe minuta (Polychaeta: Sigalionidae) in Galway Bay, west coast of

I Ireland, with special reference to settlement and recruitment patterns. Marine Biology 84:323-329.

Heffernan, P. 1988. Ultrastructural and histochemical studies of the digestive system of PhoIoe minuta

(Polychaeta: Sigalionidae). Journal of the Marine Biological Association of the United Kingdom

I 68:447-464.

Heffernan, P. and B.E Keegan. 1988a. Quantitative and ultrastructural studies on the reproductive biology

I of the polychaete Pholoe minuta in Galway Bay. Marine Biology 99:203-214.

Heffeman, P.and B.E Keegan. 1988b. The larval development ofPholoe minuta (Polychaeta: Sigalionidae)

in Galway Bay, Ireland. Journal of the Marine Biological Association of the United Kingdom 68:339-

I 350.

Heffernan, P., O'Connor, B., and B. Keegan. 1983. Population dynamics and reproductive cycle of Pholoe

I minuta (Polychaeta: Sigalionidae) in Galway Bay. Marine Biology 73:285-291.

International Code of Zoological Nomenclature (ICZN). 1985. Third Edition. Adapted by the XX General

Assembly of the International Union of Biological Sciences. University of California Press, Berkeley

I and Los Angeles, xx + pp.

338

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I 187
II

Johnson, H.E 1897. A preliminary account of the marine annelids of the Pacific coast, with descriptions of •

new species. Proceedings of the California Academy of Sciences, Zoology 1(5): 153-198.

Johnston, G. 1839. The British Aphroditacea. Annals & Magazine of Natural History, London, series 1, !

2:424-441. m

Kinberg, J.G.H. 1858. Part 3: Annulater. KongIia Svenska Fregatten Eugenies Resa Omkring Jorden uner •

Bet'al af C.A. Virgin/_ren 1851-1853: Vetenskapliga Iakttagelser, Zoology, 2:1-32. Uppsala and il

Stockholm. Almquist and Wicksells.

Kropp, R. and D. Carroll. 1990. Chapter 8. Life-history studies of selected macroinfaunal invertebrates n

form the Santa Mafia Basin, California. In, Steinhauer, M. and E. Imamura (eds.), California OCS m

Phase II Monitoring Program, Year-Three Annual Report. Volume I. Pp. 8-1 to 8-24. II

Lacalli, T.C. 1980. A guide to the marine flora and fauna of the Bay of Fundy: Polychaete larvae from I

Passamaquoddy Bay. Canadian Technical Report of Fisheries and Aquatic Sciences No. 940:1-27.

Lacalli, T.C. 1981. Annual spawning cycles and planktonic larvae of benthic invertebrates from Passamaquoddy I

Bay, New Brunswick. Canadian Journal of Zoology 59:433-440.

Laubier, L. 1975. Adaptations morphologiques et biologiques chez un Aphroditien interstitiel: Pholoe •

swedmarki sp. n. Cahiers de Biologic Marine 16:671-683. II

Lie, U. 1968. A quantitative study of benthic infauna in Puget Sound, Washington, U.S.A., in 1963-1964.

[Section on polychaetes by K. Banse, K.D. Hobson, and EH. Nichols]. Fiskeridirektoratets Skrifter

serie Havunders_kelser, 14(5):230-556. II

MarenzeUer, E. yon. 1893. Zoologische Erbegnisse, 1I:Polychaten des Grundes, gesammelt 1890, 1891, und •

1892. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch- l

Naturwissenschafttiche Classe 60:25-48, 4 plates.

Moore, J.P. 1908. Some polychaetous annelids of the northern Pacific coast of North America. Proceedings I

of the Academy of Natural Sciences of Philadelphia 60:321-364. U

Moore, J.E 1910. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern •

California in 1904. Potynoidae, Aphroditidae and Segalionidae. Proceedings of the Academy of II

Natural Sciences of Philadelphia 62:328-402, pls. 28-33.

Pettibone, M.H. 1982. Classification of Polychaeta. In: Parker, S.E (ed.). Synopsis and Classification of I

Living Organisms. McGraw-Hill. pp. 3-43. U

Pettibone, M.H. 1992. Contribution to the polychaete family Pholoidae Kinberg. Smithsonian Contributions II

to Zoology No. 532:1-24. I

Pleijel, E 1983. On feeding inPholoe minuta (Fabricius, 1780) (Polychaeta: Sigalionidae). Sarsia 68:21-23.

n

Pruvot, G. 1895. Coup d'oeil sur la distribution grnrrale des invertrbr_s dans la rrgion de Banyuls (Golfe du I

Lion). Archives de Zoologie Exprrimentale et Grnrrale Paris, srries 3, 2:629-658.

Sveshnikov, V.A. 1967. Arehiannelid and polychaete larvae from Possjet Bay, Sea of Japan. Issledovania g

Fauni Morei, Academia Nauk SSSR, Zoological Institute 5(13): 125-159. [In Russian] U

Wolf, ES. 1986. A new genus and species of interstitial Sigalionidae and a report of the presence of venom •

glands in some scale-worm families (Annelida: Polychaeta). Proceedings of the Biological Society of I

Washington 99:79-83.

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6. FAMILYSIGALIONIDAEKINBERG, 1856

|

James A. uYBlake 1

!

i Introduction

i The Sigalionidae includes highly active scaleworms that have along, narrow bodies, with numerous segments andpairs of elytra. Most species are large, andthey are carnivorous polychaetes that are also able

toburrowrapidly into sediments. These scaleworms are widely distributed,with many species being relatively

i common andfamiliar. Sigalionids occur from the intertidal to the deep sea.

i Morphology

i The bodies of sigalionids axe elongate and narrow, with numerous segments (up to 300). The elytra either cover the dorsum completely, or leave the middorsum uncovered. The dorsum and elytra are sometimes

covered with a thick layer of sand grains. The ventral surface of the body may be smooth or thickly papillated.

The prostomium is generally small, somewhat oval or pentagonal in shape, and is fused with the first tentacular

i segment, pair long, tapering palps are present are usually antennae present.

A of ventral and there 3 The

median antenna is longest and has a distinct ceratophore and sometimes diagnostic auricular lobes or ctenidia

are present as well. The lateral antennae tend to be short. The prostomium may have 1 or 2 pairs of eyes,

I nuchal be or absent. The first is vestigial and bears an acicula, a bundle of organs may present segment

notosetae, and dorsal and ventral tentacular cirri. It is called the tentaculophore and is directed forward and

lies dorsal to the palps. The tentaculophores form the upper lip of the mouth and may bear a facial tubercle.

I The second segment is the buccal segment and bears the first pair of elytra, biramous parapodia, and the lateral lips of the mouth; the parapodia bear long ventral buccal cirri that are longer than the ventral cirri of

subsequent segments. Segment 3 forms the lower lip. The pharynx is eversible, large, and muscular, with a

I papillated oral opening and 2 pairs of jaws.

The elytra are attached to knoblike elytrophores on segments 2, 4, 5, and 7, on alternate segments on

24 or 27, and then on every segment. The lateral margins of the elytra may bear papillae and the surface may

1 bear micmtubercles. Segments lacking elytra bear knoblike dorsal tubercles and usually lack dorsal cirri.

w

Branchiae are cirriform, heavily ciliated, and attached to the lateral sides of the elytrophores and dorsal

tubercles on all segments except a few anterior ones. Ciliated grooves occur along the dorsolateral sides of the

I body; these contain ciliated ctenidia.

grooves

Parapodia are supported by aciculae; notopodia are generally smaller than neuropodia, generally

i elongate and narrow and bear papillae or stylodes and long, fanlike spreading fascicles of simple capillary setae. Neuropodia are triangular to conical and surrounded by basal bracts 2with fringes of papillae and distal

stylodes. Neurosetae are compound spinigers or falcigers, some of which may be long and multiarticulated.

Ventral cirri are narrow and tapering to fine tips; sometimes they bear papillae. The pygidium bears a pair of

I long anal cirri.

ENSR Consulting and Engineering, 89 Water Street, Woods Hole, Massachusetts 02543

I _Pre-and postsetal parapodial lamellae that encompass setae and sometimes bear papillae.

I 189
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Taxonomic History I

i

Approximately 15 genera of Sigalionidae may be considered as valid according to Fauchald (1977). •

Major revisions of some of these genera, have been published by Pettibone (1969, 1970a-c, 1971a-b, 1992a) t

and Mackie and Chambers (1990). Only ten species appear to have been recorded from California. Six dlm

species are described in this Atlas. 1

I

Major faunistic treatments of sigalionids may be found in Hartman (1939) for the eastern Pacific,

Day (1967) for South Africa and Wolf (1984) for the Gulf of Mexico. The genera Pholoe and Pholoides have !

been removed from the Sigalionldae and referred to the Pholoidae. Readers are referred to Pettibone (1992b) •w

and Blake (1995, this volume) for a review of pholoid systematics and biology. The main characteristics that

differentiate the two families are summarized in Chapter 5, Table 5.t. I

Biology I

Sigalionlds are a group of burrowing carnivores that generally live in sand or mud. They range from •

the intertidal zone to continental shelf and slope depths. Some species have been recorded from coarse 1

substrates including fine gravel, coarse sand, and associated with coralline algae, shells, and bryozoans

(Pettibone, t970a). Species of Sthenelanella are unique among the Sigalionidae in having notopodial spinning •

glands that produce threads used to construct tough fibrous tubes. Species of the genus Psammolyce attach 1

sand grains to their bodies and elytra producing an effective protective covering. Little is known concerning

the feeding habits of sigalionids, although they axe thought to feed on small invertebrates (Fauchald and •

Jumars, 1979). I

The reproductive biology of sigalionids are poorly known, however, Pettibone (1970a) reports that

large specimens ofEuthalenessa oculata (Peters) have been observed swimming at the surface. Many species n

have been reported as having large, yolky eggs. Blake (unpublished) has observed plankfic larvae of a species l

of Sthenelais in northern California. The trochophore, metatroehophore, nectochaete stages of Sthenelais

boa were described by Cazaux (1968). It would appear that this species has a short, pelagic phase during its •

development. I

Key to the Genera and Species from California I

IA. Lateral antennae located on prostomium; antennae small, without ceratophores (Fig. 6.6A); elytra

with bipinnately branched lateral filaments (Fig. 6.6B) ........................................ Sigalion spinosus

1B. Lateral antennae, when present, fused to tentacular segment; median antenna with large ceratophore •

(Fig. 6.3A) ...................................................................................................................................... 2 II

2A. Median antenna with basal auricles (Fig. 6.1A) ............................................................................. 3 I

2B. Median antenna without auricles .................................................................................................... 9

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I 3A. Middle body segments with notopodial spinning glands producing long, threadlike setae extending far beyond parapodia (Fig. 6.5F) ..................................................................... Sthenelanella uniformis

I 3B. Notopodial spinning glands absent .................................................................................................. 4

4A. Segment 3 with long dorsal cirri ......................................................................... Genus Neoleanira 3

I 4B. Segment 3 without dorsal cirri ......................................................................................................... 5

5A. Neurosetae including at least some falcigers ............................................... Genus StheneIais ...... 6

I 5B. Neurosetae all spinigers ...................................................................................... Genus Sthenolepis 4

I 6A. Middle group (C-shaped) of compound neurosetae include stout, short-bladed, bidentate falcigers (Fig. 6.1F); lateral antennae present or absent ................................................................................ 7

6B Middle group (C-shaped) of compound neurosetae without short-bladed falcigers; lateral antennae

I absent ............................................................................................................ Sthenelais verrucuIosa

I 7A. Ventral surface of body and ventral sides of parapodia densely papillated (Fig. 6.2A) .....................

........................................................................................................................... Sthenelais berkeleyi

7B. Ventral surface smooth or with very few, sparse papillae (Fig. 6.2B) ............................................ 7

!

8A. Stout compound neurosetae of middle segments numbering 5 or more; first pair of elytra unpigmented,

i 8B. subsequent Stout compound ones pigmented neurosetae inofvarious middle patterns segments....................................................... numbering only 1-2; all elytra semi-translucent, Sthenelaisfusca

lacking pigmentation, but may be encrusted with rusty colored sediment particles ...........................

i ...................................................................................................................... StheneIais tertiaglabra

9A. Dorsal cirri present on segment 3; elytra encrusted with sand .......................... Genus Psammolyce s

I 9B. Dorsal cirri absent on segment 3; elytra not encrusted with sand ............................ Genus Leanira 6

!

I 3Neofeanira areolata (Mclntosh, 1885) has been recorded from the California continental slope (Pettibone, 1970c).

Leanira catcis Hartman, 1960 is a synonym.

I _Twospeciereferred s of Sthe to note Neoleanira pis werebyrecPeUi ordedbone fro(m1970c) California and Sb.yfimbr Hartma iaruran (196 (Hartman, 8): S. are1939). ofata (Mcl The nlatter tosh)species , subsequently ranges

from southern California to westernMexico from the intertidal to continental shelf depths.

_Psammolycespinosa Hartman, 1939has been recorded from southern California. This speciesismore widelydistributed

I 6Leanira a/ha in shMoore, allow su1b9tropical 10 is wide areasspread of theineastern continenPacific talslo(Hartman, pe depths from 1939;Oreg 1968)on. to southern California (Pettibone,

1970b).

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Descriptions of Species I

Genus Sthenelais Kinberg, 1855

==

Type Species. Sthenelais helenae Kinberg, 1855, by monotypy. 1

Diagnosis. Body elongate, vermiform, with numerous segments; middorsal ridge on some anterior •

segments, bordered by few pairs of small ctenidia. Elytra on segments 2, 4, 5, 7, and alternate segments to 27, II

then continuing on all following segments. Elytra large, covering dorsum, with microtubercles and lateral

fringes of papillae. Dorsal tubercles on segments 3, 6, 8, and alternate segments to 26. Prostomium rounded, •

fused to first or tentacular segment; median antenna with cylindrical ceratophore beating lateral auricles and I

terminal style; lateral antennae fused to inner dorsal sides of tentacular parapodia; 4 eyes present; palps

emerging ventral to tentacular parapodia; paired oval nuchal organs present. Tentacular parapodia extending •

anteroventral to prostomium, each with a pair of tentacular cirri, single acicula, 2 bundles of capillary setae, 1

L-shaped inner tentacular lobe with ciliated ridge and fused to inner palpal sheath; ciliated dorsal ctenidium.

- Parapodia of second or buccal segment extending anteriorly,with ventral buccal cirri longer than those following; •

small ctenidia on lateral lips and medial to ventral cirri on some anterior segments. Third segment with dorsal I

tubercles fused to posterior sides of elytrophores of segment 2. Neurosetae of segments 2 and 3 compound

falcigers with articulated blades and bifid tips. •

Branchiae cirriform, ciliated, on external borders of all elytrophores and dorsal tubercles except for a 1

few anterior ones. Parapoctial ctenidia cup-shaped, ciliated, 3 per parapodium, from segment 2. Parapodia

with accessory bracts and stylodes. Notopodia clavate, with bracts nearly encircling acicular lobes. Notosetae []

numerous, arranged in semicircular row and directed posterodorsally, finely spinous, and tapering to capillary 1

tips. Neuropodia with conical acicular lobes; bilobed C-shaped posterior bracts directed anteriorly on upper

and lower margins; crescent-shaped anteroventral and anterior upper bracts. Neurosetae arranged in 3 groups: •

(1) upper group with anterior upper bracts, (2) C-shaped group of stouter neurosetae within posterior bracts, I

and (3) lower arched group of more slender neurosetae within anteroventral bracts. Neurosetae simple spinous

(in upper group) and compound falcigerous with some blades articulated and tips bifid; distal parts of stems •

smooth and spinous. Ventral cirri subulate, with outer basal knobs. Pharynx with 11pairs of papillae and 2 |

pairs of jaws.

Remarks. This diagnosis follows Pettibone (1971a-b). Sthenelais can be distinguished by three •

antennae and well-defined basal auricles, no dorsal cirrus on segment 3, and neurosetae that include simple II

spinous capillaries and two kinds of falcigers: short, stout ones with bidentate appendages and longer, more

slender ones with articulated appendages. StheneIais differs from other eastern Pacific genera by the characters []

presented in the key. Four species are known from the eastern Pacific, all of which occur in the general II

vicinity of the Santa Maria Basin.

Sthenelaisfusca Johnson, 1897 I

Figure 6.1 !

Sthenelaisfusca Johnson, 1897:185-186, pl. 9, figs. 60-61, pl. 10, fig. 64.--Monro, 1933:16.--Berkeley and

Berkeley, 1935:767; 1941:26.--MacGinitie, 1935:690.--Hartman, 1939:61, pl. 13, figs. 163-162 1

(in part, not fig. 60, fide Pettibone, 1971b).--Pettibone, 1971b:1394-1395, figs. 1-3.--Blake,

1975:177, figs. 83-84. Not Johnson, 1901:397.--Pettibone, 1953:73. (=S. berkeleyi, fide Pettibone,

1971b). I

192 |
!

DtC man

A

I luricte 2:

!

| °

'0 C D E

Figure 6.1. Swit thenelaisfusca: A, anteriorend, dorsalview; B, anteriorparapodium,posteriorview; C, elytron h insets showing tubercles andpapillae;D, simple neuroseta;E, superiorcompoundneuroseta;

F,stoutneuropodialfalciger.(A, afterBlake, 1975;B-C, afterPettibone,I97lb; D-F, afterHartman,

i 1939)

!

Material Examined. California, Bodega Harbor, subtidal, coll. Nov. 1975 (2).

I alcohol: Des pinkcription. with elytrAa elarlegan ge species tly pig,mente up tod13(s0eembelow). m long, Ve 5-7ntrmmal suwride, faceandsmooabthout . E2ly00trasegments. ellipsoid, Cconolocave r in

on anterior margin (Fig. 6.1C); rusty-brown in coloration, with some considerable darkpigment forming

I reticulated leaving lateral patterns; bare areas; first pair microtubercles of elytra relativlow, elysubconical; unpigmented; lateral withborder micrsotubercles with row ofunifor fringing mly distrib papillaeuted, andbu3-t

6 submarginal rows.

I Prostomium with large lateral auricles (Fig. 6.1A), ciliated ventrally; lateral antennae short, cirriform; 4 eyes partially concealed by auricles, equally spaced, anterior pair larger than posterior pair; palps extending

to about segment 8; nuchal organs large, located at posterior border of prostomium, at anterior border of

I mi ventral ddorsaltentacular ridge. Tenta cirri cul about ar par halfapoasdialongwithas dors dorsal al tentac cirri; uinner lar cirtentacular ri subequallobeto about mediansame antenna length(Figas. ventral 6.1A);

tentacular cirrus, fused to shorter, rounded inner palpal sheath; dorsal ctenidium elongate to oval.

I Parapodia of segments 2 and 3 with pair of small lateral ctenidia. Compound falcigerous neurosetae with bifid tips and 3-10 articles on blade; upper and lower setae more slender than middle ones; blades longest

in upper setae, shortest in lower ones.

!

i 193
!

!

I

Parapodia of anterior and middle body segments similar. Notopodial bracts papillated, with lower i

anterior papillae or stylodes longer than dorsoposterior ones. Neuropodial acicular lobes with stylodes only

in anterior parapodia. Upper lobes of bilobed posterior bracts with several stylodes on upper anterior part, 1

lower lobes without stylodes. Anterior upper bracts indistinct, anteroventral bracts low, not fimbriated. Cshaped group of stout neurosetae with short blades and smooth stems (Fig. 6.1F). Upper anterior group of

neurosetae including simple spinous capillaries (Fig. 6.1D)and compound falcigers with blades having 3-5 •

articles (Fig. 6.1E) and shafts with 4-6 rows of minute spinelets. Anteroventral slender neurosetae with 1-4 • i

articles on blades and 3-5 rows of spinelets on shaft.

Remarks. Sthenelaisfusca is distinguished from other eastern Pacific congeners in having many 1

heavy, non-articulated bifid falcigers inthe neuropodia, while at the same time having a smooth, non-papillated i

ventral surface. The specimens from Bodega Harbor exhibit a rather elegant body coloration in which the

unpigmented first pair of elytra contrast with the following elytra that are heavily colored with mottled brownish i

patterns. ll

Biology. An intertidal to shallow-water species, common under rocks, among algal holdfasts, and i

rhizomes of Phyllospadix. 1

among

II

Distribution. Central California to Peru.

I

Sthenelais berkeleyi Pettibone, 1971 i

Figure 6.2 I

Sthenelais berkeleyi Pettibone, 197lb: 1395-1400, figs. 4-7.--Blake, 1975:177, fig. 85.---Banse and Hobson,

1974:35, figs. 6k-m. I

lI

Material Examined. California, Santa Mafia Basin, offPort San Luis, 49 m (1).

Description. A large species, up to 400 mm long, 12 mm wide, and about 270 segments (Pettibone, 1

1971b). Ventral surface thickly covered with small papillae, these continuing onto ventral sides of parapodia. i

Elytra ellipsoid, concave on anterior margin; pale to rusty-brown; microtubercles low conical, uniformly 1

distributed, but leaving lateral bare areas, and some additional bare areas on more posterior elytra; lateral 1

borders with row of fringing papillae and 4-7 submarginal rows (Fig. 6.2B); sometimes with additional I

minute sensory papillae.

I

large lateral auricles, ciliated ventrally; lateral antennae short, subulate; 4 eyes I

Prostomium with

partially concealed by auricles, trapezoidally arranged, with anterior pair larger than posterior pair; palps

i

extending to about segment 7-8; nuchal organs large, located at posterior border of prostomium, at anterior i

border of middorsal ridge. Tentacular parapodia with dorsal tentacular cirrus subequal to median antenna; 1

ventral tentacular cirri about half as long as dorsal cirri; inner tentacular lobe about same length as ventral

tentacular cirrus, fused to shorter, rounded inner palpal sheath; dorsal ctenidium elongate to oval. I

Parapodia of segments 2 and 3 directed anteriorly. Compound falcigerous neurosetae with bifid tips I

and 3-11 articles on blade; upper and lower neurosetae more slender than middle ones; blades longest in upper

setae, shortest in lower ones.

Parapodia of anterior and middle body segments similar. Notopodial bracts papillated (Fig. 6.2A), i

with lower anterior papillae longer than dorsoposterior ones. Neuropodial acicular lobes with stylodes only

I

194 I
!

|

dTu

I A

, posterior C D -- I

(_ vC ,o

I

i Figure 6.2. Sthenelais berkeleyi: A, anterior parapodium; B, elytronwith insets showingtubercles andpapillae;

' I C-E, compound neurosetae from middle group as insets. (all after Pettibone, 1971b)

!

on anterior parapodia. Upper lobes of bilobed posterior bracts with numerous stylodes on upper and lower

i anterior parts. Anterior upper bracts indistinct, anteroventral bracts low, not fimbriated. C-shaped group of stout neurosetae with blades short to longer, with 1-4 articles and smooth stems (Fig. 6.2C-E). Upper anterior

group of neurosetae including simple spinous capillaries and compound falcigers with blades having 7 articles

and shafts with about 4 rows of minute spinelets. Anteroventral slender neurosetae with 1-4 articles on

I blades.

Remarks. The details that distinguish Sthenelais berkeleyi from the closely related S.fusca were

I defined presentedbyinPettibone her paper.(1971b), The twoandspecies the descriptions are similar of in the having twomany speciesheavy, presented uniarticulated here closelyandfollows bifid falcigerous the order

neurosetae. S. berkeleyi, however, has a thickly papillated ventral surface instead of one that is smooth.

I Further, both the the upperbilobed and lower posterior parts,bracts whereas of theinneuropodia S.fusca, thereinS.areberkeleyi a few stylodes are provided in the with uppernumerous part and none stylodes on theon

lower part.

I depths fromBiolosediments gy. Little that is known containconcerning gravel or the include biologyshellsof this mixedspecies. with mud. It hasThe mostlyspecimen been taken examined in shallow here

was taken from sandy silt sediments.

I Distribution. British Columbia to southern California; intertidal to 110 m.

!

i 195
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|

Sthenelais verruculosa Johnson, 1897

Figure 6.3 I

Sthenelais verruculosa Johnson, 1897:187, pl. 9, fig. 62, pl. 10, fig. 65.--Treadwell, 1914:184.--Berkeley, •

1923:216.--Berkeley and Berkeley, 1942:189.--Hartman, 1939:62-63, pl. 14,figs. 167-175; 1961:10; 1

1968:167-168, figs. 1-5.--Hartman and Reish, 1950:8. Not Pettibone, 1953 (=S. tertiaglabra). Fide

Banse and Hobson, 1974. Ii

Material Examined. California, Santa Maria Basin, off Point Sal, Sta. R-4 (1); off Point Arguello,

Sta. 58, 99 m (1) II

1

Description. A moderate to large species, at least up to 75 mm long, 4.5 mm wide, and with at least

150 segments; a complete specimen from Sta. R-4 is 48 mm long, 3.0 mm wide, with 118 segments. Color in •

alcohol: dull yellow to light tan, with elytra semi-transparent, micmtubercles of posterior segments sometimes 1

appearing as orange spots on elytra. Ventral surface smooth. Elytra suborbicular to subreniform, concave on

anterior margin (Fig. 6.3B); with microtubercles more or less coveting most of elytral surface; microtubercles ll

low, conical to globular, some pointed, not surrounded by flocculent sediment as in S. tertiaglabra; lateral and 1

sublateral margins with row of fringing papillae (Fig. 6.3B).

Prostomium rounded, with broad lateral auricles (Fig. 6.3A), ciliated ventrally; lateral antennae absent; •

4 eyes, equally spaced, both pairs about same size; palps extending to about segment 10; nuchal organs large, 1

located at posterior border of prostomium, at anterior border of middorsal ridge. Tentacular parapodia with

dorsal tentacular cirrus subequal to median antenna; ventral tentacular cirri about one-half as long as dorsal •

cirri; dorsal ctenidium elongate to oval. Setae numerous, long, finely serrated capillaries. 1

Parapodia of segments 2 and 3 with pair of small lateral ctenidia. Compound neurosetae including

(1) a few anterodorsal falcigers with elongate, narrow blades having 1-2 articles, distally bifid, and with shaft •

ornamented with rows of delicate spines; (2) these grading ventrally into smooth, compound falcigers with

less spinose shafts, but with numerous (9-11) and more distinct articles on blades.

Parapodia of anterior and middle body segments similar (Fig. 6.3C-D). Branchiae from segment 4. _l

Notopodial bracts bearing a row of slender papillae on anterodorsal face (Fig. 6.3D); fewer in posterior

segments. Neuropodial acicular lobes with row of papillae. Neurosetae arranged in 3 groups: (1) upper

anterior group of 2-3 simple spinous capillaries (Fig. 6.3E), sometimes absent; (2) main C-shaped group of II

compound falcigers with long, bifid-tipped blades having 9-10 articles (Fig. 6.3F); stout, short-bladed falcigers

absent; (3) numerous anteroventral slender compound fatcigers with long blades and slender, delicate bifid

tips. I Remarks. S. verruculosa differs from S. fusca, S. berkeleyi, and S. tertiagIabra in completely

lacking the stout, short-bladed, and bidentate falcigers in middle andposterior neuropodia. As an offshore

species, S. verruculosa is most readily confused withS, tertiaglabra. The latter species typically has arusty- l

colored floccular material surrounding the microtubercles on the elytra. This substance is lacking in S.

verruculosa.

Biology. Sthenelais verruculosa is only known from subtidal depths ranging to about 100 m in 1

1

sediments of mixed sand and silt. The specimen from Sta. R-4 had been collected in October 1988, and its me

coelom was filled with numerous small oocytes. Hi

Distribution. Western Canada to Mexico; shallow subtidal to about 100 m. 1

am

!

I
!

!

!

I l_ B

I

,I t

i

I ." "' .7"]{":,. br

I

li

!

| _°

|

I Figure 6.3. Sthenelais verruculosa: A, anterior end, dorsal view; B, elytron with insets and showing papillae

tubercles; C, anterior parapodium in posterior view; D, middle parapodium in anterior view; E,

simple neurosetae; F, compound neuroseta with 9 articulations. (all after Hartman, 1939)

!

,!

i 197

i
!

Sthenelais tertiaglabra Moore, 1910 m

Figure 6.4

.!

Sthenelais tertiaglabra Moore, 1910:395-398, pl. 33, figs. ll3-120.--Hartman, 1961; 1968:165-166, figs.

1-6.--Banse and Hobson, 1974:35, figs.6 n-o. •

Sthenelais hancocki Hartman, 1939:65-67, pl.16, 188-202. l

Sthenelais verruculosa: Pettibone, 1953:75-76, pl. 38, figs. 344-354. Not Johnson, 1897. Fide Banse and

Hobson, 1974. m

Material Examined. California, off Half Moon Bay, Sta. 2-1, 37°28.18'N, 122°51.15'W, 100 m m

(l).---Santa Maria Basin, off Point Buchon, Sta. 12, 98 m (1); off Point Sal, Sta. PJ-3 (1, 1); Sta. PJ-7 (1, 1); m

Sta. PJ-lO (1); PJ-11 (1); off Purisima Point, Sta. 42 (1); off Point Arguello, Sta. 65, 107 m (1); off Point m

Conception, Sta. 79, 98 m (1). m

Description. A moderate to large species, up to 133 mm long, 4-5 mm wide, with up to 182 segments m

(Pettibone, 1953 as S. verruculosa). Color in alcohol: dull yellow to colorless, appearing pale, with elytra

semi-transparent. Ventral surface smooth. Elytra suborbicular to subreniform, concave on anterior margin;

with microtubercles confined to medial halves of most elytra, but leaving anterior and lateral bare areas (Fig. m

6.4B); microtubercles low, conical to globular; lateral borders with row of fringing papillae and some in

m

marginal rows; with rust-colored flocculent material around tubercles, il

Prostomium rounded, with small lateral auricles (Fig. 6.4A), ciliated ventrally; lateral antennae absent; |

4 eyes, equally spaced, both pairs about same size; palps extending to about segment 8; nuchal organs large,

located at posterior border of prostomium, at anterior border of middorsal ridge. Tentacular parapodia with m

dorsal tentacular cirri subequal to median antenna (Fig. 6.4A); ventral tentacular cirri about one-fourth as 11

long as dorsal cirri; dorsal ctenidium elongate to oval. Setae numerous, long, finely serrated capillaries.

Parapodia of segments 2 and 3 with pair of small lateral ctenidia. Compound neurosetae including [mR

(1) a few anterodorsal falcigers with elongate, narrow blades having 1-2 articles, distally with minute bifid []

tips, andshaft ornamented with numerous rows of heavy andconspicuous spines (Fig. 6.4C); (2) these grading

ventrally into smooth, compound falcigers with less spinose shafts, but with numerous (6-9) and more distinct •

articles on blades. l

Parapodia of anterior and middle body segments similar. Notopodial bracts beating 2-6 papillae on

anterodorsaj face. Neuropodial acicular lobes with few sparse papillae. Neurosetae arranged in 3 groups: (1) m

Upper anterior group ofsimple spinous capillaries (Fig. 6.4D); (2) main C-shaped group of compound falcigers,

larger, but not stout as in S.fusca, including 1-3 short to moderately long bladed bifid falcigers (Fig. 6.4F),

and several unidentate falcigers with articles on blades (Fig. 6.4E); (3) numerous anteroventral slender compound m

falcigers with slender, delicate bifid tips (Fig. 6.4G-H).

Remarks. Superficially, S. tertiaglabra has a less robust, and more fragile appearance than either S.

fusca or S. berkeleyi. Unlike these species, S. tertiaglabra lacks lateral antennae; has unusual falcigers in m

segments 2-4 that bear heavy, almost knobbed spines on the shaft that are possibly analogous to the spinous

simple setae found in subsequent segments; and has only 1-3 short, bifid falcigers in middle and posterior

neuropodia. These latter setae are not as numerous, nor as stout as those found in S.fusca and S. berkeleyi, m

There also appears to be less development of parapodial papillae or stylodes in S. tertiaglabra. The rustycolored sediment that accumulates on the elytra of some species is apparently not present in the other species

of Sthenelais found on this coast, m

Biology. A shelf species, occurring in muddy sediments.

Distribution. British Columbia and Puget Sound to southern California; in shelf depths to about 300 m. I

198 m
I Figure 6.4. Sthenelais neuroseta from tertiaglabra segment: A,2;aD, nteri simple or end,spinose dorsal neuroseta; view; B, elytron E, slender, with details unidentate of tubercles; neutropodial C, cofalciger; mpound

F, stout bidentate neuropodial falciger; G-H, anteroventral compound neuropodial falciger. (A-D

i after Moore, 1910; rest original)

I Genus StheneIaneUa Moore, 1910

Type Species. SthenelanelIa uniformis Moore, 1910, by monotypy.

I Synonym. Euleanira Horst, 1916.

Diagnosis. Body slender, dorsoventrally compressed, with up to 75 segments. Elytra numerous, on

i segments 2, 4, 5, 7, then alternating to 25 or 27, continuing on following segments to end of body. Branchiae

short, conical, lateral to dorsal tubercles or elytrophores from segment 2. Without dorsal cirri. Prostomium

rounded, fused with first tentacular segment; ceratophore of median antenna with lateral auricles and short

i style; lateral antennae very short, fused to tentacular parapodia, without sheaths; 2 pairs of eyes present on

i 199
!

raised ocular cushions; tentacular parapodia uniramous, extending anteriorly medial to palps, each with single I

acicula, 2 tentacular cirri, well-developed fan-shaped fascicles of capillary setae, and medial tentacular lamella.

Parapodia of segments 2-4 directed anteriorly, buccal segment (2) with ventral buccal cirrilonger than following II

ventral cirri. Parapodia biramous, with notopodial ctenidia; without parapodial stylodes. Notopodia with l

conical acicular lobes and inflated rounded upper lobes; notosetae simple, spinous capillaries; notopodial

spinning glands from about segment 14-16, producing long, threadlike notosetae. Neuropodia with rounded •

presetat and postsetal lobes. Neurosetae all compound, generally with short blades; some with spinous shafts. I

Ventral cirri slender, elongate. Pharynx with 13 or more pairs of distal papillae and 2 pairs of interlocking

teeth. Worms inhabiting long, tough, fibrous tubes reinforced from threads ofnotosetae secreted by spinning •

glands. I

Remarks. The genus SthenelaneUa was revised by Pettibone (1969) to include two species. The

type species, S. uniformis Moore, 1910 is an eastern Pacific species, originally described form off California.

Species of Sthenelanella are unique among the sigalionids by having notopodial spinning glands that produce

long, silky threadlike strands of setae.

I

Sthe.elanella .niformis Moore, 1910

Figure 6.5 I

Sthenelanella uniformis Moore, 1910:391, pl. 33, figs. 105-112.--Hartman, 1939:69, pl. 18, figs. 226-231; •

1961:54; 1968:169, figs. 1-6.--Reish, 1968:72.--Pettibone, 1969:431-434, figs. 1-3. II

SthenelaneUa atypica Berkeley and Berkeley, 1941:26, pl. 5, figs. 1-3. Fide Pettibone, 1969.

Material Examined. California, Santa Mafia Basin, off Point Sal, Sta. R-8 (1); Western Santa 1

i

Barbara Channel, Sta. 79, 98 m (1); Sta. 85, 113 m (1).

Description. Specimen from Sta. R-8 complete, 23 mm long, 2 mm wide, for 75 segments; recorded []

up to 26 mm long and 2-3 mm wide (Pettiborte, 1969). Color in alcohol: light tan, with mottled, rust-colored l

pigment on some anterior elytra. Elytra delicate, transparent; first pair round, with fringe of short, crowded

papillae on anterior margin (Fig. 6.5C); subsequent elytra subreniform (Fig. 6.5D) to oval, smooth or with •

scattered sensory papillae on lateral margin. U

Prostomium recessed into segment 2; median antenna with lateral auricles in middle of ceratophore

(Fig. 6.5A); with 2 pairs of eyes, anterior pair larger than posterior pair (Fig. 6.5A-B). Dorsal tentacular cirri •

subequal to median antenna; ventral tentacular cirri shorter, subequal in length to ventral buccal cirri of

segment 2; lateral antennae short, subulate, on inner dorsal bases of tentacular parapodia. Capillary setae of

tentacular parapodia f'mely spinous or smooth; with elongate tentacular lamellae medial to setal bundles (Fig. •

6.5B). Neurosetae of segments 2-4 compound, with rows of minute spines on shaft and elongate, slender and II

slightly hooked blades (Fig. 6.5G).

Notopodia forming conical acicular lobes and inflated rounded upper lobes with ciliated ctenidia; 2 1

additional ctenidia between notopodia and branchiae. Notosetae all spinous capillaries, arranged in spreading

fascicles. Notopodial spinning glands from about segment 14; slender threads extending far beyond body

(Fig. 6.5F). Neurosetae with short blades (Fig. 6.5H), upper ones rod-shaped, rest conical and pointed.

Ventral cirri short, with bulbous base, tapering apically. Pharynx with 2 pairs of chitinous teeth and 13 or

more pairs of papillae.

Remarks. SthenelaneUa uniformis is readily identified by the mottled red spots on the anterior pairs i

of elytra and by the long, silky thread-like notosetae emerging from the notopodial spinning glands. !

i
I

I

_ i '_hI' mAn ..DtC

tentacular lamella /¢_.. _- _

\ )

• . f. • "._,.._

I \_ ,,.-_ ,..,

'_! I _ _///_._v,c .,,,,,.(_

4' I , _ ....... ...__ _._

: _E ..i_ "_ _ H

!

Figure 6.5. Sthenelanella uniformis: A, anterior end, dorsal view; B, anterior end, ventral view; C, first elytron

I with inset showing marginal papillae; D, fifth elytron; E, third parapodium, anterior view; F, middle parapodium, anterior view; G, neurosetae from segment 2; H, neuropodial falcigers from

middle parapodium. (A-C, E-G, after Pettibone, 1969; D, H, after Hartman, 1939)

I

I

Biology. S. uniformis inhabits long tubes that have thickened wails, that are covered with mud or

I sand andDisreinforced by the long felt or setai threads produced from the spinning glands. tribution. Central California to Ecuador; intertidal to 113 m.

I

i 201
!

Genus Sigalion Audouin and Milne Edwards, in Cuvier 1830 I

Sigalion Audouin and Milne Edwards, in Cuvier 1830:207. Type species: Sigalion mathildae Audouin and 'l

Milne Edwards, in Cuvier 1830:207, designated by Hartman, I959. II

Thalenessa Baird, 1866:34. Type species: Sigalion edwardsi Kinberg, 1856:387, by original designation.

Fide Mackie and Chambers, 1990. j

Eusigalion Augener, 1918:112-113. Type species: Eusigalion vazensis Augener, 1918:113-118, fig. 3, pl. m

14, figs. 44-46, by monotypy. Fide Mackie and Chambers, 1990.

Diagnosis. Body elongate with numerous segments. Elytra on segments 2, 4, 5, 7, and alternating I

segments to 27, then continuing on all following segments. Elytra with lateral fringe of bipinnate papillae.

Dorsal tubercles on segments 6, 8, and alternate segments to 26. Prostomium subrectangular to subpentagonal,

situated dorsally on segments 1 and 2; with 3 small antennae lacking ceratophores; lateral antennae arising n

from anterior margin; median antenna arising from dorsal surface of prostomium; 4 small subdermal eyes

present; palps long, tapering, arising ventral to tentacular parapodia. Pharynx with a pair of jaws, 2 pairs of

oblique muscular ridges, and up to 2 middorsal papillae. Tentacular parapodia anteriorly directed, each

bearing a pair of short tentacular cirri, 2 aciculae, and 2 groups of long, thin, bifid simple setae. Pair of

n

nuchal organs present between prostomium and elytrophores of segment 2. Facial tubercle present or absent. m

Segment 3 partially or entirely reduced dorsally, with single pair of laterally directed dorsal cirri. E

Branchiae cirriform, from segment 4-7, sometimes absent from segments with dorsal tubercles.

Parapodia biramous from segment 2, with each ramus bearing a single acicula. Notopodia lacking bracts, /t

with a single presetal stylode. Neuropodia lacking bracts, with a single presetal tubercle and up to 2 postsetal II

lobes; superior postsetal lobe cirriform, inferior postsetal lobe rudimentary; with long, tapering ventral cirri.

Three large ctenidia between notopodia and elytrophores/dorsal tubercles. Additional ctenidia may variously I1

occur ventral to neuropodia, dorsal to elytrophores and dorsal tubercles, and elsewhere on parapodia. Notosetae

all simple with bifid tips. Neurosetae include simple spinous setae, compound falcigers with blades ranging

from long and multiarticulate to short with a single article. Pygidium with 2 filiform anal cirri. II

Remarks. Following an examination of the type species of Sigalion, Thalenessa, and Eusigalion, l

Mackie and Chambers (1990) determined that the two latter genera were junior synonyms of the first. Sigalion

has usually been distinguished as lacking a median antenna, but the type species was determined to have one. Iii

A single species, Sigalion spinosus (=Thalenessa spinosum) has been found in collections examined as part II

of this study. Specimens labeled as Sigalion sp. A in the Phase I collections proved to be Sigalion spinosus

that had lost the median antenna. I

Sigalion spinosus (ttartman, 1939)

Figure 6.6 I

Eusigalion spinosum Hartman, 1939:57-59, pl. 11, figs. 134-144, pl. 12, figs. 146-147. i

Thalenessa spinosa: Hartman, 1968:177, figs. 1-5.

u

Thalenessa cf. spinosa: Wolf, 1984:25-17, fig. 25-13, 25-14a-o. n

Sigalion spinosus: Mackie and Chambers, 1990:52. •

I

Material Examined. California, off Half Moon Bay, Sta. 2-1, 37°30.23'N, 122°45.95 'W, 77 m, coll. n

23 Sep. 1991 (1); Sta. 2-2, 37°28.18'N, 122°51.15'W, 100 m, coll. 23 Sep. 1991 (l).---Santa Maria Basin, !1

off Port San Luis, Sta. 21, 49 m (1, 2); off Point Sal, Sta. R-8 (1).

!

202 I
!

I Figure 6.6. Sigalion spinosus: A, prostomium, dorsal view; B, elytron with inset showing detail of marginal papillae; C, anterior parapodium; D, middle parapodium; E, simple neuroseta; F-G, compound

multiarticulated neuroseta. (all after Hartman, 1939)

!

Description. A large species, up to 100 mm long, 5.0 mm wide for about 115 segments; present

I specimens light tan, pink, smaller, or opaque largest (Sta white. . 2-2),Elytra 33 mmtranslucent, long, 2.5 mmmorewide or less for about covering150dorsum. segments.Body Colorcompressed, in alcohol:

subrectangular in cross section, widest at about segments 8-10; posterior segments narrow. Pygidium with 2

![ i longcirri.Prostomium trapezoidal, about as wide as long, widest anteriorly; with 2 short lateral antennae on

frontal margin and a short median antenna in posterior half of pmstomium (Fig. 6.6A), all 3 antennae subequal,

I dig present. itiform;Palps 4 small, long, cslender, ircular tape eye ripresent, ng apically, arranged extending in a square posterianterior orly to about to median segment antenna 7. Tentacular . Facial segment tubercle

with ventral tentacular cirrus slightly longer than dorsal one; with 2 bundles of slender capillary notosetae.

I Probo papillascison wventral ith 13 dorsal side; with and two 13 ventral copper-colored papillae oteeth n anteri orojaws r margin on dorsal inddand itionventral to a single, sids.larger, subterminal

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i 203
I ii

Elytra pale or white, translucent, smooth, slightly concave on anterior margin, with row of elegant, I

bipinnately branched filaments on lateral margin (Fig. 6.6B); without tubercles.

Parapodia elongate throughout, with notopodia initially shorter than neuropodia (Fig. 6.6C), then '[]

notopodia gradually increasing in length, becoming longer than neuropodia by middle of body (Fig. 6.6D). g

Notopodia slightly inflated, terminating in short cirrus. Neuropodia obliquely truncate, terminating in triangular

lobe beating acicula; ventral cirri cirriform, thin, about as long as neuropodium. Branchiae from segment 4, Ill

each long, thick (Fig. 6.6D). []

Notosetae long, simple capillaries with fmely serrated edge. Neurosetae arranged into supracicular

and subacicular fascicles; supracicular fascicle consisting of an anterior group of7-10 f'mecompound fatcigers I

and 6-9 spinose simple setae (Fig. 6.6E), and a posterior group of 12-15 heavier, bifid compound falcigers lib

with blades ranging from short to long. Subacicular fascicle with 3-5 heavy, stout compound falcigers with

short blades and 40 or more long, multiarticulated compound falcigers (Fig. 6.6F-G). !

gl

Remarks. Sigalion spinosus has been reported as Eusigafion spinosum or Thalenessa spinosa from

collections ranging from central California south to central America and the Galapagos Islands. The species

was recently reported from the Gulf of Mexico by Wolf (1984). The present collections include material from 1

off Half Moon Bay, extending the northern range slightly north of Monterey Bay. In some specimens the w

median antenna has been lost, confirming the deciduous nature of this structure in species of Sigalion. m

Biology. The species is found in continental shelf depths to about 100 m in sediments consisting of i

mixed sand and silt. According to Hartman (1939), there were no tubes associated with the species when it

Illll¢

was collected and she assumed it was a burrower. Ill

Distribution. Northern California to Central America; Galapagos Islands; Gulf of Mexico; 18-119 m. il

Literature Cited 1

Augener, H. 1918. Polychaeta. In Beitr_ige zur Kenntnis der Meeresfauna West-Afrikas. Herausgegeben von I

W. Michaelsen 2(2):67-625, pls. 1-6, 111 text figs.

w

Baird, W. 1866. Contributions to a monograph of the Aphroditacea. Journal of the Linnaean Society (Zoology) []_

9(33):31-38. I

Banse, K. and K.D. Hobson. 1968. Benthic polychaetous annelids from Puget Sound, Washington, with

remarks on four other species. Proceedings of the United States National Museum 125:1-53. i

Banse, K. and K.D. Hobson. 1974. Benthic errantiate polychaetes of British Columbia and Washington.

Bulletin of the Fisheries Research Board of Canada 185:1-111. I

[] Berkeley, E. 1923. Polychaetous annelids from the Nanaimo district. Part 1. Syllidae to Sigalionidae.

Contributions to Canadian Biology, New Series 1(i 1):205-218. I1

Berkeley, E. and C. Berkeley. 1935. Some notes on the polychaetous annelids of the Elkhorn Slough, Monterey I

Bay, California. American Midland Naturalist 16:766-775.

Berkeley, E. and C. Berkeley. 1941. On a collection of Polychaeta from Southern California. Bulletin of the I

Southern California Academy of Sciences 40:16-40. I

Berkeley, E. and C. Berkeley. 1942. North Pacific polychaeta, chiefly from the west coast of Vancouver 1

Island, Alaska, and Bering Sea. Canadian Journal of Research Section D Zoological Sciences !1

20(7):183-208.

!
!

I Blake, J.A. 1975. Phylum Annelida: Class Polychaeta. Pp. 151-243, In, Smith, R.I. and J.T.Carlton (eds.),

Light's Manual: Intertidal Invertebrates of the Central California Coast. University of California

i Press, Berkeley. Blake, J.A. 1995. Chapter 5. Family Pholoidae Kinberg. In: Blake, J.A., B. Hilbig and P.H. Scott (eds.),

Taxonomic Atlas of the Santa Mafia Basin and Western Santa Barbara Channel. Volume 5:175-188.

i Santa Barbara Museum of Natural History, California.

Cazaux, C. 1968. l_tude morphologique du dgveloppement larvaire d'annglides polych&es (Bassin d'Arcachon).

t Cuvier, G.L.C.ED. I. Aphroditidae, Chrysopetalidae. Archives de Zoologie Exprrimentale et G_nrrale 109:477-543. 1830. Le R_gne Animal distribu6 d'apr_s son organisation, pour servir de base h l'histoire

naturelle des animaux et d'introduction _t1'anatomie comparre. Nouvelle &tition, revue et augmentre.

I Tome 3. Paris.

Day, J.A. 1967. A monograph on the Polychaeta of Southern Africa. Part 1. Ermntia. Publication of the

i British Museum of Natural History No. 656:1-458, 108 figs. Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families and genera. Natural

History Museum of Los Angeles County, Science Series 28:1-190.

I Fauchald, K. and EA. Jumars. 1979. The diet of worms: A study of polychaete feeding guilds. Oceanography

and Marine Biology Annual Review 17:193-284.

I Hartman,Expeditions O. 1939. Polychaetous annelids. Part 1. Aphroditidae to Pisionidae. Allan Hancock Pacific 7:1-156, 28 plates.

i Hartman, O. 1960. The benthic fauna of the deep basins off southern California. Allan Hancock Pacific Expeditions 22(2):69-216, pls. 1-19.

Hartman, O. 1961. Polychaetous annelids from California. Allan Hancock Pacific Expeditions 25: 1-226.

I Hartman, O. 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation,

University of Southern California, Los Angeles, California. 828 pp.

I Hartman,No. O. 6:1-64. and D.JOregon . Reish.State 1950.College The marine Press,annelids Corvallis,of Oregon. Oregon. Oregon State Monographs in Zoology

i Johnson, H.P. 1897. A preliminary account of the marine annelids of the Pacific coast, with descriptions of new species. Proceedings of the California Academy of Sciences, Zoology 1(5): 153-198.

Johnson, H.P. 1901. The Polychaeta of the Puget Sound region. Proceedings of the Boston Society of Natural

I History 29:381-437.

Kinberg, J.G.H. 1856. Nya sl_igten och arter af Annelider. 0fversigt af Konglia Vetenskaps-Akademiens

i Mackie, Frrhan A.S.Y.dand lingarS.J. 12Chambers. :381-388.1990. Revision of the type species ofSigaIion, Thalenessa, andEusigalion

(Polychaeta: Sigalionidae). Zoologica Scripta 19:39-56.

I McGinitie, G.E. 1935. Ecological aspects of a California marine estuary. American Midland Naturalist

16:629-765.

I Monro, Cand.C.the A. 1933 Galapagos . The Islands Polychaduring eta Erranthetiaexpedition collectedofbytheDr.S.Y. C. Cr 'St.osGeorge.' sland atProceedings Col6n, in theofPanama the Zoological region,

Society of London, Part 1:1-96.

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Moore, J.E 1910. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern i

California in 1904. Polynoidae, Aphroditidae and Segalionidae. Proceedings of the Academy of

Natural Sciences of Philadelphia 62: 328-402. i

U

Pettibone, M.H. 1953. Some Scale-beating Polychaetes of Puget Sound and Adjacent Waters. University of

Washington Press, Seattle, Washington. 89 pp. m

Pettibone, M.H. 1969. The genera Sthenelanella Moore and Euteanira Horst (Polychaeta, Sigalionidae). U

Proceedings of the Biological Society of Washington 82:429-438.

Petfibone, M.H. i970a. Revision of the genusEuthalenessa Darboux (Polychaeta: Sigalionidae). Smithsonian I

Contributions to Zoology No. 52:1-30.

Pettibone, M.H. 1970b. Revision of some species referred to Leanira Kinberg (Polychaeta: Sigalionidae). i

Smithsonian Contributions to Zoology No. 53:1-25. m

Pettibone, M.H. 1970c. Two new genera of Sigalionidae (Polychaeta). Proceedings of the Biological Society

of Washington 83:365-386. B

Pettibone, M.H. 1971a. Partial revision of the genus Sthenelais Kinberg (Polychaeta: Sigalionidae) with

diagnoses of two new genera. Smithsonian Contributions to Zoology No. 109:1-40. I

Pettibone, M.H. 197lb. Descriptions of Sthenelaisfusca Johnson, 1897 and S. berkeleyi n. sp. (Polychaeta:

Sigalionidae) from the Eastern Pacific. Journal of the Fisheries Research Board of Canada 28:1393-

1401. i

Pettibone, M.H. 1982. Classification of Polychaeta. In: Parker, S.P. (ed.). Synopsis and Classification of

Living Organisms. McGraw-Hill. pp. 3-43. ml

n

Pettibone, M.H. 1992a. Two new genera and four new combinations of Sigalionidae (Polychaeta). Proceedings

of the Biological Society of Washington 105:614-629. IIi

Pettibone, M.H. 1992b. Contribution to the polychaete family Pholoidae Kinberg. Smithsonian Contributions I

to Zoology No. 532:1-24.

Reish, D.J. 1968. A biological survey of the Balda de Los Angeles, Gulf of California, Mexico. II. Benthic n

polychaetous annelids. Transactions of the San Diego Society of Natural History 15:67-106. g

Treadwell, A.L. 1914. Polychaetous annelids of the Pacific coast in the collection of the zoological museum i

of the University of California. University of California Publications in Zoology 13:175-234. II

Wolf, P.E. 1984. Chapter 25. Family Sigalionidae Malmgren, 1867. In: Uebelacker, J.M. and P.G. Johnson

(eds.), Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Vol. 3:25-1 to 25-39. B

Barry A. Vittor & Associates, Inc., Mobile, Alabama. i

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I 7. FAMILYAMPHINOMIDLAAMARC E K, 1818

i Uy

Jerry D. Kudenov _

I Introduction

t The amphinomids, along with other members of the Order Amphino_dasensu Fauchald, 1977a, are

strongly isolated in the Class Polychaeta (Clark, 1969). Certain species such as Eurythoe complanata, the sol called "fn-eworm," can be dangerous to humans. They produce an acidic neurotoxin that is transmitted to unwary humans or erstwhile predators through detachable, and highly brittle, calcareous harpoon notosetae.

Only those amphinomids that produce setal associated neurotoxins can rightly be called fireworms; many

I amphinomids apparently do not. GeneraLly speaking, most members of this family tend to be much smaller thanAmphinome, Eurythoe orHermodice, which can approach lengths of 500 mm and widths in excess of 15

mm, excluding setae.

!

i Morphology

Members of this family are brilliantly pigmented, and sometimes have characteristic color patterns,

I wNohtopygos ich is particul speciesarlycompl true feorx b(otKudh Chloeia enov, 1991 and)Notendtopytogosbe. The grublikbodies e andofoval suchinamcross-s phinomection ids asfothreaChllimit oeiaednumber of segments, whereasAmphinome, Eurythoe, Hermodice, Pareurythoe, and similar forms are elongate

i and subrectang posterior lobes ular inandcrosstendssectiono beandtightly havesurrounded many morebysegme anterior nts. The setigers postomium and denseisreprese fasciclesntedof bynotosetae. anterior

The anterior lobe is rounded to slightly incised anteriorly, rounded dorsally, has a pair each of palpal and

i lateral of eyes,antenn and aamedian e, and acaruncle pair of eyes. that generally The posterior extendslobeposteriorly is wider than in configurations long, with a median such as:antenna, (1) a flatanother unadorned pair

plate in Paramphinome, (2) a sinuous lobe in Pareurythoe, (3) a median keel with a pair of lateral ridges in

i Eurythoe and Pherecardia , or (4).a medHipponoa ian keel lacks with complexly a caruncle folded (Kudenov margi, n1977, s as in1994). the ChloThe eia-phNotopygos arynx is uspecies narmed,complex highly

muscular, and has an extensive series of characteristic transverse ridges. A peristomial segment is apparently

i absent. Parapodia are biramous, and the notopodial and neuropodial rami are widely separated. Notopodia

have setae arrayed in radiating circular whorls, with one or two dorsal cirri. Branchiae may be present on

some notopodia as in Paramphinome or Linopherus, or on most as in Branchamphinome and Pareurythoe,

i and are usually absent from the first few setigers. They are generally dendritic in most genera, and can be bipinnate in Chloeia and related taxa, or palmate in Branchamphinome (Kudenov, 1993). Neuropodia also

have setae arrayed in radiating circular whorls and bear one ventral cirrus. All setae are simple, calcareous

and brittle. Notosetae include bifurcate and harpoon setae, unadorned or spurred spines, and spurred and

I nonspurred capillaries. Neurosetae may include recurved as Hipponoa and Amphinome, biflLrcate

hooks in

setae, unadorned or spurred spines, and spurred or nonspurred capillaries. Aciculae are generally blunt

i distally, club-shaped and located in a narrow arc just anterior to the dorsal cirri in the notopodia and in a

i t Department of Biological Sciences, University of Alaska Anchorage, 3211 Providence Drive, Anchorage, Alaska 99508

i 207
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narrow superior arc in the neuropodia. The pygidium is either dorsal with unpaired midventral anal papilla in I

such genera as Amphinome and Eurythoe, or terminal with a pair of anal cirri, as in the Chloeia-Notopygos l

species complex (Kudenov, 1991).

Amphinomids and other members of the Order Amphinomida are unusual, compared to other l

polychaetes, in having such features as calcareous setae, two pairs of longitudinal nerve cords including an

!ll

additional lateral tract, and a series of ventral epidermal pigment canals that parallel the nervous system lli

(Gustafson, 1930; Kudenov, 1974). All other known modem day polychaetes reported to date lack these 1

features, although a few interstitial forms may have two pairs of nerve cords. Ill

Specific characters having taxonomic importance include the morphology of the prostomium, in

development of prostomial appendages, eyes, caruncle, setiger 1, parapodia of middle body segments, the il

kinds of noto- and neurosetae, development and numbers of noto- and neuroaciculae, distribution and placement

of branchiae and parapodial cirri, and development and kinds of anal cirri, n

Taxonomic History 1

The first four amphinomids ever to be described in western literature were assigned to the genus

Aphrodita by Pallas (1766). The family Amphinomidae was later formally recognized by Lamarck (1818), l_

as noted by Pettibone (1982), and not Savigny (1818) as reported by Hartman (1959:128; for comments see il

Kudenov, 1992). The primary systematic study for amphinomids is that of Gustafson (1930:305-471) who

focused on comparative anatomy. Other important references for this group have been summarized by Kudenov i

(1974, 1992). n

This family is not well understood systematically. The Order Amphinomida to which it is assigned is

clearly monophyletic (Fauchald, 1974, 1977a) and not closely related to any other extant Order in the Class 1

(Clark, 1969). Only recently has the Order Amphinomida been restricted to include the families Amphinomidae

and Euphrosinidae by Fauchald, 1977a (see also Pettibone, 1982); the spintherids were removed and placed

into a separate Order. Prior to 1977, the Order Amphinomida embraced amphinomids and euphrosinids either _l

as the family Amphinomidae containing two subfamilies, or as two separate families which applies here. U

Historically, spintherids were included because of their superficial resemblance to the other taxa (Manton,

1967). Kudenov (1991) has since established the Archinomidae, a new family in this Order from the Galapagos I_

hydrothermal vents, based upon a single, unusual species,Archinome rosea (Blake). Kudenov (1991) provides II

a brief historical overview of the Amphinomida.

In all, 19 genera and around 130 species of amphinomids generally considered valid have been described !

(Hartman, 1959, 1965; Fauchald, 1977a). This will undoubtedly change as each genus is revised based on n

type specimens. For example, the only generic revisions performed to date on amphinomids are those of

Bindra (1927) who worked on the genus Eurythoe, and Fauchald (1977b) who proposed Linopherus as the II

senior synonym of Pseudeurythoe. Both were based on the literature. Eurythoe is complex in that its extreme

morphological variation may be the result of its reproductive behavior (at least in Eurythoe complanata, see

Kudenov, 1974); it is also fraught with numerous synonymies, some of which seem to be highly dubious I_

(Kudenov, unpublished data). Pettibone (in litt.) did not agree with Fauchald, and severely questioned his II

proposal to treat Pseudeurythoe as the junior synonym of Linopherus.

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208 I
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l Biology

I Amphinomids are most commonly distributed in shallow tropical and subtropical waters; a number have also been reported from abyssal depths and polar habitats. Most amphinomids tend to be small and

cryptic, especially in intertidal and shallow subtidal habitats where they are associated on rocks, in or under

I dead coral or other hard surfaces inhabited by other organisms. Other taxa such as Amphinome live on driftwood, and Hipponoa is an inquilinistic commensal inhabiting the mantle cavity of Lepas (Kudenov,

1977). Other amphinomids, some of which may be strongly reduced, live in sediments (Kudenov and Blake,

I 1985). Most amphinomids are stow moving carnivores that feed on soft sessile taxa such as sponges, anemones,

hydroids and ascidians, although Pherecardia preys upon injured crown-of-thorn starfish in the Caribbean

I (Glynn, 1984). When disturbed, most assume a defensive by arching their bodies dorsally to display posture

expansive fascicles of notosetae. Some taxa such as Chloeia and Notopygos are relatively active predators

while others such asEurythoe and Pareurythoe are omnivorous with decided preferences for animal remains.

I Some larger and rather conspicuous taxa such as Hermodice can be observed feeding on coral polyps during daylight hours, in contrast to Eurythoe complanata which exhibits an aversion to light and may be nocturnal

or crepuscular in its feeding habits. The feeding habits of this family are poorly known, and summarized by

I Fauchald and Jumars (1979).

Relatively little is known about the reproductive biology of amphinomids. The literature was

summarized by Kudenov (1974) and Schroeder and Hermans (1975). Eurythoe complanata exhibits an

I annual of architomic in the followed release of later in the cycle scissipafity spring by a gametes summer.

Some species are known to swim using natatory setae (Fauvel, 1951; Hartman, 1951) and are attracted to

night lights (Fauvel, 1951). The rostraria larva is attributed to amphinomids, although the complete larval

I has not been described for this to date.

cycle group

I Description of Species

i Genus Chloeia Lamarck, 1818

Type Species. Aphroditaflava Pallas, 1766

I Diagnosis. Body ovate, oval in cross section, with around 30 segments. Prostomium with anterior

and posterior lobes. Anterior lobe with pair of palpal antennae; posterior lobe with 2 pairs of eyes, pair of

I lateral pleatedantennae, lateral folds. andBranchiae an unpairedfirstmedian presentantenna. from setigers Caruncle 4-5, bipirmate. well developed, Dorsalwith cirri high numbering medianonekeelor two and

per anterior segment. Notosetae including thick bifurcate setae, heavy spines with minute spurs and slender

I bifurcate capillaries. Neurosetae including bifurcate setae and heavy spines with minute spurs.

Chloeia pinnata Moore, 1911

I Figures 7.1, 7.2

I Chloeia 1968:193-194, pinnata Moore,figs. 1911-4; 1:239-2 --Banse 43, pl.and15,Hobson, figs. 1-6; 1974:37 --Hartman, (key only). 1940:206-207, pl. 31, figs. 10-13;

i
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,_ pa -_--- dC

_ n

Figure 7.1. Chloeia pinnata: A, anterior end, dorsal view; B, same, anterodorsal view; C, same, anteroventral I

view; D, entire worm, ventral view; E, setiger I0, anterodorsal view; F, left setiger 12, posterior

view; G, posterior end, dorsal view.Setae omitted for clarity. All scales = 1 mm.Scale 1, D; 2, E; 3, i

F; 4, A-C; 5, G.

!

Material Examined. California: Santa Maria Basin, off Point Estero, Sta. 3, 291 m, (90, USNM);

offPort San Luis, Sta. R-2, 161 m, (7, USNM); off Point Arguello, Sta. 53, 196 m, (100, SBMNH). I

Description, Present materials measuring up to 13 mm long, 5 mm wide excluding setae, for 24 I

segments. Species reported up to 30 mm long, 7 mm wide without setae, for 17-28 segments. Body generally

pate pink-orange in alcohol, without pigmented bands or pattern; with purple pigment on prostomium anterior Ill

to paired lateral antennae, ceratophores of all dorsal cirri from setiger 4, dorsal cirri from setigers 4-6, and on I

medial and paired lateral ridges of caruncle.

Prostomium with anterior lobe rounded anteriorly, with paired palpal antennae long, slender, cirriform; •

posterior lobe smaller, with paired lateral antennae, cirriform, longer than palpal antennae, arising from II

anterior margin, two pairs of pronounced purple eyes of which anterior pair largest, and median antenna

arising from anterior margin of caruncle, 2x longer than lateral antennae, 0.6x length of caruncle (Fig. 7.1 A- "l

C). Caruncle as described for the genus, extending posteriorly to middle of setiger 4 (Fig. 7.1A, B); fused to

dorsum of setigers 1-2, free thereafter. Palps fused, converging midvenlxally into longitudinal groove leading

to mouth (Fig. 7.1C). Mouth located between palps and posterior lip formed by setiger 3 (Fig. 7.1C, D). Im

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I Parapodia well developed with widely separated rami in all segments (Fig. 7.1E, F). Notopodia set

off from body wall by incised groove, elliptical in cross section, conical, terminating in long cirrophores for

i dorsal cirri; neuropodia set off from body walt by low narrow rim, circular in cross section, pad-shaped (Fig. 7.IC). Setiger 1strongly reduced, both noto- and neuropodia directed dorsally (Fig. 7.1C); setiger 2 reduced,

notopodia directed dorsally, neuropodia laterally; setiger 3 completely developed.

I or cappedAllbysetae a clearhollow. matrix Notosetae (Fig. 7.2A',yellow, L-O), ofthicker, three kinds less numerous including:than(1) long neurosetae, bifurcatetipssetae, distally mostensheathed numerous,

having long prongs straight and smooth in all fascicles of anterior half of body, ratio of long to short prongs

I ranging body, withfrombarb7.1-28.6:1 (Fig. 7.2A, B, F-K); (2) bifurcate harpoon setae in all fascicles of posterior half of s offset from small prong, not 180° apart (angle of line running from harpoon barbs to center

of seta and base of small prong approximately 150° when viewed in cross-section), ratio of long to short

I prongs smooth,ranging numbering from around 8.1-8.8:115(Fig. per fascicle, 7.2C-E); arrayed and (3) spinous as outermost notosetae, whorlshort, alongthinner notopodial than bifurcate border (Fig. notosetae, 7.2LM). Notoaciculae spinose, entire, numbering 3-4 per notopodial fascicle, arrayed directly in front of notopodial

I cirrus (Fig. 7.20). All notosetae with regular pattern of minute, file-like teeth set in alternating spiral rows, best developed on all surfaces proximal to small prong, generally absent distal to small prong (Fig. 7.2B').

Neurosetae paler yellow, considerably thinner, and much more numerous than notosetae, all bifurcate,

i including long spurredan outer capillaries ring of(Fig. relatively 7.2T-W, heavier Z, AA), neurosetae of which(Fig. superior 7.2P-S,onesX, Y, longest, AB) surrounding thinnest (Fig.dense 7.2Z). tuftsAll of

neurosetal fascicles especially dense in posterior half of body. Neuroaciculae spinose, not as heavy as

I notoaciculae, All neurosetaenumbering with regular10-12 pattern per fascicle, of rasp-like arrayed teethalset ongindoalternating rsal superiorspiralegiorows, n of fasc besticledeveloped (Fig. 7.2AC). on all

surfaces proximal to small prong, generally reduced distal to small prong (Fig. 7.2X-AB).

I and ventral All setigers cirri with parapodial cirri. Setigers 1-3 with relatively thick cirri, including true dorsal, lateral (Fig. 7.1A). Dorsal cirri arising medial to lateral cirri, lacking cirrophores. Lateral cirri

(called dorsal cirri on first branchial parapodia) associated with notopodia, long, length of cirrophores about

I equal cirrophore to length conspicuous, of segment;about cirrostyle 0.5x length slender,ofcirriform, dorsal cirrophores about 3.5xinlonger midbodythansegments; cirrophores.cirrostyle Ventral gradually cirri with

increasing in length to about 0.5x body width of midbody segments, longer than dorsal cirri (Fig. 7.1D).

Branchiae bipinnate, present from setiger 4 to end of body, with 8 alternating branches arising from primary axis, each terminating in smaller branches off'me, digitiform pinnae or terminal filaments on posterior

surfaces only; anterior surface lacking pinnae (Fig. 7.1E, F). Branchiae best developed in anterior and middle

i segments, decreasing in size and development thereafter. Pygidium terminal, opening between pair of thick, cylindically appressed anal cirri (Fig. 7.1G).

i Biology. Chloeia pinnata is one of the most abundant amphinomids known off the coast of southern and central California where it inhabits green and gray muds, and poorly sorted fine to coarse sands with shell

fragments, cobbles and rocks (Jones & Thompson, 1988).

i additionalRemarks. details andChloe illustrations iapinnatawere wasprovided adequatelybydescribed Hartman but (1940, poorly1968) illustrated and otherbysubsequent Moore (1911). workers. Few

The most noteworthy finding in this study is the presence of unusual bifurcate notosetae from posterior

i segments the small with prong;a rowtheyofareharpoon not separated barbs onbytheanlongangleprong of 180 separated °. This byis the an angle very feature of approthat ximately Hartman150(1940, ° from

1968), Banse and Hobson (1974) and others have used to distinguish Chloeia entypa Chambertin, 1919,

m which has this seemingly unusual kind of notoseta, from C.pinnata. Given this and other traits, it appears that

I Chloeia pinnata may be the senior synonym of C. entypa. However, there are other apparent differences

between these two species, and their treatment will be discussed elsewhere once the type of C. entypa can be

i examined.

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Figure 7.2. Chloeiapinnata: A, notoseta, from superior of fascicle, with A' showing detail of tip; B, same, with

B' showing detail of surface; C-E, bifurcate harpoon notosetae; F-K, spurred notoseta from inner m

setal field: F, center of fascicle; G, superior of fascicle; H, anterior of fascicle; I-J, inferior of fascicle; II

K, superior of fascicle; L-N, spinose notosetae, from outermost whorl: L, superior of fascicle; M,

. anterior of fascicle; N, inferior of fascicle; O, notoacicula; P-S, spurred neurosetae from outer field i

of fascicle: P,superior neuroseta; Q, anterior neuroseta; R, inferior neuroseta; S, posterior neuroseta; 1

T-W, spurred neurosetal capillaries from inner setal field: T,superior neuroseta; U, anterior neuroseta;

V-W, inferior neurosetae; X-AB, surface details of neurosetae: X, Y, AA, spurred neurosetae from mira

outer field of fascicle; X, superior neuroseta; Y, posterior neuroseta; AA, anterior neuroseta; Z, AB, •

spurred neurosetal capillaries from inner field of fascicle: Z, longest capillary from center of fascicle; i

AB, shorter capillary from inferior of fascicle; AC, neuroacicula. Scales: 1 = .001 ram, X-A_B;2 =

.001 ram, B'; 3 = .01 mm, A-W, AC and .005 ram, A'. I

II

Chloeiapinnata can thus be characterized as a relatively small Chloeia species, lacking any apparent i

color patterns on the dorsum when in alcohol, although purple pigment is present on the prostomium anterior II

to the paired lateral antennae, notopodial cirri from setiger 4, and anal cirri. Black pigment specks noted by

Harttnan (1940) on the dorsum were not noted in any of the nearly 200 specimens examined in this study. The i

caruncle extends posteriorly to setiger 4. Two dorsal cirri (actually the true dorsal and lateral cirri; see I

Gustafson, 1930; Kudenov, 1987, 1991) are present on setigers 1-3. Branchiae are bipinnate and occur from

setiger 4 to the end of the body. Notosetae of three kinds including: (1) bifurcate setae with long smooth i

212 i
!

I prongs in the anterior half of the body, (2) bifurcate harpoon notosetae in the posterior half, and (3) spinous

notosetae that are smooth and short, present in an outer whorl around the periphery of notopodial fascicles.

I Neurosetae neuropodial are all bifurcate and include relatively short slender spines arrayed around the periphery of

fascicles, and an inner tuft of very long capillaries which are best developed in the posterior onehalf of the body. All setae are hollow, with characteristic file- or rasp-like teeth arranged in alternating spiral

i rows on nearly all surfaces, and are either distally ensheathed or are tipped with hyaline-like caps. The second trait was referred to by Moore as being granular.

Note that Moore (1911) assumed the peristomium to be entirely fused with the prostomium, and

I coassunumpts segmen tion (Gutsstinafboson, th the 193amphinomi 0). ds and euphrosinids accordingly. There is no evidence supporting his

Distribution. Southern California, in shelf and slope habitats to depths of 567 m.

!

Literature Cited

!

Banse, K. and K.D. Hobson. 1974. Benthic errantiate polychaetes of British Columbia and Washington.

Bulletin of the Fisheries Research Board of Canada 185:1-111.

i Bindra, S.S. 1927. Fauna of Karachi. 1. A study of the genusEurythoe (family Amphinomidae). Memoires of

the Department of Zoology, Punjab University 1:1-18.

I Chamberlin, R.V. 1919. The Annelida Polychaeta. Memoirs of the Museum of Comparative Zoology, Harvard 48:1-514.

I Clark, R.B. 1969. Systematics and Phylogeny: Annelida, Echiura, Sipunculida. In Florkin, M. and B. Scheer (eds.), Chemical Zoology Vol. 4. Academic Press, New York. Pp. 1-68.

Fauchald, K. 1974. Polychaete phylogeny: a problem in protostome evolution. Systematic Zoology 23:493-

I 506.

Fauchald, K. 1977a. The Polychaete Worms Definitions and Keys to the Orders, Families and Genera. Natural

I History Museum of Los Angeles County, Science Series 28:1-190.

Fauchald, K. 1977b. Polychaetes from intertidal areas in Panama, with a review of previous shallow-water

records. Smithsonian Contributions to Zoology 221:1-81.

I Fauchatd, K. and P.Jumars. 1979. The diet of worms: A study of polychaete feeding guilds. Oceanography

and Marine Biology Annual Review 17:193-284.

I Fauvel, P. 1951. Annrlides polych_tes du Golfe de Tadjoura recueillies par M.J.L. Dantan en 1933, au cours

de pSches nocturnes _ la lumi_re. Bulletin de la Musre Nationale d'Histoire Naturelle Paris, Ser. 2,

23:287-294, 381-389, 519-526, 630-640, 4 figs.

I Glynn, Peter. W. 1984. An amphinomid worm predator of the crown-of-thorns sea star and general predation

on asteroids in eastern and western Pacific coral reefs. Bulletin of Marine Science 35:54-71.

I Gustafson, G. 1930. Anatomische Studien iiber die Familien und Polychaeten Amphinomidae Euphrosynidae.

Zoologiska Bidrag frhn Uppsala 12:305-471.

I Hartman,Expeditions O. 1940. Po7(3): lychae 173-287. tous annelids. Part II. Chrysopetalidae to Goniadidae. Allan Hancock Pacific

Hartman, O. 1951. The littoral marine annelids of the Gulf of Mexico. Publications of Texas

i University Institute of Marine Science 2:7-124.

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Hartman, O. 1959. Catalogue of the polychaetous annelids of the world. Part I. Allan Hancock Foundation I

Occasional Paper 23:1-354.

Hartman, O. 1965. Catalogue of the polychaetous annelids of the world, including additions and emendations m

since 1959. Allan Hancock Foundation Occasional Paper 25:1-197.

Hartman, O. 1968. Atlas of the Errantiate Polychaetous Armelids from California. Allan Hancock Foundation, ml

University of Southern California, Los Angeles, 828 pp. l

Jones, G.F. and B.E. Thompson. 1988. The distribution and abundance of Chloeia pinnata Moore, 1911 IN

(Polychaeta: Amphinomidae) on the southern California borderland. Pacific Science 41:122-131. m

Kudenov, J.D. 1974. The reproductive biology of Eurythoe complanata (Pallas, 1766) (Polychaeta:

Amphinomidae). University of Arizona, Doctoral Dissertation. Pp. 1-128. m

Kudenov, J.D. 1977. Brooding behavior and protandry in Hipponoe gaudichaudi (Polychaeta: Amphinomidae).

Bulletin of the Southern California Academy of Sciences 76: 85-90.

m

Kudenov, J.D. 1987. Review of the primary species characters for the genus Euphrosine (Polychaeta: m

Euphrosinidae). Bulletin of the Biological Society of Washington 7:184-193.

Kudenov, J.D. 1991. A new family and genus of the Order Amphinomida (Polychaeta) from the Galapagos m

hydrothermal vents. In Petersen, M.E. and J. Kirkegaard (eds.), Proceedings of the Second International

Polychaete Conference: Systematics, Biology and Morphology of World Polychaeta. Ophelia, m

Supplement 5:111-120. m

Kudenov, J.D. 1993. Amphinomidae and Euphrosinidae (Annelida: Polychaeta) principally from Antarctica,

the southern ocean and subantarctic regions. Biology of Antarctic Seas XXII, Antarctic Research m

Series 58:93-150. m

Kudenov, J.D. 1994. Redescription ofHipponoa gaudichaudi Audouin & Milne-Edwards, 1830 (Polychaeta:

Amphinomidae). In Dauvin, J.C., L. Laubier and D.J. Reish (eds.), Actes de la 4_me Conferrnce m

internationale des Polych_tes. Mrmoires du Musrum National d'Histoire Naturelle (A) 155: g

Kudenov, J.D. and J.A. Blake. 1985. A new species of ofPseudeurythoe (Polychaeta: Amphinomidae) lm

from central California. Bulletin of the Southern California Academy of Sciences 84(1): 38-40. l

Lamarck, J.B. 1818. Histoire naturelle des Animaux sans Vertrbrrs prrsentant les charact_res grnrraux et

particuliers de ces animaux, leur distribution, leurs classes, leurs families, leurs genres, et la citation m

des principales esp_ces qui s'y rapportent; prrcrdre d'une Introduction offrant la drtermination des U

charact_res essentiels de l'Animal, sa distinction du vrgrtal et des autres corps naturels, enfm,

l'expostion des principes fondamentaux de la zoologie. Paris Vol. 5, Pp. 1-612 (Annrlides, Pp. 274- i_

374). m

Manton, S. 1967. The polychaete Spinther and the origin of the Arthropoda. Journal of Natural History, 111

London 1:1-22. m

Moore, J.P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern

California in 1904. Euphrosynidae to Goniadidae. Proceedings of the Academy of Natural Sciences m

of Philadelphia 63:234-318.

Pallas, P.S. 1766. Miscellanea Zoologica, quibus novae imprimis atque obscurae Animalium species m

describunture et observationibus iconibusque illustrantur. Hagae Comitum, 244 pp. |

Pettibone, M.H. 1982. Classification of Polychaeta. In Parker, S.P. (ed.), Synopsis and Classification of

Living Organisms. McGraw-Hill. Pp. 3-43. I

214 m
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I Savigny, J.S. 1818. Ann61ides. In Lamarck, J.B. De. 1818. Historie naturelle des animaux sans vert6br6s.

Paris 5(3):274-374.

I Schroeder, P.C. and C.O. Hermans. 1975. Annelida: Polychaeta. In Giese, A.C. and J.S. Pearse (eds.) Reproduction of Marine Invertebrates. Vol. 11I.Academic Press, New York. pp. 1-213.

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I 8. FAMILY EUPHROSINIDAE WILLIAMS, 1851

i uy

Jerry D. Kudenov _

I

l Introduction

Eup_osir_ds, like their amphinomid relatives, represent another farmly in the OrderAmphinomida

l Fauchald (1977), which is isolated in the Class differ

sensu strongly Polychaeta (Clark, 1969). Euphrosinids

from amphinomids in lacking harpoon notosetae, or any kind of notoseta that readily detaches. They apparently

do not produce associated setal neurotoxins, and are therefore not called fireworms. Generally speaking, most

I members of this family tend to be less than 40 mm long, 10 mm wide, and have fewer than 50 segments.

I Morphology

I are apparently Members absent. of theBodies Euphrosinidae are somewhat tend tosemi-circular be pigmentedin across-section brilliant orange, and but elliptical characteristic in outline;color the pdorsum atterns

is arched and the ventrum is flat.

I ventrum,Thetightly prostomi wedged um between is an elongate, anteriornarrow setigersridge and highly extendingobscured over the by dense anteriorfascicles end froofmnotosetae. the dorsumTheto

dorsal part of the prostomium bears a pair of large eyes located lateral to the median antenna. The caruncle

I keel is fusedandtthree o the dors pairsumof inlateral Euphrosine ciliated, ridges and generally are present free oinn both a variable Euphros number inella ofandanterior Euphrossegments. inopsis; aAmedian pair of

small palpal antennae arise lateral to the smaller pair of eyes; all are placed anteriorly to a pair of elliptical,

medial pharynxpalps is unarmed, which, inhighly turn, are muscular, anteriorandto thehasmouth. an extensive The mouseries th isofsucharacteristic rrounded by thetransverse first fourridges. setigers. The

Parapodia are biramous, with rami projecting only slightly above the body surface; notopodial and

I neuropodial notosetae arrayed rami are in transverse separated. rows. Notopodi Braanchiae are present occur aeither s transverse singly orridges withwith up tosev13eralin tiers a transverse of bifurcrowate

posterior to the dorsal and lateral cirri. Neuropodia are circular in outline, with setae arrayed in radiating

I circular whorls. A ventral cirrus is present. Branchiae axegenerally present from setiger 1, either as a single cimform filament, as several filaments

arising directly from the body wall, or as dichotomously branching, arborescent processes; or they may be

I absent (Kudenov, 1993).

All setae are simple, calcareous and brittle. Notosetae include bifurcate and ringent setae. Neumsetae

include bifurcate setae, and delicate spurred capillaries in some sexually mature specimens. Acicu|ae are

I generally bifurcate, and apparently located in a transverse tier just anterior to dorsal and lateral cirri in the

notopodia, and in a narrow superior arc in the neuropodia. All bifurcate noto- and neurosetae vary consistently

both within fascicles and along the body in two species ofEuphrosine (Vogt and Kudenov, 1994).

I

_Department of Biological Sciences, University of Alaska Anchorage, 3211 Providence Drive, Anchorage, Alaska 99508

I

i 217
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The pygidium is terminal with a pair of short, cushion- to cup-shaped anal cirri surrounded by posterior

setigers. I

Euphrosinids and other members of the Order Amphinomida are unusual compared to other polychaetes

in having such features as calcareous setae, two pairs of longitudinal nerve cords including an additional

lateral tract, and a series of ventral epidermal pigment canals that parallel the nervous system (Gustafson, •

1930; Kudenov, I974). All other known modem day polychaetes reported to date lack these features, although

a few interstitial forms may also have two pairs of nerve cords. m

Specific characteristics having taxonomic importance include the number of body segments; U

morphology of the prostomium; development of prostomial appendages; eyes; caruncle; palps; setiger 1;

parapodia of middle body segments; the distribution, development and placement of branchiae, especially in 81

relation to lateral parapodiai cirri; the number of setal tiers in the notopodia; the notosetae from superior and |

inferior regions of each tier; the neurosetae from various aspects of the neuropodia; development and numbers

of noto- and neuroaciculae; and the development and kinds of anal cirri, m

Taxonomic History N

The first two euphrosinids to be described in western literature were assigned to the genusEuphrosine

by Savigny from the Red Sea (Egypt) in a manuscript that was published by Lamarck (1818). A type for the m

genus was not designated until Hartman (1959) cited E. myrtosa Savigny, 1818. The correct authority is

Lamarck, 1818, as noted by Pettibone (1963) and Kudenov (1992). The primary systematic study for

euphrosinids is that of Gustafson (1930:305-471) who focused on comparative anatomy. Other important n

references for this group have been summarized by Kudenov (1987, 1992).

As is true for the amphinomids (Chapter 7, this volume), euphrosinids are not well understood

systematically. The Order Amphinomida to which the euphrosinids are assigned is clearly monophyletic !

(Fauchald, 1974, 1977) and not closely related to any other extant Order of the Polychaeta (Clark, 1969). g

Only recently has the Order Amphinomida been restricted to include the families Amphinomidae and

Euphrosinidae by Fauchald, 1977 (see also Pettibone, 1982); the spintherids were removed and placed into a I

separate Order. Prior to 1977, the Order Amphinomida embraced amphinomids and euphrosinids either as u

the family Amphinornidae containing two subfamilies, or as two separate families which applies here.

Historically, spintherids were included because of their superficial resemblance to the other taxa (Manton,

1967). Kudenov (1991 ) added a new family, Archinomidae, to the Amphinomida, based on a single, unusual

deep-sea species, Archinome rosea (Blake, 1985), from the Galapagos Rift hydrothermal vents. Kudenov

(1991) provides a brief historical overview of the Amphinomida. m

IN In all, three genera (Euphrosine, Euphrosinella and Euphrosinopsis) and over 55 species of

euphrosinids generally considered to be valid have been described (Hartman, 1959, 1965; Fauchald, 1977;

Palmyreuphrosine Fauvel is here excluded from the above account of morphology and taxonomic history. Its I

affinities are quite uncertain and will be reported on elsewhere). These numbers will change as each genus is I

revised, based on type specimens and new collections. For example, Kudenov (1992) includes descriptions of

one new genus and eight new species of euphrosinids from Antarctica. •

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218 N
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Biology

I Euphrosinids seem to be most commonly distributed in cold shelf to abyssal habitats and polar seas;

a number have also been reported from shallow-water tropical and subtropical habitats. Most euphrosinids

tend to be small and cryptic, especially in intertidal and shallow subtidal habitats where they are associated

l with sponges, corals, barnacles, kelp holdfasts or found in rock crevices. Otherwise they tend to be associated with sandy, shelly or silty-muddy sediments in considerable depths. Their unusual body shape and that of the

anal cirri, strongly suggests that some species may be epibiotic or ectoparasitic, although this needs to be

I confirmed.

Most euphrosinids are considered to move somewhat faster than amphinomids and all are thought to

be carnivorous (Fauchald and Jumars, 1979), feeding on sponges, corals, and hydrozoans. Deep-water species

I from both the Atlantic and Antarctic Oceans also feed on foraminifera. The feeding habits of euphrosinids, however, are poorly known (Fauchald andJumars, 1979).

When disturbed, most species assume adefensive posture by rolling their bodies into a coil, displaying

I expansive fascicles of noto- and neurosetae, and looking very much like a hedgehog or porcupine.

Relatively little is known about the reproductive biology of euphrosinids. Specimens containing

i gametes 1992). Theand having delicate spurred capillary neurosetae are occasionally noted in the literature (Kudenov,

rostraria larva may be attributable to euphrosinids, although neither the complete reproductive or

larval cycle has been described for any species of this family to date.

I Key to the Species of Euphrosinidae

I 1A. Prostomium with 3 antennae (Fig. 8.1B); caruncle generally attached to dorsal body wall (Figs. 8.1A,

8.3A) ........................................................................................................... Euphrosine ................ 2

I lB. Prostomium with 5 antennae; caruncle generally not attached to dorsal body wall

...................................................................................................... Euphrosinella paucibranchiata 2

I 2A. Branchiae sessile, numbering 6-7 pairs per segment where best developed in midbody segments, each

usually terminating in 2, sometimes 1 or 3, cirriform filaments (Figs. 8.3D, E) ...............................

I .......................................................................................................................... Euphrosine bicirrata

2B. Branchiae arborescent, dichotomous (Fig. 8.1D), numbering 5 pairs per segment where best developed

I in midbody segments, each terminating in 4-5 distal branches, each branch having 8-10 subdistally

expanded sugar beet-like tips (Fig. 8.1E) ............................................................... Euphrosine arctia

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I 2AlthoughEuphrosinellapaucibranchiata(Hartman)wasnotfoundin thepresentmaterial,itwasoriginallydescribed from submarinecanyonsoffsouthernCalifornia,andis includedin thekey.It willbe redescribedelsewhere.

i

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Description of Species

I

The Euphrosinidae is a relatively small andseldom encountered family. The fauna of Santa Maria

Basin and Western Santa Barbara Channel includes two species: I

Euphrosine arctia Johnson, 1897

Euphrosine bicirrata Moore, 1905 I

Specimens originally identified asEuphrosine species A are juvenile specimens ofE. bicirrata. Measurements I

and counts of segment numbers of the present materials are cited in closed parentheses in the descriptions that

follow.

I

Euphrosine arctia Johnson, 1897 1

Figures 8.1, 8.2 I

m

Euphrosyne arctia Johnson, 1897:159, pl. 5, figs. 5-7.

Euphrosine arctia: Hartman, 1940:208-209; 1968:207-208, figs. 1-4.--Banse and Hobson, 1974:37, Fig. 7. I

Not Euphrosine arctia: Monro, 1930:34-36, fig. 4a-e; 1936:79; 1939:93-94.--Hartman, 1964:45,

pl. 14, figs. 1, 2. [=E. monroi Kudenov, 1993].

ill

Material Examined. California, Santa Maria Basin, off Purisima Point, Sta. BRA-20, 90-130.5 m, m

from rocks, (1, USNM).--Western Santa Barbara Channel, off Point Conception, Sta. BRC-2, 110-126 m,

from a rock (1, SBMNH). 1

g

Deseription. Reported less than 20 mm long (7ram); width of present specimens 2.2 mm, 3 mm

without setae; segments numbering 22 (23). Body light tan in alcohol, lacking pigmentation pattern; color in

life ochraceous, with broad red middorsal stripe about 0.25x body width. 1

Prostomium well developed, with two pairs of eyes (Fig. 8.1A-C); dorsal pair largest, each eye

present lateral andbasal to ceratophore of median antenna; ventral pairabout one-tenth size of dorsal pair, I

located between and dorsal to palpal antennae, at extreme anteroventrat prostomium. Median antenna short, !

about 0.2x length of caruncle, ceratophore tapering, about 2x longer than basal width, distal style cirriform,

about 0.5x length of ceratophore (Fig. 8.1B). Palpal antennae long, cirriform, about as long as style of m

median antenna (Fig. 8.1C). Caruncle appearing bilobed dorsoventrally, lobes attached to one another through •

middle of setiger 5, free from one another thereafter; dorsal lobe with median keel, 3 pairs of lateral ridges,

lira

each with single rows of 12-13 ciliated tufts, extending to middle of setiger 6; ventral lobe to anterior margin m

setiger 6

(Fig. 8.1B). Patps large, elliptical, paired, separated by midventral groove leading directly to mouth 1

(Fig. 8.1C). Mouth ventral, opening between setigers 2-3, latter forming posterior lip of mouth (Fig. 8.1C).

m

Parapodia biramous, with spreading notosetal and whorl-like neurosetal fascicles separated by II

interramal gap occupied by the fifth branchia; all with dorsal, lateral and ventral cirri from setiger 1to end of 1

body (Fig. 8.1 D, E). Notopodia not raised above dorsum. Neuropodia resembling distally round, circular

pads (Fig. 8.1D). .,

1

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220 I
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t pAn

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I

Figure 8.1. Euphrosine arctia: A, anterior end, dorsal view; B, prostomium, dorsal view; C, anterior end,

antero lateralvcirri, entralposte view;riorD,view; left setiger F, pygidium, 13, posterior posteoventral view; E,view. Branchiae All scales No.=1-5, 1 mm. depicting Scale 1, doA, rsalC,anD,d

F; 2, E; 3, B.

I

Notosetae all orderly, short, uniform in appearance, hollow, arrayed in a continuous outer whorl

stfrr 8.2A-D) oundingin outer innerwhorl tier, of(tiers 2 basic1, 3), kinall ds:short, bifurcaofteuniform and ringent length, notosetae. clear, glass-like Bifurcateinnoappearance, tosetae present with(Figs. long

prongs straight, stout, distally blunt, having smooth cutting surfaces; short prongs relatively short, stout, all

i distally notosetae_ntire (Fig. (Figs. 8.2E), 8.2A-D); absent fromwithtierregular 1 notosetae; spiral ratio patternof long of rectangular to short prongs file-like decreasing teeth onwithin shafta of fascicle, tier 3

ranging from 4.1 :1 superiorly to 3.9:1 inferiorly in tier 1, and increasing slightly from 3.9:1 to 4.1:1 in tier 3.

Ringent notosetae in tier 2, all category HA, of 2different lengths and shapes: long, heavy forms (Fig. 8.2F), similar in general appearance to bifurcate notosetae (Fig. 8.2A-D), numbering 4 per tier, arising from

superiormost part of tier 2; short, light forms (Fig. 8.2G, H), characteristic of category HA ringent notosetae

(Kudenov, all with 1987), present throughout remainder of tier 2, alternating with long, heavy ringent setae superiorly; regular spiral pattern of minute, scale-like teeth on shaft (Fig. 8.2I); b:a ratios (Kudenov, 1987:184,

Fig. 2) varying slightly within a fascicle, ranging from .68 superiorly to .66 inferiorly.

Neurosetae of one kind, all bifurcate, hollow, longest in superior half of neurofascicle, these longer than notosetae, becoming shorter inferiorly (Fig. 8.2J-S); long prongs smooth, characteristically narrow and

curved, appearing sword-shaped, those along posterior outer margins distally bidentate (Fig. 8.2S); with

regular spiral pattern of poorly deemed rectangular file-like teeth on some superior neurosetae (Fig. 8.2T); ratio of long to smooth prongs changing within a fascicle, ranging from 4.9:1 superiorly to 5.4:1 centrally to

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Figure 8.2. Euphrosine arctia: A, superior notoseta, tier 1; B, inferior notoseta, tier 1; C, superior notoseta, tier I

3; D, inferior notoseta, tier 3; E, middle notoseta, detail of shaft, tier 3; E long superior ringent

notoseta, tier 2; G, short superior ringent notoseta, tier 2; H, short inferior ringent notoseta, tier 2; I

I, fingent notoseta, detail of shaft, tier 2; J, long superior neuroseta; K, long neuroseta from center

of fascicle; L, long inferior neuroseta; M, short inferior neuroseta; N, subinferiormost neuroseta; O,

inferiormost neuroseta; P, short anterior neuroseta; Q, long anterior neuroseta; R, long posterior

neu suproerior seta;acSicu, shor la. Sct poalesste:ri1or=ne.001 urosemm, ta; E, T, supe I, T;ri2or= n.00eur5osram, eta, A-D, detailF-ofHsh, Jaf-S,t; U, U-Va.nterior acicula; V, I

14.4:1 inferiorly (Fig. 8.2J-O), and from 8.5-4.8:1 anteriorly to 5.4:1 centrally to 4.8-7.4:1 posteriorly (Fig. I

8.2K, P-S). Neuroaciculae bifurcate, numbering 5-6 per fascicle, present in anterosuperior row, prong ratios

ranging from 6.6-6.8:1 (Fig. 8.2U,V). I

Branchiae present from setiger 1 to end of body, numbering 5 per segment where best developed in

midbody segments, slightly less than 0.5x length of notosetae, 0.67 to 0.75x length of dorsal, lateral cirri;

dichotomously branching 2 to 2.5x, trunks and branches stout, short, terminating in characteristically thick I

distally pointed and subterminally enlarged sugar beet-shaped lobes (Fig. 8.1A,D,E) Lateral cirri arising

between gills 2-3.

Paired anal cirri terminal, inflated, with ventral cup-shaped depression in each cirrus, about as wide I

as long; anus terminal (Fig. 8.1F).

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I Biology. Relatively little is known about the biology ofEuphrosine arctia. It was originally obtained

from fish traps set at 100 and 55 m in Monterey Bay. The specimens from the Santa Maria Basin were

I collectedRemarks. from rocksType at depths of 69-135 m. specimens of Euphrosine arctia Johnson, 1897 are apparently no longer available.

The above description must therefore temporarily characterize E. arctia until adequate materials from the

I type and thelocality specimens of Monterey on whichBaythiscandescription be obtained.is based However, generally the Santa agreeMaria well Basin with Johnson's is adjacentdescrip to Monterey tion. Bay,

Johnson's original description ofEuphrosine arctia is generally accurate as far is it goes, but is quite

I incomplete. For example, his description did not include the prostomium, parapodia and setal fascicles, and the branchia he illustrated has led to confusion in the literature (Kudenov, 1992).

Perhaps the most glaring discrepancy between Johnson's description and the present specimens is

I that Johnson described and illustrated one kind of ringent notoseta forE. arctia. In fact, there two variants

are

of the category KA ringent notoseta: a few long, heavy forms reminiscent of bifurcate notosetae orE. abyssalis

Kudenov, 1992, confined to superior notofascicles that alternate with the more numerous short, light forms

I that are present throughout the tier. Johnson illustrated the latter. Given this highly characteristic feature, E.

arctia can be readily identified.

Ill This species is further characterized by having a bilobed camncle (K-1) extending to midsetiger 6, a

|

very short median antenna, and 5 pairs of arborescent branchiae per midbody segment where the gills are best

developed. Branchial trunks and branches are stout, branching dichotomously 2 to 2.5x, terminating in stout,

subdistally enlarged, sugar beet-shaped lobes. Johnson's branchial illustration is similar to those presented

I general form; portrayal of the distal branchial lobes is exactly what one would expect to see if the

above in his

gill were compressed on a slide by coverglass.

i Euphrosine Euphrosine arctia is assigned to Group II (Kudenov, 1987:187, 189), which is characterized by species having ringent notosetae (category HA) and dorsoventrally bilobed caruncles (category

K-I), and is similar to five other species in having dichotomously branching branchiae with subterminally

expanded tips. These five species include: E. armadillo Sars; E. capensis Kinberg; E. dumosa Moore; E.

I maorica Augener; and E. monroi Kudenov.

Euphrosine arctia is an example of how a poor or incomplete description can become convoluted

I (Kudenov, on branchial1993). and ringent Briefly,notosetal Monro (1930, traits. 1936, Monro1939) also identified suggested Ethat . arct/a E, armadi as a species Uoidesfromwas Antarctica the senior primarily synonym

ofE. arctia, but deferred synonymizing the two, which Hartman did in 1964. Orensanz (1972) later suspected

i that Monro's Distribution. citations actually Californiarepresented . another species, which was confirmed by Kudenov (1993).

I Euphrosine bicirrata Moore, 1905

Figures 8.3-8.4

I Euphrosyne bicirrata Moore, 1905:532-534, pl. 34, figs. 8-12; 1908:339; 1911:234.--Berkeley, 1923:211.--

Hartman, 1940:208, 209, 210-211, pl. 32, figs. 21-23.

I Euphrosine bicirrata: Hartman, 1968:211-212, figs. 1-4.--Banse and Hobson, 1974:37, Fig. 7g;

Material Examined. California, Santa Maria Basin, off Purisima Point, Sta. BRA-13 A/B, 92-100

I m, from a rock (1, USNM); Sta. BRA-14, 96-105 m, from a rock (1, SBMNH); Sta. BRC-14, 96-105 m, from a rock (1, SBMNH); Sta. BRA-20, 90-130.5 m, from a mck (1, USNM).

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I 223
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A !

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Figure 8.3. Euphrosine bicirrata: A, anterior end, dorsal view; B, prostomium, dorsal view; C, anterior end,

anteroventralview;D, fight segment 9, posterior view; E,leftsetiger 12,posterior view;F,pygidium,

posteroventral view. All scales = 1ram. Scale 1, A; 2, D-E; 3, C-F; 4, B. I

!

Description: Reported up to 20 mm long (4.5 mm), 6.5 mm wide (2 mm) without setae, for 23-25

(19) segments. Body light tan in alcohol, lacking pigmentation pattern, usually coiled, resembling bristly

balls. I

Prostomium well developed, with 2 pairs of eyes (Fig. 8.3A-C); dorsal pair largest, lateral and basal

to ceratophore of median antenna (Figs. 8.3A, B); ventral pair about half as large, between, dorsal to palpal B

antennae (Fig. 8.3C). Median antenna nearly as long as caruncle, ceratophore 2x longer than basal width, II

distal style cirriform, 2.5x longer than ceratophore (Fig. 8.3B). Palpal antennae short, stout, distally pointed

(Fig. 8.3C). Caruncle appearing bilobed dorsoventraUy, both lobes totally attached to one another from setigers •

2 to the middle of setiger 4; dorsal lobe with median keel and 3 pairs of lateral ridges, extending to anterior |

third of setiger 5 (setiger 4), ventral lobe to posterior one-third of setiger 4 (mid-setiger 3) (Fig. 8.3A,B).

Palps paired, each ovoid, separated by midventral groove leading directly to mouth (Fig. 8.3C). Mouth ventral, •

opelaing between setigers 2-3, forming posterior lip of mouth (Fig. 8.3C).

Parapodia biramous, with spreading notosetal and whorl-like neurosetal fascicles separated by narrow

interramal gap occupied by gill 7 (Fig. 8.3D); all with dorsal, lateral and ventral cirri from setiger 1 to end of

body. Notopodia not raised above dorsum (Fig. 8.3D,E). Neuropodia resembling flat circular pad (Fig. 8.3D,E).

Notosetae exceedingly long, much longer than neurosetae, radiating in various directions from fascicle, i

all hollow, arrayed in continuous outer whorl surrounding 2 inner tiers (Fig. 8.3D,E), of 2 basic kinds:

bifurcate and ringent notosetae. gO

l
!

!

!

!

,I° I

I Figure 8.4. Euphrosine bicirrata: A, superior notoseta, tier 1; B. inferior notoseta, tier 1; C, superior notoseta, tier 4; D, inferior notoseta, tier 4; E, superior notoseta, detail of shaft, tier 4; F, inferior notoseta,

detail of shaft, tier 4; G,superiormost notoseta, tier 2; H, superior notoseta, tier 2; I, inferior notoseta,

I tier neuroseta; 2; J, superi M, orsuperior ringentneuroseta; notoseta, tier N, long 3; K,neuroseta inferior rinfromgent center notosetofa, fascicle; tier 3; L,O, larges same, t sufrom periormost point

below center of fascicle; P, subinferiormost neuroseta; Q, inferiormost neuroseta; R, short anterior

i neuroseta; S, long anterior neuroseta; T, long posterior neuroseta; U, short posterior neuroseta; V,

superior neuroseta, detail of shaft; W, anterior acicula. Scales: 1 = .001 ram, E, F, V; 2= .005 ram,

A-D, G-U, W.

I Bifurcate notosetae present in outer whorl (tiers 1, 4) and anteriormost inner tier (tier 2); all opaque

white proximal to prongs, latter clear, glass-like, heavily calcified (Fig. 8.4A-D); notosetae of tiers 1 and 4

I half as long as those of tier 2, with long prongs having characteristic change in curvature at a point about equal to length of short prong from base of cleft (Fig. 8.4A-D); short prongs relatively well developed,

generally entire except bidentate in 1-3 superiormost setae of tier 1 (Fig. 8.4A); with regular spiral pattern of

1 square file-like teeth on shaft of tiers 1, 4 notosetae (Fig. 8.4E,F); ratio of long to short prongs increasing

i II slightly within a fascicle, ranging from 3:1 superiorly to 3.1:1 inferiorly in tier 1 (Fig. 8.4A,B), and decreasing

from 3.2:1 to 1.9:1 in tier 4 (Fig. 8.4C,D). Notosetae of tier 2 longest, best developed (Fig. 8.4G,H), generally

I

" I 225
!

II

longer than body width, interfacing with similar notosetae of same and neighboring segments; long prongs •

distally smooth, sometimes subtly dentate, having characteristic change in curvature similar to those of tiers

1,4, proximally striated (Fig. 8.4C); short prongs very well developed; file-like teeth not present on shafts;

ratio of long to short prongs decreasing within a fascicle, ranging from 3.2:1 superiorly (Fig. 8.4G,H) to 2.3:1 II

inferiorly within tier 2 (Fig. 8.4I).

Ringent notosetae (category 1113)present in tier 3 (Fig. 8.4J,K), decreasing in size inferiorly within a B

fascicle; b:a ratios (Kudenov, 1987:184, Fig. 2) constant at .97:1 for both superior and inferior ringent notosetae

of tier 3.

Neurosetae longest superiorly, becoming abruptly short in middle and inferior regions of fascicle, all I

hollow, arrayed in tuft-like whorl; all of one kind, bifurcate, generally having characteristic appearance similar

to bifurcate notosetae with opaque white bands proximal to clear, heavily calcified prongs (Fig. 8.4L-V);

nearly all with short prong distinctly bidentate; some neurosetae unusual in having ringent-like dentitia on i

inner surface of long and outer surface of short prongs (Fig. 8.4L); with regular spiral pattern of minute filelike teeth well developed on all neurosetae in superior half of fascicle (Fig. 8.4V); ratios of long to short

prongs changing within a fascicle, ranging from 2.8:1 superiorly to 2.5:1 centrally to 2.6-3.3:1 inferiorly (Fig.

8.4L-Q), and from 2.5-2.8:1 anteriorly to 2.5:1 centrally to 2.7:1 posteriorly (Fig. 8.4N,O,R-U). Neuroacicula II

bifurcate, numbering 4-5 per fascicle, present in anterosuperior row, prong ratios around 3.4:1 (Fig. 8.4W).

Branchiae present from setiger 1to end of body, numbering 6-7 pairs per segment where best developed I 

in midbody segments, around 0.1 x length of longest notosetae, 0.5x length of dorsal and lateral cirri; having

2 cirriform filaments per gill, occasionally one, sometimes three in adults, only single filaments in juveniles. i

Branchiae unevenly spaced from one another, with large gap present between gills 1-2 especially conspicuous •

in juveniles. Lateral cirri arising between gills 2-3, all closely associated. g

Paired anal cirri terminal, cup-shaped, about as long as wide; anus terminal (Fig. 8.3F).

Biology. Relatively little is known about the biology of Euphrosine bicirrata. It has been collected H

from depths of 18-369 fms in green mud, and coarse sand-shells-rock. Prong lengths and ratios of bifurcate

notosetae in E. bicirrata vary systematically along both the body axis and within setal fascicles in relation to •

body size, based upon univadate and multivariate analysis of covadance (Vogt and Kudenov, 1994). Ii

Remarks. Euphrosine bicirrata is a highly characteristic species which can be readily identified by

the presence of bifurcate notosetae that tend to: (1) be longer than the width of the body; (2) interface with one

another to completely conceal branchiae, cirri, prostomium, camncle and dorsum; (3) have very well developed |

short prongs; (4) have fine parallel striae on the proximal cutting surface of the long prong; (5) be calcified

over both distal prongs; (6) have a deep cleft between both prongs; and (7) have a characteristic shape, i

Its neurosetae are, perhaps, even more characteristic in having (1) the short prong of most setae

distally bidentate; and (2) some of the larger neurosetae with ringent-like teeth on the inner surface of the long

prong, and outer surface of the short prong. The short neurosetal prong in most described species of Euphrosine

is entire; E. setosissima may be the only other congener suspected of having the short neurosetal prongs

distally bidentate (Kudenov, 1993). In addition, no other described Euphrosine species is known to have

neurosetae with the ringent-like notosetal dentitia described above forE. bicirrata (Kudenov, 1987; 1993). It i

may eventually be demonstrated that this kind of neuroseta represents a second category of euphrosinld

neurosetae; until then, it is here considered to be a bifurcate neuroseta.

Branchial features of adult E. bicirrata are easily distinguishable from those of other eastern Pacific

Euphrosine species. However, the branchiae of juvenile E. bicirrata are present as single, cirriform filaments,

and highly reminiscent of such species as E. armata Kudenov and E. setosissima Ehlers. Several obvious

features allow ready identification of such juvenile E. bicirrata: characteristic setal morphology; the relatively

large spacing between gills 1-2; and the tight association between gills 2-3 and the lateral cirri.

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226 I
!

I Euphrosine bicirrata is assigned to Group 1I (Kudenov, 1987:187, 189), which is characterized by

having ringent notosetae (category HA) and dorsoventrally bilobed caruncles (category K- 1), and is similar to

I seven expanded othertips. species Thesein having seven species dichotomously include:brEa.nching abyssalis branchiae Kudenov; havingE. borealis cirriformOfilaments rsted; E.that echidna lack sul_distally Kudenov;

E. heterobranchia Johnson; E. notialis Ehlers; E. orientalis Gustafson; and E. superba Marenzeller.

I Mexico D(Gulf istribution. of California). Northeast Pacific: Alaska, British Columbia, Washington, Oregon, California, and

I Literature Cited

I Banse, K. and K.D. Hobson. 1974. Benthic errantiate polychaetes of British Columbia and Washington. Bulletin of the Fisheries Research Board of Canada 185:1-111.

I Berkeley,toE.Canadian 1923. Polychaetous Biology, Ottawa, annelidsn.s.of1:203-218. the Nanaimo district. 1. Syllidae to Sigalionidae. Contributions

Blake, J.A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific I:

I Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae, and Maldanidae. Bulletin of the Biological Society of Washington 6:67-101.

I Clark, R.(eds.), B. 1969. Chemical Systematics Zoology and Phylogeny: Vol. 4. Academic Annelida, Press,Echi Newura,York. Sipunculida. Pp. 1-68.In: Florkin, M. and B. Scheer

Fauchald, K. 1974. Polychaete phylogeny: a problem in protostome evolution. Systematic Zoology 23:493-

I 506.

Fauchald, K. 1977. The Polychaete Worms Definitions and Keys to the Orders, Families and Genera. Natural

I History Museum of Los Angeles County, Science Series 28:1-190. Fauchald, K. and P.Jumars. 1979. The diet of worms: A study of polychaete feeding guilds. Oceanography

and Marine Biology Annual Review 17:193-284.

I Gustafson, G. 1930. Anatomische Studien tiber die Polychaeten-Familien Amphinomidae und Euphrosynidae.

Zoologiska Bidrag fr_ Uppsala 12:305-471.

I Hartman, O. 1940. Potychaetous annelids. Part II. Chrysopetalidae to Goniadidae. Allan Hancock Pacific Expeditions 7(3): 173-287.

i Hartman,Occasional O. 1959. Catalogue Paper 23:1-354. of the polychaetous annelids of the world. Part I. Allan Hancock Foundation

Hartman, O. 1964. Polychaeta Errantia of Antarctica. Antarctic Research Series 3:1-131.

I Hartman, O. 1965. Catalogue of the polychaetous annelids of the world, including additions and emendations

since 1959. Allan Hancock Foundation Occasional Paper 25:1-197.

I Hartman, O. 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation, University of Southern California, Los Angeles, 828 pp.

i Johnson, H.P. 1897. A preliminary account of the marine annelids of the Pacific coast, with descriptions of

new species. Euphrosynidae, Amphinomidae, Palmyridae, Polynoidae and Sigalionidae. Proceedings

of the California Academy of Sciences 1:153-190.

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i 227
|

Kudenov, J.D. 1974. The reproductive biology of Eurythoe complanata (Pallas, 1766) (Polychaeta: I

Amphinomidae). University of Arizona, Doctoral Dissertation. Pp. 1-128. IR

Kudenov, J.D. 1987. Review of the primary species characters for the genus Euphrosine (Polychaeta: •

Euphrosinidae). Bulletin of the Society of Washington 7:184-193. Ii

Kudenov, J.D. 1991. A new family and genus of the Order Amphinomida (Polychaeta) from the Galapagos

hydrothermal vents. In: Petersen, M.E. and J. Kirkegaard (eds.), Proceedings of the Second Intemational I

Polychaete Conference: Systematics, Biology and Morphology of World Polychaeta. Ophelia,

Supplement 5:111-120.

i

Kudenov, J.D. 1993. Amphinomidae and Euphrosinidae (Annelida: Polychaeta) principally from Antarctica, I

the southem ocean and subantarctic regions. Biology of Antarctic Seas XXII, Antarctic Research

Series 58:93-150.

Lamarck, J.B. 1818. Histoire naturelle des Animaux sans Vertrbrrs prrsentant les charact_res g_nrraux et m

particuliers de ces animaux, leur distribution, leurs classes, leurs families, leurs genres, et la citation

des principales esp_ces qui s'y rapportent; prrcrd_e d'une Introduction offrant la d&ermination des n

charact_res essentiels de l'Animal, sa distinction du vrg6tal et des autres corps naturels, enfm, l'expostion

des principes fondamentaux de la zoologie. Paris Vol. 5, Pp. 1-612 (Annrlides, Pp. 274-374).

Manton, S. 1967. The polychaete Spinther and the origin of the Arthropoda. Journal of Natural History, I

London 1:1-22.

Monro, C.C.A. 1930. Polychaete worms. Discovery Reports 2:1-222. N 

Monro, C.C.A. 1936. Discovery worms II. Discovery Reports. 12:59-198.

Monro, C.C.A. 1939. Polychaeta. Antarctic Research Expedition, 1929-1931. Adelaide, Australia. Reports, i

Series B. (Zoology and Botany) 4(4):89- i56.

Moore, J.P. 1905. New species of polychaetes from the north Pacific, chiefly from Alaskan waters. Proceedings i

of the Academy of Natural Sciences of Philadelphia 57:525-554. U

Moore, J.P. 1908. Some polychaetous annelids of the northern Pacific coast of North America. Proceedings of

the Academy of Natural Sciences of Philadelphia 60:321-364. I

u

Moore, J.P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern

California in 1904. Euphrosynidae to Goniadidae. Proceedings of the Academy of Natural Sciences

of Philadelphia 63:234-318. I

Orensanz, J.M. 1972. Los Annelidos Poliquetos de la Provincia Biogeografica Argentina. I. Palmyridae

(=Chrysopetalidae), Amphinomidae y Euphrosinidae. Physis 31:485-501. n

l

Pettibone, M. H. 1963. Marine annelid worms of the New England region. 1. Aphroditidae through

Trochochaetidae. Bulletin of the United States National Museum 227:1-356.

Pettibone, M.H. 1982. Classification of Polychaeta. In: Parker, S.P. (ed.), Synopsis and Classification of

Living Organisms. McGraw-Hill. Pp. 3-43.

Vogt, K.D. and J.D. Kudenov. 1994. Morphometric variation in bifurcate notosetae of two Euphrosine I

species (Polychaeta: Euphrosinidae). In: Dauvin, J.C., Laubier, L. and D.J. Reish (eds.) Actes de la

4_me Confrrence intemationale des Polych_tes. Mrmoires de Musrum National D'Histoire Naturelle n

(A) 162: 291-298. !

Williams, T. 1851. Report on the British Annelida. Report of the British Association for the Advancement of

Science, London 21:159-272. U

228 I
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- 9. FAMILYONUPHIDAEKINBERG, 1865

I by

Brigitte Hilbig 1

l Introduction

I Onuphids are midsized to large worms with dorsoventrally flattened bodies anda distinctive array of

prostomial appendages, including five occipital antennae, two frontal palps, and two labial palps. This

I arrangement of antennae and palps is consistent in all onuphids, in contrast to the morphologically similar eunicids where the number of antennae varies from one to five. The first few segments, particularly the

parapodia, areoften specialized and, in combination with the head appendages, make the Onuphidae the most

I spectacular and morphologically diverse family among the Eunicida. The largest onuphids are the Australian beachworms (Australonuphis), living representatives of which have been known to reach a length of 300 cm

with more than 1000 segments.

I

Morphology

I

The prostomium is usually round to pentagonal and, because of its relatively small size, is often

I othat bstructe maydbebyclosed the ceratophores to a circle byofthethesmall, occipital simple antennae frontal(Fig. palps9.1(also A). The calledantefrontal nnae are antennae) arrangedthatinarise a crescen fromt

the frontal margin of the prostomium. The ceratophores of the occipital antennae are of varying length and

I ulength suallyofringe thedcer(rarel atophores y smooth); relativethe tonuthe mberprostomial of tings islengoftethnisimused portant to distinguish for species genera. discriminAantion, other apair nd theof

simple, short appendages are the ventrally situated labial palps (Fig. 9.1C). Eyes are often present, one pair

i nearthe bases of the frontal antennae and one pair between the bases of the outer (anterior) and inner (posterior)

lateral antennae. The posterior pair is usually the larger, and may be lensed in a few taxa. A pair of crescentic

to straight nuchal grooves is located near the posterior margin (Fig. 9.1B). The peristomium consist of a

i single apodous The eversible ring thatandmaymuscular or may npharynx ot bear aispair armedof tentacular with mandibles cirri, usanduallyfiveonmaxillae. the anteriorThemargin. mandibles

consist of two elongate shafts that bear a roughly crescentic, often dentate, calcified cutting plate (Fig. 9.1D);

I the adulremains t ones. The of themaxilla juvenilee aremandibles asymmetcan rical;oftethneybetyspical een asly bhavlack, e shoL-shaped rt and wide, structuoften res insquare the cucattrringierplaslink te ofedtheto

falcate MI enveloping a pair of triangular MII with serrated cutting edge and a left MIII of similar shape and

I deandntiatipair on (ofthesrimall, ght MII usuall I are y uniden absentt)at. eAnte or srmooth ior to MI MV-MHI, are located. a pair ofThe rounrideghd tMIV MIV with is often serrat largedercuthan ttingtheedges left

one, bears more teeth, and extends further back than the latter (Fig. 9.1E). A pair of small MVI is present in

i a few genera. As in the other eunicemorph families, the morphological details of the maxillary apparatus are

among the most important taxonomic characters for species discrimination.

I l ENSRConsultingandEngineering,89WaterStreet,WoodsHole,Massachusett02543 s

i 229
I

I

oLan_--,Pa I

tC nuO I

oLan -- _

' I

cest B

iLan _._ ceph

"" _ __--- lip

. C li

A "_ in

..,,._,_:_ _._._ ,

cu,_, ,v_ _ _!_:-,v

,,,_.._ _ ...... l

.....! ,

,.a. i ' ' II

' '

D f,,,, _mC I _ E |

Figure 9.1. Morphological features of onuphids: prostomial and peristomial appendages and jaws. A, anterior I

end, dorsal view; B, C, schematic dorsal and ventral views ofprostomium; D, mandibles; E, maxillae.

(all from Paxton, 1986a). I

230 I
!

I The anterior 1-8 setigers usually bear modified parapodia that are directed forward rather than to the

sides, and often displaced dorsally (Hirsutonuphis) or ventrally (Americonuphis). In some genera, the first

I fewothersparapodia they extend are extremely about as farprolonged as the frontal and project palps andwelldiffer beyond fromtheunmodified prostomiumpampodia (Longibrachium only slightly ), while in thein

relative length of dorsal and ventral cirri and postsetal lobes. The genus Nothria is characterized by bluntly

i rounded, present infleshy, a smallauricular number presetal of anteriorlobes segments on the and modified replacedparapodia by glandular (Fig. 9.2A). pads inVentral subsequent cirri setigers. are generally The

position of the first parapodium with padlike ventral cirri is used for species discrimination. Dorsal cirri are

i present throughout, although they are largest in anterior setigers. The bases of the dorsal cirri are the only remainders of the notopodia, supported by a bundle of fine notoaciculae that extends into the proximal part of

the dorsal cirrus (Fig. 9.2B). There are no notosetae. Branchiae are usually present and may be simple,

pectinate, or spiraled (Diopatra) (Fig. 9.2C).

I The setae include a wide variety of simple and compound limbate (Fig. 9.2F), and pectinate setae

(Fig. 9.2D, E) and compound, pseudocompound, and simple hooded hooks (Fig. 9.2G-K). Their distribution

I genus along the andbody and morphological details, such as number of teeth, are of great taxonomic importance for species discrimination. The division of the onuphids into two subfamilies, the Onuphinae and

Hyalinoecinae, is based on the presence of a lower fascicle of simple limbate setae throughout (Hyalinoecinae)

i or up to the first setiger with subacicular hooks (Onuphinae). In the first case, the subacicular hooks are positioned in the middle of the fascicle ("intmfascicular hooks" of Fauchald (1982)), in the second case these

hooks occupy the ventralmost position of each fascicle.

I discriminating The presence charactersof very so mansimple; y countable however,mosome rphologi of these cal features charactersmakes are growth- the establishment or size-dependent of species (see

below), and individual variability has to be kept in mind when following keys or descriptions.

I

Taxonomic History

I

Due to their generally large size and richness of morphological features, onuphid species have largely

i been valid since the majority were described about 100 to 150 years ago. In contrast, generic concepts have been changed several times, and the establishment of monophyletic taxa with clearly defined character sets

was not achieved until the early 1980's when Fanchald (1982a) and Paxton (1986a, b) published their invaluable

i revisions of the family. The onuphids now include 22 genera and about 220 species. For more details, the reader is referred to the excellent accounts by Fauchald (1982a) and Paxton (1986a, b).

I Distribution and Biological Notes

I Onuphids are distributed worldwide and in can be found from the intertidal to abyssal depths. Fauchald

(1982a) provided a comprehensive overlook over distributional patterns of some onuphid genera, and it appears

that the eastern Pacific Ocean is a center of radiation for this family. All onuphids are tubicolous, although

I not necessarily sessile; Orensanz (1990) def'med three different ecotypes, including (1) species thatlive partially buried in the sediment and produce permanent tubes with a more or less elastic inner layer and an outer layer

consisting of sediment or shell particles (many genera); (2) species that can move around on the sediment and

I carry their tubes with them; these tubes are quill-like (Hyalinoecia) or flat, two-layered, with a cover of flat shell or rock particles (Nothria); and (3) species that are epibenthic scavengers on sandy beaches and produce

delicate, elastic, temporary tubes (Australonuphis). The tubes are often much longer than the animal.

I

I 231
I

A /dC I

,

_ vC dC I

br I

|

I'/ ;/ I

.oo, ./ , .oo°> |

E _

tt

!

G H I J, K'' |

Figure 9.2. Morphological features of onuphids: pampodia and setae. A, first parapodium, Nothria; B, anterior 1

parapodium, Onuphis; C, middle parapodium, Diopatra; D, E, scoop-shaped and flat pectinate I

setae; F, compound spiniger (Mooreonuphis); G, subacicular hook; H, I, pseudocompound hooks; J,

simple hooded hook ("large median hook"); K, pseudocompound hook (Paradiopatra). (all from

Paxton, 1986a). I
!

I Feeding habits of onuphids were summarized by Fauchald and Jumars (1979). From the often

contradictory accounts in the literature, they concluded that most onuphids are probably ominvomus scavengers

i that opportunistic may specibehavior alize in aincertain Hyalinoecia food itemspp.;if itseveral is the only hundred one available. specimens Dayton were seenandtoHessler aggregate(1972) aroundobserved a fish

cadaver. At least some populations of Diopatra ornata seem to feed exclusively on kelp (Fauchald and

i Jumars, The 1979). reproductive biology ofonuphids is not well known; sexes are generally separate, and no asexual

reproduction is known. According to the few existing investigations, eggs are generally large and yolky, and

I brood pelagic,protection and the juveniles in the maternal may stay tubeinseems the parent's to be common tube until (Hsieh theyandareSimon, about 171987). to 20Larvae setigersarelong; oftendire non- ct

development is known forHyalinoecia araucana (Carrasco, 1983). Many taxonomically important characters,

I such 1986a;asHsieh the setae, and jaws, Simon,parapodial 1987); a synopsis cirri, andofbranchiae, larval morphology differ in larvae of onuphids and adults was compiled (Blake, 1975b; by Bhaud Paxton, and

Cazaux (1987). From these and other studies, it is clear that some characters used for species discrimination

i are occurrence only validmayinmove adults; for example, gills are always first present on setigers 6-8, and the setiger of their fu-st to a more anterior or posterior position during postlarval development; ventral cirri and

postsetal lobes are cirriform and prominent in a smaller number of anterior parapodia in younger specimens.

i Pseudocompound hooks may occur in a greater number of anterior setigers with increasing body length, and special larval setae, reminiscent of eunicid compound setae, are present in very young individuals; subacicular

hooks may start on a more anterior setiger in younger specimens. Fanchald (1982a, b) was the first to draw

i the attention of systematists to these growth dependent changes by providing tables and graphics depicting ranges of many key taxonomic characters. Some larval characters, such as the distribution of larval setae,

may be species specific (Hsieh and Simon, 1987).

I

I Key to the Onuphidae

1A. First three setigers greatly modified, with long, slender, distally recurved hooks, with setaI

very sacs

extending back to setiger 50, at least through setiger 25; antennae no longer than first 2-3 setigers.

....................................................................................................... Rhamphobrachium longisetosum

I lB. _mterior setigers modified, but carrying pseudocompound or simple hooded hooks; at least inner

lateral antennae longer than first 2-3 setigers .................................................................................. 2

I 2A. Branchiae spiraled (Fig. 9.2C) ................................................................................ genus Diopatra 2

2B. Branchiae simple, pectinate, or absent ............................................................................................. 3

I

3A. First parapodia with prominent, auricular presetal lobes in addition to subulate postsetal lobes (Fig.

i 9.2A); peclinate setae scoop-shaped (Fig. 9.2D); second parapodia with foliaceous, prominent presetal lobes; hooks in setiger 1 simple, in setigers 2 and 3 pseudocompound; branchiae from setiger 8 .....

.......................................................................................................................... Nothria occidentalis

I 3B. Presetal lobes inconspicuous throughout; pectinate setae flat (Fig. 9.2E) ....................................... 4

i 2Notfoundinpresentmaterialb,utcommonin shallowerwater.

I
!

!

4A. Ceratophores of occipital antennae distinctly longer than prostomium; styles of outer lateral antennae

shorter than ceratophores ............................................................................ genus Onuphis ............ 5 i

4B. Ceratophores of occipital antennae as long as prostomium or shorter; styles of outer lateral antennae

longer than eeratophores .................................................................................................................. 9

I

5A. Branchiae present from setiger 3-6 .................................................................................................. 6

5B. Branchiae present from setiger I (2 or 3 in smaller specimens) ....................................................... 7 i

6A. Pseudocompound hooks in setigers 1-3 ....................................................... Onuphis geophiliformis

6B. Pseudocompound hooks in setigers 1-4 ..................................................................... Onuphis affinis I

7A. Branchiae in middle setigers winged, distinctly wider and thicker than in preceding and subsequent i

segments; pseudocompound hooks in setigers 1-4, subacicular hooks from setiger 8 or 9 ................

............................................................................................................................... Onuphis elegans

7B. Branchiae slender throughout, not winged ....................................................................................... 8 •

8A. Subacicular hooks first present in setiger 12 or 13 ............................................ Onuphis iridescens i

8B. Subacicular hooks first present in setiger 8 or 9 ............................................ Onuphis intermediates

9A. Hoods of pseudocompound hooks projecting well beyond apical tooth, acutely pointed (Fig. 9.2K); I

branchiae pectinate, from setiger 2-4 through about 30, maximally 7 filaments ...............................

........................................................................................................................... Paradiopatra parva •

9B. Hoods of pseudocompound hooks not prolonged, oblique or bluntly rounded ............................... 10 1

i

10A. Lower fascicle of limbate setae compound (Fig. 9.2F) ......................... genusMooreonuphis ...... 11 i

lJ

10B. Lower fascicle of limbate setae simple; branchiae pectinate, first present between setigers 2 and 4,

with maximally 10 filaments; pseudocompound hooks in 7 or 8 setigers, subacicular hooks from i

setiger 18-25 ........................................................................................... Kinbergonuphis vexiUaria |

11A. Tridentate pseudocompound hooks in 8 setigers, median ones often simple in last 3 or 4 parapodia; 1

compound spinigers present from setiger 7 to 19, then replaced by subacicular hooks .....................

.................................................................................................................... Mooreonuphis nebulosa

11B. Pseudocompound hooks in 4 setigers ............................................................................................. 12 i

1

12A. Branchiae simple, f'n-stoccurring between setigers 11and 20 ....................... Mooreonuphis exigua •

12B. Branchiae pectinate, first occurring between setigers 7 and 15, with maximally 5 filaments ............ II

.......................................................................................................... Mooreonuphis segmentispadix

!

!

234 I
I

I Description of Species

I with ten The or 11onuphid species fauna in six genera; in the Santa two species Maria were Basinfound and Western that are newSantatoBarbara science. Channel The onlyisgenus fairlycommon diverse

to the eastern Pacific, but not represented in the Phase I and II collections, isDiopatra. The following species

I are described: Kinbergonuphis vexillaria (Moore, 1911)

Mooreonuphis exigua (Shisko, 1981)

i Mooreonuphis nebulosa (Moore, 1911) segmentispadix (Shisko, 1981)

Nothria occidentalis Fauchald, 1968

Onuphis affinis Hilbig, new species

I Onuphis elegans geophiliformis (Johnson, (Moore, 1901) 1903)

Onuphis iridescens (Johnson, 1901)

i Onuphis sp. intermediates sensu Hobson, 1971 Paradiopatra parva (Moore, 1911)

Rhamphobrachium longisetosum Berkeley and Berkeley, 1938

I Genus Kinbergonuphis Fanchald, 1982

i Type Species. Onuphis tenuis Hansen, 1882

Diagnosis. Ceratophores of occipital antennae short, usually no longer than prostomium, with 3 to

i 10 rings; outer (anterior) styles at least as long as their ceratophores. Pseudocompound hooks with short, blunt hoods, usually tridentate; median ones often becoming simple after setiger 4 ("large median hooks" of

Fauchald, 1982). Compound spinigers absent. Modified parapodia slightly or not at all prolonged. Branchiae

i usually pectinate, rarely simple or absent; first occurring on setiger 6, rarely before or after. Body size generally small, length rarely surpassing 100 mm, width typically to 2 mm.

Remarks. The generic definition is taken from Paxton (1986) and includes her minor revisions of

I Fauchald's main distribution (1982) original diagnosis. Kinbergonuphis is a large genus with more than 30 species, with its in shelf depths off both coasts of the Americas. While the majority of Pacific species occur

off Central and South America, only three species have been described from off California, the poorly known

i K. paradiopatra (Hartman, 1944) from intertidal depths, and the slope species K. proalopus (Chamberlin, 1919) and K. vexilIaria (Moore, 1911). Only the latter was found in the Phase I and II material, and the Santa

Maria Basin may be the northern boundary of its distribution.

I Kinbergonuphis vexillaria (Moore, 1911)

I Figure 9.3

Onuphis vexillaria Moore, 1911:266-269, pl. 17, figs. 69-76.--Fauchald, 1982:54-55.

I Kinbergonuphis vexillaria: Paxton, 1986a:54.

sp. A Steinhauer and Imamura, 1990:F-1.

Onuphis sp. A Lissner et aI., 1986:D-11.

I Onuphis sp. B Lissner et al., 1986:D-11.

Material Examined. California, Santa Maria Basin, off Point San Luis, Sta. 27 (1); off Point Sal,

I Sta. 35 Sta. R-7 off Point Sta. 77

(1), (1); Arguello, (2).

i 235
!

I

I

Description. All specimens incomplete; length to 159 rnm, width to 4 mm (measured behind pharyngeal •

region, parapodia excluded), setigers to 242. Body wide and dorsally flattened, with crowded segments,

except for pharyngeal region encompassing first 10 setigers. Color in alcohol pale with iridescent cuticle. 1

Prostomium small, rounded, almost concealed by bases of occipital antennae. Frontal antennae

ovoid, shorter than prostomium; ceratophores of occipital antennae shorter than prostomium, each with 5-8

rings; outer (anterior) lateral antennae as long as peristomium and first 2-3 setigers, styles much longer than II

their ceratophores; inner (posterior) lateral antennae as long as peristomium and f'u-st8-14 setigers; median

antenna as long as peristomium and first 5-10 setigers; all styles gently tapering toward long, filiform distal

portion (Fig. 9.3A). Eyes and nuchal organs not observed. Peristomium shorter than following setigers, with i

very long, slender tentacular cirri projecting just beyond frontal margin of prostomium.

Maxillae hard, calcified, with dark teeth; maxillary carriers very short, only 1.5 times as long as

wide; MI gently curved, about 3 times as long as carriers; MII 5-6+7-9, MIII 7-8+0, MIV 6-8+6-8, MV very i

small, delicate, squarish plates without teeth (with single tooth according to Moore, 1911) (Fig. 9.3B).

First 4 parapodia modified, slightly enlarged, pointing forward, with long, slender, tapered dorsal and 1

ventral cirri and postsetai lobes (Fig. 9.3C); ventral cirri replaced by pads in setiger 9 or 10, last ventral cirrus 1

sometimes spherical rather than subulate; postsetal lobes subulate through setiger 16-30, then gradually

i

shortening and turning conical; dorsal cirri always as long as longest branchial filament, becoming filiform in

middle and posterior setigers. Branchiae present from setiger 2-4, simple at first, quickly becoming pectinate 1

(Fig. 9.3D, E), with up to 12 filaments arising from thick stem, diminishing toward posterior end, in one

i

specimen simple from setiger 70. Pseudocompound hooks tridentate, with short, oblique hoods (Fig. 9.3F), 1

present in setigers 1-7 or 8, median one sometimes turning into large median hook in last 2 or 3 setigers; 1

simple limbate and pectinate setae present from setiger 2; limbate setae densely covered with surficial spines 1

(Fig. 9.3G); pectinate setae oblique, with about 10 long, slender teeth, terminal tooth on one side much longer

than others in anterior setigers, all teeth equal inposterior setigers (Fig. 9.3H, I). Subacicular hooks present 1

from setiger 18-25 (Fig. 9.3J). Neuropodia supported by 2-4 neuroaciculae, number declining toward posterior i

end; base of dorsal cirri with dense fascicle of about 12-15 delicate, transparent notoaciculae.

Pygidium unknown, i

Remarks. Kinbergonuphis vexillaria differs from allits congeners by the presence of gills on setigers

2-4; it is related toK. vermillionensis (Fauchald, 1968),K. nannognathus (Chamberlin, 1919), andK. proalopus •

(Chamberlili, 1919) that all occur slightly to the south of the type locality ofK. vexillaria and are characterized I

by pectinate branchiae. Aside from the first occurrence of gills, K. vexillaria differs from K. vermillionensis

by the distribution of subacicular hooks (present from setiger 12 inK. vermillionensis, not before setiger 18 III

in K. vexilIaria) and the number of maxillary teeth (MII 14+12 in K. vermillionensis, maximally 6+9 in K. I

vexillaria). K. nannognathus and K. proaIopus both have pseudocompound hooks in 6-7, rather than 7-8

setigers.

Type Locality and Type Specimens. Pacific Ocean off California, Soledad Hill, Point La Jolla, 1

Albatross Sta. 4326: holotype (USNM 19061).

Habitat. Moore (1911) found K. vexillaria in soft green mud; the Phase 1and 2 specimens occurred i

in fine sediments with at least 75% silt and clay.

Distribution. Southern and central California, 442 - 611 m.

I

!

236 i
I Figure 9.3. Kinbergonuphis view; D, middlevexillaria parapodium, : A, anteri anterior or end,view; dorsalE,view; posterior B, maxill parapodium, ae; C, parapodi anterior um 2,view; anteriorF,

pseudocompound seta; G, simple seta; H, peetinate seta, setiger 2; I, same, setiger 10; J, subacicular

i hook.

I 237
!

Genus Mooreonuphis Fauchald, 1982 I

Type Species. Onuphis nebulosa Moore, 1911

Diagnosis. Ceratophores of occipital antennae short, no longer than prostomium, with maximally 5 I

rings; outer (anterior) styles at least as long as their ceratophores. Pseudocompound hooks bi- to tridentate,

with short, blunt hoods; median ones sometimes becoming simple ("large median hooks" of Fauchald, 1982). l

Lower setae compound spinigers in segments between last one with pseudocompound hooks and first one with

subacicular hooks. Modified parapodia not prolonged. Branchiae simple, rarely pectinate, usually present

after setiger 17, rarely from 6 or 7. Body size small, length less than 100 mm, width up to 2 mm.

Remarks. To distinguish this genus from the similar Kinbergonuphis, several parapodia should be

examined carefully for the presence of compound spinigers. These setae may be difficult to observe especially

in small individuals. Differences in the development of branchiae are also helpful; they are usually pectinate I

and present from setiger 6 in Kinbergonuphis, usually simple and present after setiger 17 in Mooreonuphis.

The distribution of Mooreonuphis is somewhat similar to that of Kinbergonuphis, with the majority of species

being found along the Atlantic and Pacific coasts of the Americas. Four species are known from California,

and all but one (M. veleronis (Fauchald, 1980)) are present in the Santa Maria Basin.

Mooreonuphis exigua (Shisko, 1981) I

Figure 9.4 I

Nothria exigua Shisko, 1981:973-976, fig. 3.

Mooreonuphis exigua: Paxton, 1986:56.---Steinhauer and Imamura, 1990:F-1. I

Material Examined. California: Santa Mafia Basin, off Purisima Point, Sta. R5 (1).

1

Description. Length to 24 mm, width to 1 nun, segments to 100. Body slender, flattened; Per•store•urn 1

and tilrst 20 segments with wide reddish brown dorsal pigment bands; first setiger light colored, subsequent

segments gradually darkening. I

Pmstomium relatively large, rounded, bearing globular frontal antennae and occipital antennae arranged I

in a crescent. Ceratophores of lateral antennae with 3-4 rings, ceratophore of median antenna smooth; outer

(anterior) lateral antennae as long as peristomium and first 1or 2 setigers, inner (posterior) lateral antennae as •

long as peristomium and tin'st 10 setigers, median antenna as tong as peristomium and tin'st 8 or 9 setigers. I

Two pairs of eyes, located behind and in front of inner lateral antennae (Figure 9.4A). Peristomium about half

as long as first setiger; tentacular cirri reaching bases of inner lateral antennae.

Jaws poorly sclerotized; larval mandibles distinctly visible as black feature on weakly calcified cutting

plate; maxillae transparent with dark teeth. Maxillary formula: MI 1+1,strongly falcate; MII 6+6, MIII 7+0,

MIV 7+8, MV oval plates without dentition. •

Anterior parapodia only slightly modified (Fig. 9.4B); fLrSt2 pairs directed anterolaterally. Postsetal I

lobes digit•form through setiger 8, papilliform after a few transitory setigers; ventral cirri replaced by pads on

setiger 5. Branchiae simple, first present on setiger 11 in examined specimen (range 11-20). Simple limbate •

setae present throughout, winged part timely serrated and covered with surficial spines (Fig. 9.4C); pectinate

setae present on most setigers, with few teeth. Slender and stout pseudocompound hooks present in setigers 1-

4, tridentate, with proximal tooth in slender hooks minute and easily overlooked (Fig. 9.4D, E). Compound •

spinigers present on setigers 5-16 (range 4-5 to 11-16); end of shaft and blade densely covered with surficial I

spines (Fig. 9.4F); subacicular hooded hooks starting on first setiger without compound spinigers.

Tubes not observed, thin and transparent according to Shisko (1981). i

238 I
!

!

' I I

I !i.

I -

I A _ _ / ,\

I

! ""_" C E i

I B .i.i;ii!_-_:_

I

Figure 9.4. Mooreonuphis exigua: A, anterior end, dorsal view (after Shisko, 1981); B, anterior parapodium,

i anterior view; C, simple seta; D, slender pseudocompound hook; E, stout pseudocompound hook; F,

compound spiniger.

I Remarks. This species is somewhat similar to M. nebulosa (see below), especially with regard to

pigmentation, and may be mistaken for its juvenile. The two species differ mainly in the distribution of

I pseudocompound hooks (setigers 1-4 in M. exigua, 1-8 in M. nebulosa) and the first appearance and shape of the gills (present from setiger 11-20, all simple in M. exigua, 6-8, with up to 6 branches in M. nebulosa).

Type Locality and Type Specimens. Southern California Bight: holotype (USNM 63081) and 4

I paratypes (USNM 63082-63085).

Habitat. M. exigua prefers sandy sediments, but was also found in silt and clay.

I Distribution. Southern and central California, 65 - 319 m.

239
!

Mooreonuphis nebulosa (Moore, 1911) !

Figure 9.5 I

Onuphis nebulosa Moore, 1911:269-273, pl. 17, figs. 58-68.--Hartman, 1944:75-78, pl. 4, figs. 76-85.

Mooreonuphis nebulosa: Fauchald, 1982:56-57, fig. 17a.--Lissner etal., 1986:D-11. •

Mooreonuphis nr. nebulosa: Steinhauer and Imamura, 1990:F-1 1

Mooreonuphis cf. nebulosa: Gathof, 1984:39-23, figs. 39-19, 39-20a-i.

I

Material E_mmed. California: Santa Maria Basin, off Purisima Point, Sta. R5 (1); off Point

Arguello, Sta. 64 (1); Western Santa Barbara Channel, Sta. 85 (rock) (1), Sta. 94 (1). I

Description. Length to 125 mm, width to 2.0 nun, segments to 175. Body slender, linear, dorsoventrally 1

flattened behind pharyngeal region, tapering toward pygidium within last 5 setigers (Fig. 9.5A, B). Peristomium

and first 15-25 setigers with wide, brownish transverse pigment bands across dorsum; darkly pigmented •

peristomium sometimes contrasting against relatively pale first setiger. 1

Prostomium rounded, without eyes. Ceratophores of occipital antennae short, crowded, with 3 or 4

rings; outer (anterior) lateral antennae as long as peristomium and first 1or 2 setigers, inner (posterior) lateral •

ones as long as peristomium and first 6 or 7 setigers, median one as long as peristomium and first 5 or 6 I

setigers; frontal antennae short and tapered. Peristomium only about 1/4 as long as f'trst setiger and about as

wide, bearing two relatively short tentacular cirri. •

Maxillae poorly sclerotized; with short, squarish carders about half as tong as MI; MII 7-10+8-10,

MIII 8-10+0, similar in size to MII; MIV 6+8, crescent-shaped plates; MV unidentate or smooth, roughly

triangular, small plates (Fig. 9.5C). Mandibles about as long as maxillae, with cutting plate deeply incised 1

medially and irregularly serrated (Fig. 9.5D). 1

First 7 or 8 pampodia slender, about aslong as segmental width, supported by 2 notoaciculae projecting

into base of dorsal cirri and 2-3 neuroaciculae with long, geniculate free tips (Fig. 9.5E), with digitiform II

postsetal lobes and clavate dorsal and venlral cirri projecting about as far as postsetal lobes; slightly displaced

ventrally and directed forward on first two setigers, gradually moving to lateral position within following few

setigers. Ventral cirri usually clavate to conical through setiger 10 (range 7-10), replaced by rounded pads _l

from setiger 11 (range 8-11); postsetal lobes digitiform through first 20 setigers, then gradually diminishing in

size and papilliform from about setiger 50; dorsal cirri present throughout, markedly decreasing in width

within branchial region but of almost equal length throughout (Fig. 9.5F, G). Gills present from setiger 7 1

(range 6-8), starting as single filaments, with 4 branches where best developed, absent from posterior region.

Tridentate pseudocompound hooks presentin first 8 setigers, of two kinds: (1) slender, with long shaft

and blade, with terminal tooth much larger than proximal ones (Fig. 9.5H), (2) stout, short-bladed, terminal 1

tooth smallest; some or all stout pseudocompound hooks replaced by simple hooks ("large hooks" ofFauchald)

from setiger 4 or 5 on (Fig. 9.51,J). Simple limbate capillaries present throughout (Fig. 9.5K), pectinate setae

starting on anterior setiger and continuing to almost end of body; with about 10 short, fine teeth (Fig. 9.5L); 1

compound spinigers present on setigers 7 to maximally 19, usually to setiger 13 or 15. Subacicular hooded

hooks present from setiger behind last one beating compound spinigers; two per fascicle, stout, amber colored,

present to end of body (Fig. 9.5M). 1

Tube flattened, consisting of relatively stiff mucous lining, covered with often quite large sand and

shell particles. m

Remarks. This species is readily distinguished from its congeners by the large number of segments I

bearing modified parapodia. None of the species specific characters of the single specimen originally called

Mooreonuphis nr. nebulosa were found to be outside the variability documented in the literature. The specimens 1

from the Gulf of Mexico (Gathof, 1984) were reported to have longer parapodia than the Californian ones and •

1

2o !
!

!

! _ ii

! i i J"

i .... .

!

I M

!

• ..,:.._ _ br

-_.

I

C

a oc

i Figure 9.5. Mooreonuphis nebulosa: A, anterior end, dorsal view; B, posterior end, dorsal view; C, maxillae;

D, mandibles; E, free end of acicula; F, anterior parapodium, anterior view; G, middle parapodium,

anterior view; H, slender pseudocompound hook; I, stout pseudocompound hook; J, large hook; K,

I limbate seta; L, pectinate seta; M, subacicular hooks. (C after Moore, 1911; F, G after Gathof, 1984).

I 241
!

were therefore described as M. cf. nebuIosa. However, a comparison of Moore's (1911) original illustration I

and that of Gathof (1984) does not reveal any readily visible differences in the proportions of the parapodia.

Type Locality and Type Specimens. Monterey Bay, off Point Pifios Lighthouse: holotype (USNM 1

16881) and 2 paratypes (USNM 17062).

Habitat. M. nebulosa is found in various sediments ranging from clayey silt to sand and gravel; the I

examined specimens were collected in sediment consisting of approximately 70 to 80% sand and only 3 to 5%

clay. m

Distribution. California; Gulf of Mexico; 19 - 142 m. I

1

Mooreonuphis segmentispadix (Shisko, 1981)

Figure 9.6 !

Onuphis segmentispadix Shisko, 198I :978-981.

Mooreonuphis segmentispadix: Paxton, 1986:56.--Lissner et al., 1986:D- 11.

Mooreonuphis sp. B Lissner et al., 1986:D-11.

!

Material Exara_ed. California: Santa Maria Basin, off Point Sal, Sta. 36 (2), Sta. 108 (4); off

Purisima Point, Sta. 46 (3).

Description. Length at least 28 mm, width to 2 mm, segments over 100. Body flattened except for

pharyngeal region, with crowded segments, 2 to 3 times as wide as long anteriorly, at least 4 times as wide as

long after first 5 setigers. Dorsum with wide transverse bars of reddish brown pigment through setiger 20 to •

30; pigmentation gradually fading in subsequent setigers, middle and posterior part of body pale. 1

Prostomium rounded, with globular, slightly tapering frontal antennae; eyes not observed in California

specimens, two pairs present according to Shisko (1981). Ceratophores of occipital antennae with 3 to 4

rings. Outer (anterior) lateral antennae as long as peristomium and first 2 setigers, inner (posterior) lateral

antennae as long as peristomium and first 14 setigers, median antenna as long as peristomium and fLrSt11

setigers. Posterior prostomial margin with dark transverse pigment band delineating nuchal organs (Fig. il

9.6A). Peristomium short, bearing long cirri projecting beyond antennal bases.

Maxillae with slender, triangular carders and strongly falcate MI; MII 8-9+8, MIII 9+0, MIV 8-

10+10-14, arching over MII and MIII; MV rounded plates with smooth edges (Fig. 9.6B). All maxillae 1

poorly sclerotized and calcified, transparent with dark teeth.

First 3 parapodia elongate (Fig. 9.6C), slightly displaced ventrally and pointing forward; subsequent I

parapodia inserted laterally and pointing to the sides and upward. Postsetal lobes conical through setiger 14 1

to 17, ventral cirri padlike from setiger 5 or 6; gills typically first present on setiger 7 or 8 (range 7-15), simple m

at first, pectinate where best developed, with 4-5 branches (Fig. 9.6D, E). Pseudocompound hooks tridentate, IIIIll

present in setigers 1 to 4, no large hooks (Fig. 9.6F); compound spinigers in ventralmost position of fascicle

in setigers 5 to 15, distal end of shaft and entire blade covered with surficial spines (Fig. 9.6G); subacicular !

hooks from setiger 16 (Fig. 9.6H); simple limbate setae present throughout, covered with surficial spines,

widest and shortest in ventral part of fascicle (Fig. 9.61, J); pectinate setae in median and posterior setigers, I

with about 10 subequal teeth and one outer tooth about 4 times as long (Fig. 9.6K). Notopodia with 6 fine

aciculae in anterior setigers, reduced to 3 in posterior setigers; neuroaciculae 4 to 2 from anterior to posterior,

with very long tips resembling short limbate setae (Fig. 9.6L). I

II

Remarks. Several specimens originally designatedMooreonuphis sp.B in the MMS Phase 1material

were identified as M. segmentispadix here because no differences could be detected; the original voucher sheet tim

was obviously prepared from a juvenile specimen. B

242 II
!

!

! /

I! A : _

!

!

!

|

!

Figure 9.6. Mooreonuphis segmentispadix: A, anterior end, dorsal view; B, maxillae; C, parapodium 1, anterior

i view; D, prebranchial parapodium, anterior view; E, parapodium 12, anterior view; E, far posterior

parapodium, anterior view; F, slender and stout 0eft) pseudocompound hook; G, compound spiniger,

H, subacicular hook; I, upper simple seta; J, lower simple seta; K, pectinate seta; L, free tip of

acieula.

!

I 243
!

i

Type Locality and Type Specimens. Southem California: holotype (USNM 63074) and 6 paratypes i

(USNM 63075-63080).

Habitat. M. segmentispadix lives in a wide variety of sediments from silt to sand and gravel, but n

seems to prefer sandy substrata.

Distribution. Southern and central California, 63 - 670 m. i

Genus Nothria Malmgren, 1866

Type Species. Onuphis conchylega Sars, 1835 1

Diagnosis. Ceratophores of occipital antennae barely longer than wide, smooth or with 3-5 rings. I

Anterior 2

to 3 parapodia modified, first parapodium prolonged; presetal lobes large, auricular in setiger 1, i

rounded to conical in subsequent setigers, gradually diminishing; postsetal lobes subulate in some anterior

setigers; ventral cirri subulate in 2 to 3 setigers, followed by 1or 2 transitory parapodia with spherical ventral

cirri, replaced by pads after that. Branchiae simple if present, first occurring on between setigers 8 and 13. i

Simple to pseudocompound hooded hooks in 2 or 3 setigers, uni- to bidentate; pectinate ans limbate setae

m

present from setiger 2; pectinate setae scoop-shaped, forming often dense fascicles; limbate setae geniculate

with broad wings, resembling lumbrinerid setae. Subacicular hooks first present between setigers 8 and 14, i

usually with long shafts and relatively small heads. roll

n

Nothria occidentalis Fauchald, 1968 i

Figure 9.7 i

Nothria conchylega occidentaIis Fauchald, 1968:20-21, pl. 5, figs. a-n.

Nothria occidentalis: Fauchald, 1982a:95, fig. 27d. i

Paranorthia sp. A Steinhauer and Imamura, 1990:F-1.

Material Examined. California: Santa Mafia Basin, off Purisima Point, Sta. R-5 (1). i

Description. Length more than 14 mm, width to 2 mm, segments more than 30. Body slender, linear,

slightly flattened; color in alcohol pale. i

Prostomium rounded, with antennae inserted along outer margin; two large posterior eyes and two

smaller anterior ones. Frontal palps small, conical; ceratophores of occipital antennae as long as wide, with

3-4 very short rings; outer (anterior) lateral antennae aslong as peristomium and first 2 setigers, inner (posterior) i

lateral antennae as long as peristomium and first 8-9 setigers median antenna as long as peristomium and first

3-8 setigers (Fig. 9.7A). Peristomium short, about one-third as long as prostomium, with slender, tapering

tentacular cirri extending to frontal margin ofprostomium. Maxillary formula unknown. !

i

First setiger twice as long as following ones, with broad, muscular parapodial bases; parapodia

elongate, pointing forward, anterior side facing body; with large, auricular presetal lobe, subulate postsetal i

lobe and long, tapering dorsal and ventral cirri (Fig. 9.7B). Second setiger slightly longer than following i

ones, parapodia somewhat prolonged and pointing forward, with rounded presetal lobe; postsetal lobe and II

cirri resembling those of first parapodium (Fig. 9.7C). Presetal lobes of subsequent setigers short, conical; i

postsetal lobes subulate to about setiger 15,then low and rounded; ventral cirri in setigers 3 and 4 spherical, i

replaced by pads after that; dorsal cirri long and tapering to setiger 15, then shortening and narrowing.

Branchiae starting on setiger 8, smaller than dorsal cirri at first, after setiger 15 surpassing them in width and

length, broadly foliaceous (Fig. 9.7D).

244 !
I

I

I

' /

I E F / /

G, HJ

I _-'_>_ I

I '- '.'.''.":"-..'.-..... preL

II i

- . ......:.::.:.s:..:_,_., !/_

i i LIM

I i:;_i ...... ac

• •_ • If "_

_C'-__ -------- ":__;_ '

i L _T ...... ::_: o__

!

Figure 9.7. Nothria occidentalis: A, anterior end, dorsolateral view; B, parapodium 1, anterior view; C,

parapodium 2, anterior view; D, middle parapodium, anterior view; E-F, simple and pseudocompound

I hook, setiger 1; G-H, uni- and bidentate pseudocompound hooks, setiger 2; I, nearly compound

hook, setiger 3; J, Pectinate seta; K-M, anterior, middle, and posterior simple setae.

I

I _4_
!

|

Unidentate spines with hooked tips and short, oblique hoods in setiger 1(Fig. 9.7E, F); pseudocompound •

to compound uni- and bidentate hooks with long, spinose hoods in setigers 2 and 3 (Fig. 9.7G-I); scoopshaped pectinate setae with 10-15 long, equal teeth from setiger 2, often forming dense fascicles (Fig. 9.7J); •

limbate setae present from setiger 2, geniculate, with broad wings; in posterior setigers geniculate part becoming

shorter in relation to shaft (Fig. 9.7K-M). Subacicular hooks long and slender, with relatively small heads

and short, oblique hoods; present singly from setiger 9 or 10, in pairs from setiger 13 (Fig. 9.7N). 1

Tubes fiat, with tough inner lining and outer cover of shell hash.

Remarks. The auricular presetal lobes of the first parapodia clearly mark this specimen as aNothria. i

The genus Paranorthia has been recognized as a synonym of Rhamphobrachium by Paxton (1986) and •

represents juveniles of the latter. i

Habitat. Found in sand mixed with shells and mud. !

Distribution. Central California to Colombia, 80 to 200 m. 1

i

Genus Onuphis Audouin and Milne Edwards, 1833 1

Type Species. Onuphis eremita Audouin and Milne Edwards, 1833

I Diagnosis. Ceratophores of occipital antennae distinctly longer than prostomium, usually with 10 to

25 rings; styles of outer (anterior) lateral antennae shorter than their ceratophores. Pseudocompound hooks i

bi- to tridentate, rarely quadridentate, with short, blunt hoods; large median hooks and compound spinigers i

absent. Modified parapodia prolonged, at least in anteriormost setigers. Branchiae simple to pectinate, most

II

often present from setigers 4 to 6, sometimes from sefiger 1, rarely absent. Body size moderate to small, I

length to 300 ram, width to 4 mm. 1

m

Remarks. Onuphis is a typical part of the fauna of the Eastern and Western Pacific oceans. The

genus is easily recognized by the long ceratophores of the occipital antennae. Seven previously described m

species have been reported off California; three of those, and an additional species new to science were found mE

the Santa Mafia Basin. I

Onuphis affinis Itilbig, new species I

i

Figure 9.8 I

Onuphis pallida Lissner et al., 1986:D-11.--Not Moore, 1911.

Onuphis sp. 1SAIC, 1992:A-7 i

Material Examined. California, R/V Point Sur, off the Farallon Islands, north of Pioneer Canyon,

Sta. 3-10, 37°26.00'N, 123°10.83'W, 985 m, 16September, 1991: holotype (CASIZ 099239) and 3 paratypes 1

(USNM 170423); Sta. 3-4, 37°27.83'N, 123°11.60"¢¢, 1040 m, 14 September, 1991:4 paratypes (LACMAHF Poly 1657); Santa Maria Basin, off Point Arguello, Sta. 63, 34°26.29'N, 120°58.08a_V,930 m: 1paratype

(USNM 170422); Sta. 69, 34°22.88'N, 120°54.20'W, 927 m: 1 paratype (SBNMH 142695); additional 1

specimens: off the Farallon Islands, north of Pioneer Canyon (9), axis ofPioneer Canyon (1), south of Pioneer

Canyon (10).

Description. Holotype complete, 100 mm long, 2 mm wide excluding parapodia (measured behind 1

pharyngeal region) for 247 setigers; longest incomplete specimen 135 mm long, 2 mm wide for about 225 1

setigers; other specimens 20 to 91 mm long, 1 to 3 mm wide for 38 to 176 setigers. Body slender, flattened,

rigid, with crowded segments behind pharynx, tapering toward pygidinm within last 20 setigers or so. Color I

246 !
!

in alcohol pale or dusky because of very f'me transverse dark brown lines running across dorsal and ventral

surface; cuticle highly iridescent.

I Prostomium small, bluntly triangular, bearing oval to clavate frontal antennae about as long as prostomium; ceratophores of occipital antennae crowded, arranged in semicircle; eyes not observed; nnchal

organs in pigmented specimens delineated by blackish lines along posterior prostomial margin separated by

I small with 10-15 mediantings, gap.distinctly Outer (anterior) longer occipital than ceratostyles; antennae asinner long(posterior) as peristomium lateralaantennae nd first 1-2assetigers, long as peristomium ceratophores

and first 13-18 setigers (8-9 in small individuals), ceratophores with 12-16 tings, always longer than those of

I outer individuals), lateral antennae; ceratophoresmedian with 6-10 antenna rings, as usually long asabout peristomium half as long and asfirstthose6-9ofsetigers inner lateral (rarelyantennae. 14; 4 in small Palps

massive, about as long and twice as wide as frontal antennae. Peristomium about half as long as first setiger,

i bearing two Maxillae very slender weakly and long tentacular cirri extending to anterior prostomial margin (Fig. 9.8A). scterotized, but calcified and hard, with darkened outer margins and teeth; carriers

short, triangular, laterally incised, less than half as long as MI; MII 7-9+8-10, MIII 9+0, MIV 7-8+11,

I arching over first 3 maxillae, very hard and massive; MV small, delicate, triangular plates without dentition (Fig. 9.8B). Mandibles about as long as maxillae, with slender, sclerotized shafts beating calcified, deeply

incised and irregularly serrated cutting plate (Fig. 9.8C).

I First two setigers about as long as wide, bearing modified parapodia with long dorsal and ventral cirri and postsetal lobes; parapodia I-4 slightly displaced ventrally and pointing forward (Fig. 9.8D, E). Subsequent

setigers quickly becoming wider and shorter, only as long as width of parapodial stem throughout. Ventral

I cirri cirriform through setiger 5 or 6, sometimes followed by transitory cirrus, replaced by pad in setigers 6- 8. Postsetal lobes elongate conical through setiger 11-13 (9-10 in small individuals), gradually turning

papilliform in middle and posterior setigers (Fig. 9.8F, G). Dorsal cirri present throughout, very long and

prominent in modified setigers, gradually diminishing and very short and slender through most of body. Gills simple, first present in setiger 4 (rarely 3 or 5), as long as corresponding dorsal cirri at first, quickly surpassing

dorsal cirri and reaching at least half body width; absent from last 20 or 30 setigers.

Trident,ate hooks in 1-4, stout and short-bladed or slender and longpseudocompound present setigers

bladed, with slightly pointed hoods (Fig. 9.8H); subacicular hooks present from setiger 12, (occasionally 10

or 11in small specimens, rarely 13), with short, squarish hood (Fig. 9.81); simple limbate capillaries with very

I narrow wings present throughout, pectinate setae present in most setigers, bearing about 10-12 equal teeth (Fig. 9.8J, K).

Pygidium very small, bearing 2 pairs of ventrally inserted anal cirri, upper pair longer than lower

I pair.

Remarks. The new species shows similarities with several co-occurring species that seem to form a

I clu "intermediates" ster of closelydescribed related forms by Hobson similar(1971). to thaOt

.incl affinis udingresembles O. elegans O. pallida , O. iridandescens O. similis , O. oinpaltheinanumber and theof

setigers with pseudocompound hooks, but is most similar in the occurrence of subacicular hooks to O.

I geo differphiliniformis the first , aoccurrence species withof the onlysubacicular 3 segmentshooks beating(setiger pseudocom 16 for Opound . pallidasetae. , 11-13O.

foraffiOnis. affinis and )Oand . pallmost ida

likely also the number of maxillary teeth, especially MII (6+9 for O. pallida, 7-9+8-10 for O. affinis) and

i MIV (6-7+8 for O. pallida, 7-8+11 for O. affinis). O. affinis differs from O. similis in those same two characters (subacicular hooks present from setiger 8-9, MII 5+7, MIV 5+8) and in addition in the first

occurrence of the branchiae (nearly always setiger 4 for O. affinis, setiger 6 for O. similis).

I Etymology. The species name is the Latin word for close or related and refers to the morphological resemblance to several related species.

Distribution. Central California, 800 - 2755 m, in silt and sand mixed with silt.

I
Figure 9.8. Onuphis affinis Hilbig, new species: A, anterior end, dorsal view; B, maxillae; C, mandibles; D- 1

E, parapodium 1 and 4, anterior view; F-G, middle and posterior parapodium, anterior view; H,

pseudocompound hook; I, subacicular hook; J, simple seta; K, pectinate set.a. (all from non-type

specimens). I
!

I Onuphis elegans (Johnson, 1901)

Figure 9.9

!

Northia eIegans Johnson, 1901:406-407, pl. 8, figs. 77-85.

Nothria elegans: Hartman, 1968:675 (in part).--Blake, 1975a:201, figs. 172-175; 1975b:43-61.

I Hobson, 1971:529-531, 1.--Fauchald, 1982:45. Onuphis elegans: fig.

I Lagoon, Macoll.teri11al June, Examine 1952d.(2).California: Dillon Beach, Lawson's Flat, coll. 13 July, 1973 (i); Bodega

Description. Length to 160 mm, width to 6 ram, segments to about 250. Body cylindrical through

I In-st4 or 5 setigers, then dorsoventrally flattened and linear, tapering gently toward pygidium within last 25 setigers. Color in alcohol pale or, in anterior setigers, with brown dorsolateral spots located at posterior

margin of each segment, sometimes forming a transverse line.

I Prostomium small, rounded, almost concealed by crowded bases of occipital antennae, bearing 2

pairs of eyes in front of and beside insertion of inner (posterior) lateral antennae. Frontal antennae ovoid,

almost as long as prostomium; ceratophores of outer (anterior) lateral antennae much longer than styles, with

I 10 rings, antennae as long as peristomium and first setiger; ceratophores of inner (posterior) lateral antennae with 11 rings, antennae as long as peristomium and first 7 setigers (range 3-11); ceratophore of median

antenna shortest, with 5 rings, antenna as long as peristomium and f'trst 2 setigers (Fig. 9.9A).

I Maxillary formula, according to Fauchald (1982): MI 1+1, MII 6-7+5-7, Mill 7-10+0, MIV 6-7+5-

8, MV 1+1.

I displacedFirst ventrally, 4 or 5gradually parapodiainserting modified, morewithdorsallly long cirri and and pointing postsetal to thelobes, side. Gills directedpresent forward from and firstslightly setiger,

simple and straplike throughout, between setigers 10 and 60-80 markedly wider and thicker than preceding

i and subsequent gills, typically appearing to be winged on one side (Fig. 9.9B). Ventral cirri replaced by pads from setiger 6, rarely 7; postsetal lobes cirriform through setiger 12 or 13. Slender and stout pseudocompound

hooks present in first 4 setigers, usually tridentate (Fig. 9.9C), rarely bi- or quadridentate; subacicular hooks

first present in setiger 8 or 9 (Fig. 9.9D); simple limbate and pectinate setae present throughout, the latter

I distally oblique, with about 14 teeth.

Remarks. The description ofO. elegans is added here although the main distribution of this species

I seems station.toMost be Puget of theSoindividuals und arid further fromnorth California to Canidentified ada and itsasdepth O. elegans range isbased shallower on thethan first anoccurrence y Phase I aofndthe11

subacicular hooks seem to actually belong to the "intermediates" described by Hobson (1971 ), which in turn

I do not occur in Washington and Canada. If the anterior branchiae in an individual do not match Johnson's

(1901) very accurate illustration, but subacicular hooks are present in setiger 9 (rather than 10-12 as in O.

iridescens), it should be assigned to the intermediates. Specimens of less than 0.6 mm body width should be

i identified Hobson, to species with caution because the fast occurrence of subacicular hooks is growth-dependent (see 1971); in adults of greater body width, however, this character seems to be sufficiently consistent for

species discrimination.

I (LACM-AHF Type lPoly ocation740andandtype MCZsp1895). ecimens. Washington, Puget Sound, Port Townsend region: 2 syntypes

Biology. Larvae were reported to be most abundant between January and April, and postlarval

I juveniles were found between February and October in California waters (Blake, 1975b). A 60-setiger stage, the smallest adult size, is reached within 120 days of fertilization. O. etegans seems to prefer sandy sediments

and can be hand collected on beaches.

Distribution. British Columbia to southern California, intertidal to 23 m.

i 249
I

I

I

iLan

I

oLan 1

' '_ tC

A b_ I

!

! ',

D !

C

!

Figure 9.9. Onuphis elegans: A, anterior end, dorsal view; B, middle parapodium with wide gill; C,

pseudocompound hooks, setiger 2; D, subacicular hook. (all from Blake, 1975a). 1

250 I
!

!

I Onuphis geophih'formis (Moore, 1903)

Figure 9.10

I Northia geophiliformis Moore, 1903:445-448, pl. XXV, figs. 57-59.

Nothria geophiliformis: Fauchald, 1968:22.--Hartman, 1968:677.

i Onuphis7a-j. geophiliformis: Fauchald, 1982:47-48, fig. 13a, tab. 14.--Maekawa and Hayashi, 1989:72-74, fig.

I Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. R2 (7).

Description. Length to 10 mm, width to 1mm excluding parapodia, segments more than 100. Body

I slender, gracile, flattened except for pharyngeal region, tapering toward pygidium within last 10 segments or so; pigmentation consisting of wide brown bands across dorsum in anterior segments, alternating with pale

intersegmental furrows; cuticle vividly iridescent.

I

slender; Prostomium occipital antennae small, with bluntlylongtriangular, ceratophores with divided crowdedintoantennal 10 to bases. 15 tings,Frontal those antennae of outer lateral oval, relatively antennae

distinctly longer than ceratostyles. Outer (anterior) lateral antennae as long as peristomium and first 1-4

i setigers, inner (posterior) lateral antennae as long as peristomium and first 6-11 setigers, median antenna as long as peristomium and first 5-8 setigers. Eyes usually absent (Fig. 9.10A).

Maxillae with triangular, laterally slightly incised carriers about half as long as MI; MII 7+6, Mill 7-

8+0, MIV 8+9, MV 1+1 (Fig. 9.10B). First 4 parapodia elongate, displaced ventrally and pointing forward, subsequent parapodia inserted

laterally and pointing to the sides, dorsal cirri, gills and postsetal lobes arched across dorsum. Postsetal lobes

I elongate triangular to setiger 10-12, ventral cirri padlike from setiger 5 or 6; gills first present on setiger 3-6, simple and straplike throughout, much larger than dorsal cirri, absent from last 20 to 30 setigers (Fig. 9.10CE). Pseudocompound hooded hooks present in setigers 1-3, tridentate, with pointed hoods (Fig. 9.10F);

i subacicular hooks first present on setiger 10-12 (Fig. 9.lOG); simple limbate capillaries present throughout, pectinate setae ("paddle-shaped setae" in Moore's original description) with 10-15 subequal teeth, one outer

tooth longer than others, present after setiger 10 (Fig. 9.1 OH).

I Pygidium small, bearing 2 pairs of ventrally inserted anal cirri, outer pair twice as long as inner pair;

anus dorsal (Fig. 9.101).

Remarks. The species is easily recognized by its gills that are simple and start on setigers 3-6 rather

I than 1 it differs from O. similis the number of (Fauchald, 1968), a

species occurring offSaja California, by

setigers with pseudocompound hooks (3 for O. geophiliformis, 4 for 0. similis).

Type Locality and Type Specimens. Japan, north of Sendai Bay: holotype (USNM 15711), 7

I paratypes (USNM 5362), 11 paratypes (ANSP 971).

Habitat. Found in mud and sand; the examined specimens were collected from sediments with more

I than 80%Distribution. silt. Japan; Alaska to southern California, 100 - 295 m.

!

!

|
Figure 9.10. Onuphis geophiliformis: A, anterior end, dorsal view; B, maxillae; C, first parapodium, anterior •

view; D, fourth parapodium, anterior view; E, middle parapodium, anterior view; F, pseudocompound l

hooks; G, subacicular hook; H, pectinate seta; I, posterior end, dorsal view. (A-H after Maekawa

and Hayashi, 1989). ..

I

!
!

!

Onuphis iridescens (Johnson, 1901)

Figure 9.11

I Northia iridescens Johnson, 1901:408, pl. 8, figs. 86-87, pl. 9, figs. 88-92.

Nothria iridescens: Hartman, 1944:87-88, pl. 5, figs. 99-104 (in part); 1968:681.

I Onuphis 1986:D-11 iridescens: Hobson, 1971:533-535, figs. 3a-d, 4a-c.--Fauchald, 1982:49, fig. 14a.--Lissner et al., (in part).---Steinhauer and Imamura, 1990:F-1.

I Material Examined. California: Santa Mafia Basin, off Point San Luis, Sta. 23 (1), Sta. R-3 (2).

Description. Length more than 30 mm, width to 3 mm excluding parapodia, segments more than 50.

I Body transverse linear, barstout, acrossflattened posteriorbehind part ofpharyngeal each segment region; (Fig.color 9.11A); in alcohol cuticle highly pale, anterior iridescent. segments brownish

Prostomium rounded, wider than long, nearly concealed by bases of occipital antennae; outer (anterior)

I lateral with antennae as long as peristomium and frost 2 or 3 setigers, ceratophores distinctly longer than styles, 8-14 rings; inner (posterior) lateral antennae as long as peristomium and first 14 setigers (range 7-25),

ceratophores with 9-14 rings; median antenna as long as peristomium and first 10 setigers, ceratophore with

I 9Peristomium rings, only half as long as those of inner lateral antennae. Frontal antennae ovate, shorter than prostomium. shorter than first setiger, beating relatively short tentacular cirri projecting about as far as bases

of inner lateral antennae.

i Maxillae with slender, together urn-shaped carriers and slender, gently curved first maxillae. Maxillary formula: MI 1+1, MR 7+7-8, MIII 7-8+0, MIV 5+10, arching over MI-MIII; MV squafish, delicate plates

with no teeth (Fig. 9.11B). Mandibles about as long as maxillary apparatus, with slender shafts bearing

I deeply incised, scalloped, calcified cutting plate (Fig. 9.11C).

First 3 or 4 parapodia enlarged, with long cirri and postsetal lobes, slightly displaced ventrally,

pointing forward (Fig. 9.11D); subsequent parapodia gradually moving into lateral position and pointing to

I the sides. Venlxal cirri replaced by pads in setiger 6 (rarely 7 or 8; 4 or 5 in juveniles), postsetal lobes cirriform through setiger 12-16 (8 in juveniles). Gills simple and slender throughout, present from setiger 1;

smaller than corresponding dorsal cirri at first, but quickly surpassing the latter after the first 3 or 4 setigers

I (Fig. 9.11E, F); arched over dorsum, reaching dorsomedian line where best developed. Slender and stout tridentate pseudocompounds present in setigers 1-4 (Fig. 9.1 tG, H); hooded subacicular hooks present from

setiger 12 or 13 (8-10 in small individuals) (Fig. 9.11I); limbate setae with very narrow wings in all setigers

I (Fig. 9.11J); pectinate setae in most sefigers, distally slightly oblique with about 15 teeth.

Tube parchment-like, white, with outer layer of sand grains.

i Remarks. Onuphis iridescens is very similar to two co-occurring forms, O. elegans and the Onuphis intermediates of Hobson (1971 ). As the characters that separate those forms are growth dependent and seem

to stabilize when the individuals are about 60 setigers long and about 0.6 mm wide, specimens smaller than

i that shoulTdypenotLbeocaliden itytifianded Typ to spe e Spcies. ecimens. British Columbia, Victoria: holotype (MCZ 1887).

Habitat. The examined specimens were found in silty mud, but generally occur in various types of

I mixed sediments. Distribution. British Columbia to Mexico; Sea of Okhotsk; South Atlantic; intertidal to 2400 m.

!

I 253
I

I

!

I

° i

I

I

I

',

i

° i

I

Figure 9.11. Onuphis iridescens: A, anterior end, dorsal view; dashes: pigment pattern; B, maxillae; C, mandible;

D-E, parapodia 2 and 15, anterior view; F, middle parapodium, anterior view; G-H, stout and I

slender pseudoeompound setae; I, subacicular hook; J, simple seta.

I

254 I
!

!

!

Onuphis sp. "intermediates"

I Figure 9.12

I Onuphis Onuphis sp. elegans "intermediates" : Lissner et alHobson, ., I986:D-11. 1971:535-537.

Onuphis iridescens: Hartman, 1944:87-88, pl. 5, figs. 99-104 (in part).---_ssner et al., 1986:D-11 (in part).

I

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. 21 (3). Tomales Bay,

PMS 180240, 180242 (2).

I Description. Length more than 92 mm, width to 1.5 mm, segments more than 260. Body slender,

linear, flattened after first few setigers, in posterior region becoming softer andalmost cylindrical; color in

alcohol pale, occasionally anterior segments with transverse dorsal stripes of brownpigment across posterior

I half; cuticle strikingly iridescent.

Prostomium bluntly triangular, about as long as wide; frontal antennae ovate, slightly shorter than

I prantennae ostomium; as tonglateral as peristomium occipital antennae and firstwith2 orvery 3 setigers, long cerceratophores atophores an2-3d shtimes ort styles; as longouasterstyles, (anteriwith or) late 13-15ral

rings; inner (posterior) lateral antennae as long as peristomium and first 7 or 8 setigers, ceratophores as long

I assetigers, entire oceratophore uter lateral antennae, short and wslender, ith abouwith t 16 tings; 8 tings.median Eyes antenna not observed. about asPeristomi long asuperistomi m widest umposteriorly, and first 4

laterally continuous with prostomium, with slender tentacular cirri projecting to middle of prostomium (Fig.

I 9.12A). Maxillary formula MI 1+1, MII 7+7, MIII 8+0, MIV 6+8, MV squarish plates with no dentition.

First 4 parapodia enlarged, with elongate cirri and postsetal lobes, inserted slightly ventrally and

I pointing forward (Fig. 9.12B). Vental cirri replaced by pads in setiger 6 or 7, postsetal lobes cirriform through setiger 8 or 9; gills present from setiger 1, slender and thin throughout, exceeding dorsal cirri after

first few setigers. Pseudocompound hooks present in setigers 1-4, Iridentate, sometimes with rudimentary (or

I broken?) terminal tooth, in anteriormost setigers of 3 or 4 kinds, stout to very slender with large to delicate teeth (Fig. 9.12C-F), of 2 kinds in subsequent setigers. Subacicular hooks present from setiger 8 or 9, dark

amber colored, projecting further in more posterior setigers than in anterior ones (Fig. 9.12G). Simple limbate

[ I setae present throughout, with very narrow wings; pectinate setae in most setigers, distally oblique, with

i • about 15 teeth (Fig. 9.12H). Dorsal cirri with fascicles of 3 delicate notoaciculae, neuropodia with 2-3

neuroaciculae with geniculate free tips.

I Remarks. Hobson (1971) hypothesized that these intermediates with characters reminiscent of O.

elegans (start of subacicular hooks) and O. iridescens (slender shape of branchiae) may be offspring of

interbreeding individuals of these two species. Although this hypothesis has notbeen proven through experiments

I in the laboratory, it is favoured here over the possibility of a new species (also suggested by Hobson). It

appears likely that the two established species and the intermediates are still in the process of speciation, and

they may be isolated more strongly by biological characters, such as depth range or reproductive behaviour,

I than their respective genetic makeups.

!

I 255
!

!

!

!

!

' - ' I

• i

H G I

I

I

, !

B C /

F

:J_ . "•. D E I

" |

i

Figure 9.12. Onuphis sp. "intermediates", specimen indentified and dissected by Hobson: A, anterior end, dorsal

G,vieW;su B, parapodium bacicular hook; 2H,Orpectinate 3, anteriorseta.View; C-F, pseudocompound hooks from same, dosal to ventral; i

i

Genus Paradiopatra Ehlers, 1887 ii

Type Species. Paradiopatrafragosa Ehlers, 1887

i

Diagnosis. Ceratophores of occipital antennae short, no longer than prostornium; styles of outer i

(anterior) lateral antennae distinctly longer than their ceratophores. Pseudocompound hooks uni- to tridentate,

most often bidentate, with long, slender, acutely pointed hoods; large median hooks and compound spinigers •
!

absent. Modified parapodia not prolonged. Branchiae pectinate, often restricted to short anterior part of

body between setigers 2-18 and 30-50, rarely absent. Body size generally small, length to 100 ram, width to

I Remarks. The genus Paradiopatra was revised by Paxton (1986) and now includes all species

2 iiun.

formerly assigned to Sarsonuphis Fauchald, 1982, a genus recognized asjunior synonym of Paradiopatra by

I parapodia, Paxton (1986a). readilyThdistinguish e pointed hoods Paradiopatra of the pseudocompound from Kinbergonuphis hooks,andoftenMooreonuphis visible without whodissection also haveofshort the

ceratophores. The distribution of Paradiopatra is worldwide, but most species are typical deep-water forms.

I Only two species are known from California, one of which was present in the Phase I and II material.

i Paradiopatraparva (Moore, 1911) Figure 9.13

I Onuphisparva Moore, 1911:263-266, pl. 17, figs. 51-57, pl. 18, figs. 98-99.--Fauchald, 1968:37, pl. 9, figs.

a-e.

i Sarsonuphis parva: Fauchald, 1982:76-77, fig. 20f.--Lissner et al., 1986: D- 11.--Steinhauer and Imamura, 1990:F-1.

Paradiopatra parva: Paxton, 1986a:38.

I Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. R-2 (2); off Purisima

Point, Sta. 48 (1); off Point Arguello, Sta. 65 (1).

I Description. Length to 45 mm, to 1 nun, segments to Body slender, gracile, linear,

width about 100.

only slightly flattened, tapering toward pygidium within last 5-8 setigers; color in alcohol pale.

i Prostomium rounded, about as long as wide, with clusters of 3-6 small lateral eyespots just outside

inner lateral and behind outer lateral antennae; 2 additional single eyes (rarely 3) on frontal margin. Ceratophores

of occipital antennae short, with 4-5 rings, arising from crescentic line across middle of prostomium (Fig.

i 9.13A). Outer (anterior) lateral antennae as long as peristomium and first setiger, styles much longer than ceratophores; inner (posterior) lateral antennae as long as petistomium and first 4-7 setigers; median antenna

as long as peristomium and first 5 setigers. Peristomium shorter than prostomium, about as long as subsequent

setigers; tentacular cirri slender, extending to antennal bases.

I Jaws delicate, transparent; maxillary carriers together urn-shaped, shorter than MI; MII 8-9+9, MIII

7+0, MIV 5-6+7-8, arched over MI; MV small rounded plates without teeth (Fig. 9.13B). Mandibles with

I slender shafts All and poorly mineralized cutting plate. setigers of similar width and length except for last 6-8 prepygidial ones; first 2-3 parapodia

slightly modified, with somewhat elongate cirri and postsetal lobes (Fig. 9.13C); ventral cirri replaced by

pads setigerin3setiger (rarely42(orrarely 4), simple 3 or 5),atpostsetal first, thenlobes quicklycirriform becomingthrough pectinate setigerwith9-1up2;tobr8anchiae filaments, firstarched presentover on

dorsum, about one-fourth as long as segmental width, longer than dorsal cirri (Fig. 9.13D); last branchia on

I setiger very long37-45. and pointed, Pseudocompoun present din setigers hooks bidentate, 1-3 (Fig. sometimes 9.13E); subacicular unidentatehooks or wfiithrstinypresent subdistaln setiger tooth,9,hooshort ds

and very stout at first, gradually projecting further out and becoming more slender with teeth pointing almost

straight throughout, up raleatherst than numerous formingin posterior tight angleparapodia; with shaftpecti (Fig.nate9.13F, setaeG);present slender,fromnarrowanterior ly limbate setigers, setaewidepresent with

about 15 short teeth anteriorly, gradually narrowing, with 10 long teeth in far posterior segments (Fig. 9.13H, I).

!

257
I

I

I

l '

!

:/ ,ll '

'

_ -_-, _i:__,,,_ _.,,,,_-,,. _ I

• x 1/

I

D !

Figure 9.13. Paradiopatra parva: A, anterior end, dorsal view; B, maxillae; C-D, parapodia 1 and 7, anterior !

views (after Fauchald, 1968); E, pseudocompound seta (after Fauchald, 1968); F-G, subacicular

hooks, posterioranterior end, dorsal and farview. posterior setiger; H-I, peetinate setae, anterior and far posterior setiger; J, I

258 |
I

I Pygidium with dorsal anus and 2 pairs of anal cirri, shorter pair as long as last 6 setigers, longer pair

as long as last 8 setigers (Fig. 9.13J).

I only otherRemarks known. species Paradiowi patra th thisparva character is easiisly Pr.ecognized furcasetosaby(Monro, the presen1937) ce ofwhich gills before occurssetiger in the 1Indian 0; the

Ocean. The co-occurring P. hiadentata (Moore, 1911) differs most conspicuously from P. parva in the

I branchiae; they are simple rather than pectinate and start on setiger 12 rather than 3. Type Locality and Type Specimens. California, Monterey Bay, off Point Pifios Lighthouse: holotype

(USNM 17356) and 9 paratypes (USNM 17359).

I Biology. (1911) reported species can great abundances, mostly mud,

Moore that this be found in in

but also in sandy sediments. The examined specimens were collected in sediments with about 80% silt.

i Distribution. Southern and central California; Baja California; 10 - 300 m.

Genus Rhamphobrachium Ehlers, 1887

I Type Species. Rhamphobrachium agassizi Ehlers, 1887

Diagnosis. First three setigers highly modified, bearing elongate parapodia with very long, delicate setae with spiny shafts and hooked tips, setal sacs extending back to setigers 30-60. Ceratophores of occipital

antennae short, with 2-5 rings; all styles short, no longer than peristomium and first 2-5 setigers. Branchiae

I from setiger 6-17, simple or pectinate. Body size moderate, length to more than 190 mm, width to 7 mm. Distribution worldwide, all species rare.

Remarks. Rhamphobrachium and related genera were revised by Paxton (1986b), who erected the

I subgenera R. (Rhamphobrachium) and R. (Spinigerium), the latter differing from the former by the presence

of compound spinigers in some anterior segments and a sixth pair of maxillae.

I Rhamphobrachium longisetosum Berkeley and Berkeley, 1938

i Figure 9.14

Rhamphobrachium longisetosum Berkeley and Berkeley, 1938:428-435, figs. l-8.--Hartman, 1944:48-49,

I pl Imamura, . 1, figs.1990:F-1. 1-8.--Paxton, 1986:88-89, figs. 7a-g.--Lissner et aI., 1986:D-11.--Steinhauer and

I Material Examined. California: Santa Mafia Basin, off Point Estero, Sta. 2 (1); off Point Sal, Sta.

R-8 (1).

• flattenedDescription , with highly. mLeodngthifiedmoanretethrior ans84etigmmers,;width color toin5almmcohol,seuniformly tigers morepale than, or12p7ros. Bod tomium y lin,eaper,ristomi somewhumat

andf'n-stfew setigers with brownish dorsal pigment bands.

Prostomium small, rounded, with short appendages; frontal antennae spherical; ceratophores of occipital antennae shorter than prostomium, with 4 short (range 3-6) tings and longer terminal ring; outer (anterior)

lateral antennae as long as peristomium and first 1 or 2 setigers, inner (posterior) lateral antennae as long as

1 peristomium and first 3-5 setigers, median antenna as long as peristomium and first 4 setigers. Two pairs of

eyes, usually faded in alcohol. Peristomium short, bearing fairly long, tapered tentacular cirri, almost reaching

palps (Fig. 9.14A).

I

i 259
Figure 9,14, Rhamphobrachium longisetosum: A, anterior end, dorsal view; B, maxillae; C, mandibles; D, ll

parapodium 4, anterior view; E, far posterior parapodium, anterior view; F, recurred hook; G-I,

pseudocompound setae; J-K, pectinate setae; L, subacicular hook. (A, F after Berkeley and Berkeley,

1938; B, C after Hartman, 1944; D, G-I after Paxton, 1986b). •

260 I
!

I Mandibles calcified, with long, slender shafts beating crescentic cutting plate with large tooth on

either side of median cleft. Maxillae hard and calcified, with large, squarish carriers and wide, strongly

I falcate MI. M116-7+7, MIII 7-8+0, MIV 7-8+9-11, MV flat plates with no teeth (Fig. 9.14B, C). First 3 parapodia strongly modified, progressively longer from anterior to posterior, pointing forward,

with anterior sides facing body wall, with long subulate postsetal lobes and 3 short, papilliform lobes surrounding

I each size after seta;setiger dorsal and 5; ventral ventralcirri cirrisubulate subulate.through Postsetal setiger lobes3,present sphericalthrough in setiger setigers 4 (Fig. 15-20, 9.14D), muchreplaced reduced byin

pads after that. Branchiae from setiger 8 or 9, pectinate, with maximally 8-10 short filaments (Fig. 9.14E),

I simpl two longitudinal e in first fewrowssetigers. of spines, Modisetal fiedsaparapodia cs extendingeachtowith about3setiger very lo50ng,(Fig. slend9.14F). er recurved Pseudocompound hooks, shaftssetae with

present in setigers 4 to about 15, in lower part of setal fascicles (Fig. 9.14G-I); simple limbate setae and

I pectinate hooks present setae with singly12-20 fromteeth setiger present 12-14, fromin setiger pars from4, insetigers upper part15-17 of setal (Fig.fascicles 9.14L).(Fig. 9.14J, K); subacicular

Tubes with tough, parchment-like inner layer and sandy outer layer.

I Remarks. Rhamphobrachium longisetosum is the only species of this known to occur in the

genus

northeastern Pacific; the strongly modified first three segments bearing prolonged parapodia make this species

unmistakeable even if the rather fragile hooks have broken off. The specimen from Point Estero appears to be

I the largest one reported and may also represent a range extension.

Type Locality and Type Specimens. California, off mouth of Santa Ana River: syntype (USNM

32865); off Balboa: syntype (USNM 32866).

I Habitat. Found in sandy sediments; the examined specimens were collected at stations with about

75% sand.

I Distribution. Central California to Mexico; Galapagos Islands; 20 - 730 m.

I Literature Cited

i Bhaud, M. current and C.biological Cazaux. problems. 1987. Descripti Oceanison 13(6): and identificati 596-753.on of polychaete larvae; their implications in

Blake, J.A. 1975a. Phylum Annelida: Class Polychaeta. Pp. 151-243 In: R.I. Smith and T.J. Carlton (eds.),

I Light's Manual: Intertidal Invertebrates of the Central California Coast. University of California Press, Berkeley.

m Blake, J.A. 1975b. The larval development of Polychaeta from the northern California coast. II. Nothria

[I elegans (Family Onuphidae). Ophelia 13: 43-61.

Fauchald, K. 1968. Onuphidae (Polychaeta) from western Mexico. Allan Hancock Monographs in Marine

I Biology 3: 1-82.

Fauchald, K. 1982a. Revision ofOnuphis, Nothria, andParadiopatra (Polychaeta: Onuphidae) based upon

i type material. Smithsonian Contributions to Zoology 356:1-109. Fauchald, K. 1982b. Description ofMooreonuphisjonesi, a new species ofonuphid polychaete from shallow

water in Bermuda, with comments on variability andpopulation ecology. Proceedings of the Biological

Society of Washington 95(4): 807-825.

Fauchald, K. and P.A. Jumars. 1979. The diet of worms: a study of polychaete feeding guilds. Oceanography

i and Marine Biology Annual Review 17:193-284.

I 261
!

Gathof, J. 1984. Family Onuphidae Kinberg, 1865. Chapter 39 In: Uebelacker, J.M. and RG. Johnson

(eds.). 1984. Taxonomic Guide to the Polychaetes of the Northern Guff of Mexico. Barry A. Vittor •

& Associates, Inc., Mobile, Alabama. 7 vols. |

Hartman, O. 1944. Polychaetous annelids. Part V.Eunicea. Allan Hancock Pacific Expeditions 10: 1-237,

pl. 1-18. I

Hartman, O. 1967. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises, chiefly

from antarctic seas. Allan Hancock Monographs in Marine Biology 2: 1-387. i

[] Hartman, O. 1968. Arias of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation,

University of Southern California, Los Angeles. 828 pp. m

Hobson, K.D. 1971. Some polychaetes of the superfamily Eunicea from the north Pacific and north Atlantic H

Oceans. Proceedings of the Biological Society of Washington 83(47): 527-544.

Hsieh, H.-L. and J.L. Simon. 1987. Larval development of Kinbergonuphis simoni, with a summary of i

development patterns in the family Onuphidae (Polychaeta). Bulletin of the Biological Society of i

Washington 7: 194-210. i

Johnson, H.P. The Polychaeta of the Puget Sound region. Proceedings of the Boston Society of Natural H

History 29(18): 381-437, pl. 1-19.

Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bemstein, R. Cimberg, T. Kauwting, and W. Anikouchine. i

1986. Assessment of long-term changes in biological communities of the Santa Maria Basin and i

Western Santa Barbara Channel - Phase I. Report prepared for the U.S. Department of the Interior, i

Minerals Management Service, Pacific OCS Region, under contract No. 14-12-0001-30032. H

Maekawa, N. and I. Hayashi. 1989. Onuphid polychaetes from Wakasa Bay, Sea of Japan. Memoirs of the

College of Agriculture, Kyoto University 134: 61-93. i

I Moore, J.P. 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. Proceedings

of the Academy of Natural Sciences of Philadelphia 1903: 401-490, pl. XXIII-XXVII. I

Moore, J.E 1911. The polychaetous annelids dredged by the U.S.S. "Albatross" off the coast of southern i

California in 1904:111.Euphrosynidze to Goniadidae. Proceedings of the Academy of Natural Sciences

in Philadelphia 1911: 234-318, pl. XV-XXI. I

Orensanz, J.M. 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas. Biology

of the Antarctic Seas XXI. Antarctic Research Series 52: 1-183.

i

Paxton, H. 1986a. Generic revision and relationships of the family Onuphidae (Annelida: Polychaeta). H

Records of the Australian Museum 38: 1-74.

Paxton, H. 1986b. Revision of the Rhamphobrachium complex (Polychaeta: Onuphidae). Records of the i

Australian Museum 38: 75-104. i

Science Applications International Corporation (SAIC). 1992. Detailed physical and biological oceanographic []

studies for an ocean site designation effort under the Marine Protection, Research and Sanctuaries n

Act of 1972 (MPRSA). Report prepared for the U.S. Environmental Protection Agency under EPA

contract No. 68-C8-0062. xvii + 255 pp. and Appendices A-F. •

Shisko, J.F. 1981. Five new polychaetes of the families Eunicidae and Onuphidae, collected in 1975 and

1976 during the Southern California Baseline Project. Proceedings of the Biological Society of

Washington 94(4): 968-983. i

i

Steinhauer, M. and E. Imamura (eds.). 1990. California OCS Phase II Monitoring Program. Year-Three

Annual Report. Prepared for the U.S. Department of the Interior, Minerals Management Service, m

Pacific OCS Region, under contract No. 14-12-0001-30262. |

262 i

i"
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1 10. FAMILYEumcmAE SAVIGNY, 1818

I by

Brigitte t-Iilbig 1

I

I Introduction

The eunicids are among the more spectacular polychaetes because of theirconsiderable size andvivid

I They arenot only one largest polychaete families terms of number of species, but also one

coloration. of the in

of the very few with afossil record which reaches backto the Cretaceous period (fossils are also known from

most other eunlcemorph families, although more scatteredthanfor the eunicids). The largestknown polychaete,

I reported by (1923) as being tong, a species only polychaetes

Fauvel about three meters is of Eunice. The

eaten regularly by people are swarming species caught in regions of the South Pacific (genus Palola). The

family is important ecologically because they break down coral rock. Eunicids are most closely related to the

I Onuphidae, but can be distinguished from the latter by the of five antennae (sometimes only one or

presence

three) arising from very short, cylindrical ceratophores; in contrast, onuphids always have seven prostomial

appendages, including five antennae that arise from long, usually annulated ceratophores, and two frontal

I appendages referred to as frontal antennae or frontal palps. Eunicids are not quite as iridescent as onuphids, but often more colorful, and typically have larger, more delicate gills than onuphids. In addition, eunicids

build fragile, parchmentlike tubes if any, whereas many onuphids are known for their hard, quill-like or very

I tough fibrous tubes.

I Morphology

i The prostomium is typically bilobed with deep median dorsal and ventral folds, but it may also be rounded. In most genera, it bears five occipital antennae: a pair of outer and inner lateral antennae (also

referred to as A-I and A-II) and a single median antenna (also called A-ffI). There are only two exceptions to

i this pattern; Nematonereis has only one antenna, Lysidice has three antennae. The ceratophores are inconspicuous and short; the ceratostyles may be as long as about 10 to 15 setigers, and they may be smooth,

wrinkled, or articulated, with the articles being cylindrical, moniliform, or drop-shaped. Characters of the

antennae, such as their relative lengths, are often important for species discrimination. Many species have

I two or four eyes. The pharynx is developed into an eversibte proboscis with very massive muscle layers and

a jaw apparatus consisting of ventral mandibles and several pairs of maxillary plates as is typical for the order

Eunicida. The maxillae are asymmetrical; the left side consists of five to six plates, the right side only of four

I five third maxilla is absent the All elements less sclerotized and often

to (the on fight side). jaw are more or

calcified with aragonite, the same mineral that is found in onuphid jaws.

i The peristomium consists of two apodous and asetigerous rings, the anterior one usually much longer than the posterior, which may or may not bear a pair of tentacular cirri. The presence of tentacular cirri is

used as a diagnostic character at the generic level; proportions and the presence of articulations or wrinkles

i can sometimes be used for identification to species.

i 1ENSRConsultingandEngineering,89WaterStreet,WoodsHole,Massachusetts02543

I 263
!

Parapodia are subbiramous with the notopodia reduced to the base of the dorsal cirri; a fascicle of I

fine notoaciculae extends into the cirri that may be smooth, wrinkled, or articulated. The articles are typically

cylindrical and separated by very thin folds, so that in preserved specimens they are easily overlooked. The •

neuropodia are well developed and bear large setal fascicles. Setae include simple capillaries, furcate setae, n

hooded compound spinigers (Marphysa and Euniphysa) and falcigers, and simple, uni- to tridentate, hooded

subacicular hooks (subacicular hooks are absent in the genus Pa/o/a). The color and dentition of the subacicular []

hooks is used to divide the genus into four major groups, at least some of which are natural (Fauchald, pers. i

comm.). Branchiae are often present and typically consist of several branches arising from a more or less

well-defined stem in a palmate or pectinate arrangement. The branchial region may be restricted to a small m

number of anterior segments or encompass most of the length of the animal; the number of branchiae, the U

arrangement and number of filaments, and the distribution of simple and branched gills along the body are

important characters for species discrimination, in

Taxonomic History n

The first eurficids, including the type species of Eunice, E. aphroditois, were described by Linnr,

Pallas, and Miiller in the years between 1767 and 1788 as a species of Nereis, which at that time was a genus !

used to accommodate almost all polychaetes that were not scaleworms. The wealth of eunicid species known

today, however, was described during the second half of the 19th century, during a period of numerous

expeditions by European countries that had colonies in tropical and subtropical locations. In recent years, the n

focus in the literature has shifted from taxonomy to the systematics of this family and its relationship to other

eunicemorphs (Fauchald, 1970, 1992; Orensanz, 1990), and the definition of eunicid genera is still in flux

(Fauchald, pers. comm.). Orensanz (1990) provides an excellent account on the phylogeny of the eunicemorph n

families, including fossil ones. A very extensive account of the history of the genus Eunice can be found in []

Fauchald (1992). Currently, the Eunicidae contain about 275 species in 8 genera, with 205 species alone in

the genus Eunice. U

Distribution and Biological Notes I

Although eunicids are distributed worldwide and in depths ranging from the intertidal to the continental

slope, they are typical tropical, shallow-water organisms, often associated with coral reefs. Some species are i

known to build thin, sometimes branched, parchment-like tubes, includingEunice americana Hartman, 1944, i

a species common in central California. Most species, however, bore into corals and sponges, burrow into the i

sediment, live among algal holdfasts, or dwell in fissures and cracks among rocks. Winsnes (1989) reported i

that some Scandinavian species of Eunice are found inside of cold-water corals where they may secrete athin I

tube. Eunicids are typically very active and thought to be mostly carnivorous (Pettibone, 1982; Fauchald and

Jumars, 1979), feeding on nemerteans (Winsnes, 1989), annelids, small crustaceans andbivalves, chaetogrtaths, i

and diatoms (Gathof, 1984). Some species of Marphysa are believed to be herbivorous or onmivorous. W

Gathof (1984) describes in some detail how reef-dwelling eunicids feed on small organisms attached to coral

rock. The maxillae are used to hold on to the substratum, while the mandibles function as a rasp to break 1

down the coral. Some species leave their tube or shelter to feed, while othersjust extend their anterior end.

When food is abundant, they may store food in the tube (Winsnes, 1989).

!

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I

I When disturbed, eunicids fragment readily and are able to regenerate both ends. Hofmann (1969)

investigated regeneration in Eunice siciliensis under laboratory conditions and found that (1) results were

i better the more anterior the fragment was, and (2) regenerated heads had fully developed sense organs, appendages, and brain including neurosecretory glands, but no mouth and pharynx. Apparently regeneration

of the prostomium ensures completion of one reproductive cycle even though the animals are not able to

I resume feeding. Sexes are separate, and several modes of reproduction are known (_kesson, 1967; Hofmann, 1972,

1974). The most widely known and extensively studied reproductive behavior is epitoky and swarming

I influenced Palola, Eunice by ,lunar and cycles. Lysidice.Swarming Either theduring entire full animals moonornights the shedded has beenposterior describedendsforfilled several withspecies gametesof

ascend to the surface en masse and release eggs and sperm while swarming; since posterior ends are easily

I regenerated, Eunice valensboth(as groups E. kobiensis of species in Lesson, can reproduce 1967), domorenotthan become onceepitokous in their lifetime. and reproduce Some onspecies, the seasuchfloor. as

Bhaud and Cazaux (1987) summarized two studies on larval development of eunicids, one on Eunice harassii

I by Herpin (1925) and the other on Marphysa borradailei by Pillai (1958). The pelagic stage ofE. harassii lasts for about 8 to 10 days; the settling larva has 3 setigers. About 3 days later, the jaws and the median

antenna are visible; inner lateral antennae appear after 8 months when the juvenile has 13 setigers, and outer

i lateral antennae andtentacular cirri appear even later. Species of Marphysa donot swarm, but form gelatinous egg masses. After a very short pelagic period, the larvae settle. A median antenna was observed in M.

borradailei at the ll-setiger stage, and gills first appeared at the 12-setiger stage. As in E. harassii, the

lateral antennae and tentacular cirri form very late, so that specimens with less than 80 setigers cannot even be

I identified to genus.

I Key to the Eunicidae

1A. Peristomium occurring without tentacular cirri; compound setae with spinigerous or falcigerous blades, sometimes together ....................................................................................... genus Marphysa ........ 2

lB. Peristomium with tentacular cirri; compound setae falcigerous, subacicular hooks present ..............

I .................................................................................................................... genus Eunice ............ 3

I • 2A. Subacicular hooks and aciculae yellow; branchiae large, pectinate, up to 7 filaments, present on only

I 10 segments; all compound setae falcigerous ....................................................... Marphysa conferta

2B. Subacicular hooks and aciculae dark brown to black; branchiae simple throughout, present from

I setiger 5-7 to nearly end of body; all compound setae falcigerous .......... Marphysa stylobranchiata 2

3A. Subacicular hooks and aciculae black; MIII short, arching over MI and MII ...................................

I Eunice multipectinata

3B. Subacicular hooks and aciculae yellow; MIII long, located alongside MI and MII ........................ 4

I 4A. Compound setae with pointed hoods; branchiae with up to 23 filaments, on setigers 3 through about

38 ......................................................................................................................... Eunice americana

I 4B. Compound setae with blunt hoods; branchiae with maximally 10 filaments 5

I 2NotfoundinthePhaseIandIImaterialbut , likelytooccuronhardbottominthearea.

i 265
!

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!

5A. Antennae thick, with club-shaped articles; branchial region encompassing less than half of entire

body length; peristomial cirri reaching to middle or posterior margin of prostomium ....................... 1

n

........................................................................................................................... Eunice mutticylindri

5B. Antennae slender, with cylindrical articles; branchial region encompassing more than half of entire •

body length; peristomial cirri reaching tip of prostomium .................................... Eunice vittatopsis I

Description of Species I

Eunicids are typical inhabitants of subtropical to tropical, shallow-water, hard-substratum habitats, _l

and consequently only few members of this family were found during Phases I and II. The following species

are treated in this chapter:

I

Eunice americana Hartman, 1944

Eunice multicylindri Shisko, 1981 •

Eunice multipectinata Moore, 1911

|

Eunice vittatopsis Fauchald, 1970

Marphysa conferta Moore, 1911 I

Genus Eunice Cuvier, 1817 II

Type Species. Nereis aphroditois Pallas, 1788

m

Diagnosis. Prostomium bilobed or anteriorly entire, bearing 5 antennae with short, smooth ceratophores I

and smooth or annulated styles; 1 or 2 pairs of eyes usually present. Maxillary apparatus with falcate MI,

paired MH, left MIII, asymmetrical MIV with the right one being longer and bearing more teeth than the left •

one, small edentate or unidentate MV, and occasionally small MVI resembling MV. Mandibles flat, about as I

large as maxillae, with elongate shafts bearing crescentic, medially cleft cutting plate. Peristomium consisting

of 2 apodous tings; posterior ring with apair of tentacular cirri arising from anterior margin. Setae including II

hooded compound falcigers, simple limbate capillary and pectinate setae, and bi- to tridentate, hooded 1

subacicular hooks. Branchiae rarely absent, often pectinate, starting between setigers 5 and 10, usually

present in all but last few prepygidial segments.

Remarks. The shape of the anterior prostomial margin is usually included in a species description, i

but it may not be of much taxonomic value. For the most part, the degree to which a prostomium is notched

anteriorly depends on the orientation of the prostomium. If it points anteriorly, the notch is always visible in •

dorsal view; if it points slightly downward, the anterior margin appears smoothly rounded because the actual 1

margin cannot be seen from above.

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266 |
I

1

I Eunice americana Hartman, 1944

I Figure 10.1

Eunice americana Hartman, 1944:118-121, pl. 8, figs. 164-t74, 189; 1968:727.--Fauchald, 1970:19, pl. 1,

I figs. d-e; 1992:51, fig. 8,tab. 42, 43.--Lissner et al., 1986:D-11 .---Steinhauer and Imamura, 1990:F-1.

Material Examined. California, western Santa Barbara Channel, off Point Conception, Sta. 81 (1),

I Sta. 101 (3); Santa Maria Basin, off Purisima Point, Sta. R4 (1).

Description. Length to 105 mm, width to 4 mm, segments to 133. Body robust, dorsoventrally

flattened, with delicate, conspicuous branchiae arched over dorsum of anterior segments. Color in alcohol

I pale.

Prostomium incised anteriorly, with two pairs of large eyes (occasionally only one pair visible in

i preserved specimens) between bases of inner andouter lateral antennae. Antennae with indistinct cylindrical articles; outer lateral antennae shortest, as long as peristomium and first setiger; inner lateral antennae up to

5 times longer, usually as long as peristomium and first 4 to 8 setigers; median antenna as long as inner

i laterals or only about half as long. Tentacular cirri slender, tapering, projecting to tip of prostomium or slightly beyond (Fig. 10.1A).

Maxillary formula: MI 1+1, MII 8-9+8-9, MIH 9-10+0, MIV 8+10, MV 1+1, large cones; MVI

I small, edentate plates or absent (Fig. 10.1B). Parapodia with simple, basally inflated ventral cirri and indistinctly articulated, medially inflated

dorsal cirri; pre- and postsetal lobes low, rounded folds, acicular lobes conical throughout. Branchiae present

I infilame shonrttsantinef'trst rior two bodysetigers, part, occurring then quickly frombecoming setiger 3 pectinate, through sestems tiger disti 34 onrctly35 (longer range than 33-38); dorsal withcirri 2 orand3

not much wider than filaments, with up to 23 filaments where best developed, usually on setigers 10-15 (Fig.

I serrated; 10.1C). compound Last 2 branhooded chiae shooks imple.withSetaelong,nuslender merous,blades in dense, bearingbustwo hy fasc small, icles; widely limbadiverging te setaeteeth, longesappearing t, fmely

almo st unidentate when worn; hoods with distinct mucrons (Fig. 10.1D). Pectinate setae short, with about 15

I teeth, presentthefromoutermos setigert 23-25 ones lon(setiger gest, of29equal in oneorspecimen), subequal length Iridentate, (Figusually . 10.1E).occurring Subacicular singly,hooks occasionally yellow, fi3rorst

4 hooks present including replacement hooks (Fig. 10.1F). Aciculae yellow, 2 per parapodium, with slightly

I curved andRematapered tips projecting through parapodial wall (Fig. 10.1G). rks. E. americana is one of the most common eunicids offthe California coast. The species is

easily recognized by the pointed hoods of the compound setae and the numerous filaments of the gills.

I Type Locality and Type Specimens. California, offRedondo Beach, 256-309 m: holotype (LACM- AHF Poly 0725).

Habitat. Living in branched, parchment-like tubes in sandy and silty sediments.

I Distribution. Central California to western Mexico, 27 - 357 m.

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i 267
I

Figure10.1. Euniceamericana: A,anteriorend,dorsal view; B, maxillae; C, parapodium from branchial region,1

anterior view; D,compound falciger; E, pectinateseta; F, subacicularhook; G, tipsof aciculae. (D-

G after Fauchald, 1992).•
I

Eunice muiticylindri Shisko, 1981

Figure 10.2

I Eunice multicylindri Shisko, 1981:971-973, fig. 2, tab. 1.--Lissner et al., 1986: D-11.--Fauchald, 1992:227-

228, fig. 75f-i, tab. 10, 41, 42.

I Eunice vittatopsis: Lissner et al., 1986: D- 11 (in part).

Material Examined. California: Santa Mafia Basin, off Point Estero, Sta. 6 (1); off Point Buchon,

I Sta. 16 (2); off Point San Luis, Sta. 27 (1)

Description. Length to 45 mm, width to 1mm, segments to 132. Body slender,cylindrical, uniformly

I pale in alcProstomium ohol. rounded, with 2 large, dark eyes behind bases of inner lateral antennae. Antennae with

3-6 club-shaped articles, relatively thick compared to overall body size. Peristomial cirri indistinctly articulated,

I articles cylindrical; reaching to posterior margin or middle of prostomium (Fig. 10.2A). Jaws weakly sclerotized and not calcified. Maxillary formula: MI 1+1, MII 8+8, MIII 8+0, MIV

7+10, MV 1+1, according to Fauchald (1992); MVI absent.

I with truncated and lobes and conical acicular lobes from which of

Parapodia low, pre- postsetal tips

aciculae are emerging (Fig. 10.2B, C). Dorsal and ventral cirri smooth, ventral cirri slender in first 5 to 6

setigers. Branchiae first present from setiger 3 or 4, simple in first 2 to 7 setigers, then becoming pectinate

I maximally (usually 4) to simple, region extending through

with 6 filaments; last 10 15branchiae Branchial

20 to 40 setigers, representing less than half of body length. Compound falcigers with moderately long blades

bearing blunt hoods; subdistal tooth much larger than distal one (Fig. 10.2D). Pectinate setae with about 12

I teeth, outermost ones longest, subequal (Fig. 10.2E). Subacicular hooks yellow, tridentate, with distal tooth

much smaller than subdistal one (Fig. 10.2F); hooks occurring singly, first present in setiger 13 (young

specimens) to about 30. Aciculae yellow, tapering, with free filamentous tips.

I Remarks. This species is very similar to E. vittatopsis Fauchald, 1970 and can best be separated

from the latter by the relatively thick antennae and the shorter branchial region; the absolute number of gills

overlaps between the two species.

I Type Locality and Type Specimens. Southern California: holotype (USNM 63066) and 5 paratypes

(USNM 63068-63970).

I Habit Dis at. In coarse sand or on rocky bottom; tubes with outer lining of relatively coarse sand grains. tribution. Central and southern California, 70 to 611 m.

I Eunice multipectinata Moore, 1911

Figure 10.3

I Eunice multipectinata Moore, 1911:248-251, pl. 15, figs. 20-23.--Hartman, 1944:112-113.--Fauchald,

1970:36-37, pl. 3, figs. h-i; 1992:228-229, fig. 76a-i, tab. 27, 32.--Lissner et al., 1986: D-11.

I

Material Exmed. California: Santa Mafia Basin, off Point Estero, Sta. 6 (2).

I with slighDetlsycriflat ptionten.ed Lven engtrthum; to 205 widesmm, t inwidth pharytong1e1alrarm, egion, segments gentlytotape 181.ringBodytowarobust, rd pygidium. nearly cylindrical Color in

alcohol pale, in life variously pigmented, with pearl-colored to orange-brown or yellow dorsum, banded,

I mottled, or with scattered irregular red blotches in most anterior setigers.

I 269
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I

I

.... m_ 13n I

;...ii,.:... ,_!..;]:ii_:] I .!i!i_i_i:i"::"]":: i i_i _:':]:_:_ oL,n I

A _ I

E

i_i!_. I

".._:':_.

br dC I

I

i

Figure 10.2. Eunice multicylindri: A, anterior end, lateral view; B-C, parapodia from branchial and postbranchial I region, anterior vierw; D, compound falciger; E, pectinate seta; F, subacicular hook. (A, D-F after

Fauchald, 1992).

I

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270 I
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I Prostomium deeply notched anteriorly, withdrawn into anterior peristomia] ring to antenna] bases,

less than half as deep as peristomium. Antennae irregularly articulated, median one (A-m) longest, reaching

to setiger 7-11, with 7-12 articles; inner lateral antennae (A-H) reaching to setiger 4-8, with 5-11 articles;

I outer antennae (A-I) shorter, reaching setiger -3, Eyes conspicuous, dark,

lateral much 1 with 4-7 articles. but

easily overlooked when head is withdrawn into peristomium. Anterior peristomia] ring about 5 times as long

as posterior one; peristomial cirri slender, tapered, with 5-6 articles, reaching posterior margin of prostomium

I or front of anterior peristomial ring (Fig. 10.3A).

Jaws very hard, strongly calcified and sclerotized (dark brown) in adults, less sclerotized (white to

I light MV brown) in younger specimens. Maxillary formula: MI 1+I, MII 5-8+5-8, MIII 6-9+0, MIV 4-7+10-11, 1+1, MVI I+1. Teeth on MII very coarse and offset to close tightly when withdrawn; MIII short, arched

over MI and MII; teeth on MV very large and conical, those on MVI minute. Mandibles with dark shafts and

very hard, white cutting plate with smooth anterior edge, with about 3 large, irregular teeth injuveniles.

I Parapodia with articulated dorsal cirri, at least in anterior setigers; ventral cirri digitiform in f'L,'St5

setigers, basally inflated in middle setigers (Fig. 10.3B), long and slender in far posterior setigers. Branchiae

I present from setiger 7, with 7 f'daments in holotype, 3 in largest examined specimen; number of branchial filaments quickly increasing to maximum number (5 in largest examined specimen, 11in holotype); decreasing

after about 20 setigers, only last one with single filament. Branchial region extending to end of body except

for withlast about 2 or 103 setigers. Limbate setae with narrow, very finely serrated wings; pectinate setae slightly furled, to 15 teeth, one outer tooth much longer than others (Fig. 10.3C). Compound falcigers with

ends of shafts finely and very regularly serrated; blades moderately long, bearing sharply oblique, short hoods

i with setigerserrations 32-45, occurring similar tosingly; shaft (Fig. with 10.3D). large mainSubacicular fang and smaller hooks dark distalbrown tooth toabove jet black, (Fig. first 10.3E). present Aciculae from

large, dark, tapering, straight or slightly bent, especially in posterior setigers, protruding from dorsal tip of

i acicular Pygidium lobes (Fig.small, 10.3F). ring-shaped, surrounding dorsal anus and bearing 2 pairs of anal cirri, both inserted

ventrolaterally; larger pair thick, as long as last 7-11 setigers; smaller pair much thinner and only as long as

I last 1-2 setigers Rema (Fig. 10.3G). rks. There appear to be two oversights in Fauchald's (1992) review of the eunicids considering

this species, both related to the branchiae. Both in the key and the description of E. multipectinata, the

I branchiae as being "shorter are saidthan to benotopodial distinctly cirri longer except thaninthesetigers dorsal 15-30", cirri, butwhich later agrees in the text withthethebranchiae illustrations. are described Fauchald

also describes the branchiae as having "single filaments in fu-st 8-10 branchiated setigers", which clearly

I differs unilateralfrompectinate the illustration throughout". and The alsoexfromaminedMoore's specimens, (1911)bothdescription not fully ofgrown, the branchiae are more sasimilar beingto Moore's "strictly

description in that the branchiae are shorter than the dorsal cirri and pectinate throughout except for the last

I few setigers Ty .pe Locality and Type Specimens. Southern California, off Point Loma Lighthouse: holotype

(USNM 16919).

I Habitat. E. multipectinata is found in sand mixed with mud or rocks.

Distribution. Central California to western Mexico (Baja California), 43 - 270 m.

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Figure 10.3. Eunice muftipectinata: A, anterior end, dorsal view, proboscis partly everted; B, middle parapodium, l

anterior view (after Fauchald, 1992); C, pectinate seta; D, compound falciger; E, subacicular hook;

F, tips of aciculae; G, posterior end, dorsal view. I

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i Eunice vittatopsis Figur Fauchald, 1970 e t0.4

I Eunice vittatopsis Fauchald, 1970:50-52, pl. 7, figs. a-d.--1992:340, fig. 115 j-s, tab. 41, 43.--Lissner et al., 1986:D-11 (in part).

I Material Examined. California: SantaMaria off Purisima Sta. BRA-16 1

Basin, Point, ( ).

Description. Length to 40 mm, width to 2.5 ram, segments to 115. Body slender, oval in crossi section; Prostomium color in alcohol deeply pale,incised often stroanteriorly, ngly iridesce bearing nt. slender, tapered, articulated antennae; eyes absent.

Outer lateral antennae shortest, with about 6 cylindrical articles, as long as peristomium and first 1 to 4

I set to 9igers; cylindrical inner lateralrticles.andPeristomial median antenn cirriareaching e subequal, to orasbeyond long asanterior peristomium prostomial and fimn'st ar6gin, or 7slender, setigers,withwith4 to5

6 cylindrical articles (Fig. 10.4A).

I absent. Jaws well sclerotized; maxillary formula: MI 1+1, MII 8+9, MIII 9+0, MIV 6+13, MV 1+1; MVI

Parapodia with articulated dorsal cirri, smooth ventral cirri, and low, rounded pre- and postsetal

I lobes throughout; acicular lobes bluntly conical, barely projecting beyond and postsetal lobes. Ventral

precirri digitiform through setiger 4, basally inflated with f'fliform tips through setiger 60, more slender and

digitiform to end of body (Fig. 10.4B-D). Branchiae present from setiger 3, pectinate except for first 2

I setigers, with up to 10 filaments about as long as dorsal cirri (Fig. 10.4C, D). Branchial region encompassing most of body length; last branchia of holotype on setiger 85. Simple setae with long, slender serrations along

one margin (Fig. 10.4E), appearing broadly limbate under low magnification; pectinate setae narrow, with 6

I to 8 teeth, one outer one much longer than others (Fig. 10.4F); compound falcigers with moderately long blades; shafts distally slightly inflated, smooth or f'mely serrated; blades with 2 fairly large teeth and smooth

or finely serrated hood (Fig. 10.4G). Subacicular hooks present from setiger 39, tfidentate with very small

I distal tooth (Fig. 10.4H), occurring singly; aciculae usually 2 per parapodium, straight with tapered tips;

subacicular hooks and aciculae yellow.

Remarks. E. vittatopsis is similar to E. multicylindri Shisko, 1986; it can best be distinguished by

I the shape of the antennae, which are more slender in E. vittatopsis and composed of cylindrical rather than drop-shaped articles.

Type Locality and Type Specimens. Mexico, Baja California, Ensenada de San Francisco: holotype

I (LACM-AHF Poly 0337).

Habitat. Found on hard substrata.

I Distribution. Central California to western Mexico (Baja California), intertidal to 92 m.

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A

" I

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Figure 10.4. Eunice vittatopsis: A, anterior end, dorsal view; B-D, anterior, middle, and posterior parapodia, I

anterior view; E, detail of limbate seta; F, pectinate seta; G, anterior (left) and posterior compound

falciger; H, subacicular hook. (B-D, G, H after Fauchald, 1992). •

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i Genus Marphysa Quatrefages, 1865

I Type Species. Nereis sanguinea Montagu, 1815

Diagnosis. Prostomium rounded or bilobed, with 5 occipital antennae arising directly from prostomium,

I not divided into ceratophores and styles; tentacular cirri absent; setae include hooded compound spinigers and falcigers, limbate and pectinate setae, and hooded subacicular hooks. Branchiae usually present, often pectinate,

distributed along short length of body.

I Remarks. The is currently being revised by Fauchald; it differs from Eunice mainly in the

genus

absence of tentacular cirri and the presence of both spinigerous and falcigerous compound setae. Additionally,

the branchial region tends to be shorter in Marphysa than in Eunice. Hartman (1968) lists six species for

I California; only one was found during Phase I and is treated in this chapter.

I Marphysa conferta Moore, 1911 Figure 10.5

I Marphysa conferta Moore, 191I:252-254, pl. 16, figs. 29-34.--Hartman, 1944:129; 1961:83; 1968:727.-- Fauchald, 1970:59-60.--Lissner et al., 1986:D- 11.

I Material Examined. California: Santa Mafia Basin, off Point Estero, Sta. 6 (1).

Description. Length to 33 mm, width to 2 mm including parapodia, segments to 69. Body slender,

I dorsoventrally flattened except for pharyngeal region. Color in alcohol pale to light orange. Prostomium wider thanlong, anteriorly rounded or very slightly incised, depending on whether proboscis

is extended or not. Two large eyes present; all antennae articulated, short, about aslong as peristomium, outer

I lateral antennae shortest. Anterior peristomial ring about 1.5 times longer than posterior one (Fig. 10.5A). Parapodia with low presetal lobe, conical acicular lobe, and conical postsetal lobe in prebranchial

and branchial segments; all parapodial lobes gradually reduced to very low folds in postbranchial setigers.

I Branchiae first present on setiger 7 (range 6-9), only about 11 pairs (range 7-13); first branchia with up to 5

filaments, may be simple; larger than dorsal cirri, with up to 7 filaments on remaining segments, meeting

middorsally where best developed. Dorsal cirri digitiform, tapering, longer than parapodia; ventral cirri

I short, basally inflated (Fig. 10.5B). Setae in dense fascicles, particularly in anterior setigers; simple setae long and delicate; pectinate setae with about 16teeth, outer ones prolonged (Fig. 10.5C); all compound setae

with falcigerous blades, teeth large and widely separated; hoods and distal portion of shafts covered with

I syellow urficiaI spines, appearing serrated in side view (Fig. 10.5D). Subacicular hooks first present in setiger 28, to amber, bidentate, with short, truncate hoods (Fig. 10.5E). Aciculae yellow, 1 or 2 per parapodium,

with straight, tapering free ends; notoaciculae very f'me,difficult to see (reported as absent by Moore).

I Pygidium unknown except for regenerating posterior end of holotype.

Remarks. The combination of very few gills and the absence of compound spinigers is so unmistakable

that this species is not likely to be misidentified; however, there is some confusion in the literature concerning

segment, region was reported by Moore (1911) to extend from X through

the first branchial The branchial

XVHI. This has been misinterpreted as setiger 10 through 18 (e.g., Gathof (1984) in the description ofM. cf.

conferta), when indeed Moore refers to what he calls "somites", which includes the two peristomial rings.

I Somite X is therefore setiger 8. Hartman (1944) recorded M. off southern California in conferta depths

ranging from shallow intertidal to 76 m; however, she does not refer to these records in 1961, but only

mentions that the species was found in "slope depths" (although the depth range is indicated to be to "76-200

I

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meters"); the first gill is described as occurring on setiger 7. In the Atlas of Polychaetous Annelids, Hartman g

(1968) cited the records from 1961, but erroneously described the gills as starting on setiger 4. It is possible

that she saw some specimens with gills from setiger 4 in the material she described in Hartman (1944), which I

she may later have found to be a different species because they are not cited in subsequent publications. I

Type Locality and Type Specimens. California, Santa Rosa Island, off Brockway Point: holotype

(USNM 17328, ANSP slide 6.22). I

Habitat. In sand mixed with mud and rocks.

Distribution. Central California to Baja California, intertidal to 200 m.

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I Literature Cited

I ,_kesson, B. 1967. The embryology of the polychaete Eunice kobiensis. Acta Zoologica 48:165-192.

Bhaud, M. and C. Cazaux. 1987. Description and identification of polychaete larvae; their implications in

I current biological problems. Oceanis 13(6):596-753.

Fauchald, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae,

Lysaretidae and Dorvilleidae from western Mexico. Mlan Hancock Monographs in Marine Biology

I 5:1-335.

Fauchald, K. 1992. A review of the genus Eunice (Polychaeta Eunicidae) based upon type material.

I Smithsonian Contributions to Zoology 523:1-422.

Fauchald, K. and P.A. Jumars. 1979. The diet of worms: a study ofpolychaete feeding guilds. Oceanography

i Fauvel, and P. 1923. MarinePolych_tes Biology Ann errantes. ual Review Faune17:193-284. de France 5:1-488.

i Gathof, J.M. 1984. Family Eunicidae Savigny, 1818. Chapter 401n: Uebelacker, J.M. and P.G. Jones (eds.).

1984. Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Barry A. Vittor &

Associates, Inc., Mobile, Alabama. 7 vols.

I Hartman, O. 1944. Polychaetous annelids. Part V.Eunicea. Allan Hancock Pacific Expeditions 10: 1-237.

U

Hartman, O. 1961. Polychaetous annelids from California. Allan Hancock Pacific Expeditions 25: 1-226.

I Hartm_, University O. 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation, of Southern California, Los Angeles. 828 pp.

i Herpin, R. 1925. Recherches biologiques sur la r6production et le d6vetoppement de quelques Ann61ides Polych_tes. Bulletin de la Soci&6 des Sciences naturelles de l'Ouest de la France 4:1-250.

Hofmann, D.K. 1969. Untersuchungen _iber die Regeneration des Prostomiums und des Hinterendes beim

I PDeutschen olyehaetenZoologischen Eunice viridisGeseltschaft GRAY. Zoologischer Anzeiger, Supplement-Band 33, Verhandlungen der 1969, pp. 253-260.

Hofmann, D.K. 1972. Nachweis der Epitokie als Fortpflanzungsmodus des Polychaeten Eunice siciliensis

i (Polychaeta: Annelida). Marine Biology 14(4):341-344.

Hofmann, D.K. 1974. Maturation, epitoky and regeneration in the polychaete Eunice siciliensis under field

I and laboratory conditions. Marine Biology 25:149-161.

Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bemstein, R. Cimberg, T. Kauwling, and W. Anikouchine.

1986. Assessment of long-term changes in biological communities of the Santa Maria Basin and

I - Phase I. Report prepared for the U.S. Department of the Irttedor,

Western Santa Barbara Channel

Minerals Management Service, Pacific OCS Region, under contract No. 14-12-0001-30032.

Moore, J.P. California 1911.inThe1904: polychaeto 111.Euphrosynidae us annelidstodredged Goniadidae. by theProceedings U.S.S. "Albatross" of the Academy off theofcoast Natural of southern Sciences

of Philadelphia 1911:234-318, pls. XV-XXI.

I Orensanz, J.M. 1990. The annelids from antarctic and Subantarctic eunicemorph polychaete seas. Biology

of the Antarctic seas XXI. Antarctic Research Series 52:1-183.

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Pettibone, M.H. 1982. Annelida. In: Synopsis and Classification of Living Organisms. McGraw-Hill Book I

Co., Inc., pp. 1-43.

Pillai, T.G. 1958. Studies on a brackish-water annelid, Marphysa borradailei sp.n. from Ceytan. Ceylon I

Journal of Biological Sciences 1(2):94-106.

Shisko, J.F. during1981. the Five Southemnew California polychaetesBaseline of the families Project. eunicidae ProceedingsandofOnuphidae, the Biological collected Societyin 1975 of Washington and 1976 I

94: 968-983.

Steirdaauer, M. and E. Imamura (eds.). 1990. California OCS Phase II Monitoring Program. Year-Three I

Annual Report. Prepared for the U.S. Department of the Interior, Minerals Management Service,

Pacific OCS Region, under contract No. 14-12-0001-30262.

W'msnes,I.M. 1989. Eunicid polychaetes (Annelida) from Scandinavian and adjacent waters. Family Eunicidae. I

Zoologica Scripta 18(4): 483-500.

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11. FAMILYLUMBRINERIDAEMALMGREN, 1867,

I EMENDED ORENSANZ, 1990

by

I Brigitte Hilbig _

I

Introduction

I

Lumbrinerids are long, cylindrical, burrowing worms that can reach considerable size, especially

i tropical species belonging to the genus Lysarete that attain lengths of more than 60 cm. Despite the relatively featureless morphology, a remarkably large number of species have been described. The genus Lumbrineris

s.1.alone contains about 200 species, and many others still await description. Lumbrinerids can be distinguished

from the similar looking Oenonidae by the shape of the prostomium; it is always rounded in side view in

I dorsoventrally of the pharynx will also reveal that the two

lumbrinerids and flattened in oenonids. Dissection

families have very different jaw structures.

I Morphology

I The prostomium is conical to globular and usually about as long as wide except in the genera

Lumbrinerides and Lumbrineriopsis where the elongate prostomia may be up to seven times longer than

I wi andde.twoAnte pairsnnaofe, eyes palps, areandpresent eyes are (nuchal usuallypapillae, absentocexcept casionally for Lyseen sareteeverted and Kufromwaitaa groove where atthrethee antennae posterior

margin, are not homologous to antennae). The eversible pharynx is armed with five pairs of maxillary plates

I (generally referred toforceps-shaped as MI through and MV)usually and paired unidenmand tate;iblesLysare embed te and dedLumbrinerides inthick musclepossess pads. The bi- first oridentate maxillae MI. are

Basally the first maxillae are attached to roughly triangular, often laterally incised maxillary carriers. On the

i outs are located ide of eventral ach MI,ofa the slenMI, der usually support about piece the is attsame achesidz(e"bri (except dles" for of Ore thengenus sanz, Eranno 1990). The wheresecotheyndaremaxillae much

smaller) and armed with 3 to 8 teeth with internal canals. The first and second maxillae are basatly connected

i bypiecethin,forms threadlike the connection ligaments;between in the genu MI and sEranno, MII. The a withird, de, ribbfourth, onlike,andsclerotize fifth maxillae d ligament are located or additiino

front nal jawof

MI and MII and considerably smaller. MIII and MIV are usually rounded plates with a thick, hard cutting

i edge that may form a few teeth devoid of internal canals. Ninoe and Lumbrineriopsis have MIV (sometimes also Mill) with multidentate cutting edges;Augeneria possesses large MIV with a white center and dark rim.

The fifth maxillae are not present in all species, and they may be completely fused to MIV (Abyssoninoe). In

the literature they are generally referred to as lateral supports. The mandibles consist of two shafts bearing a

hard, more or less crescent-shaped plate with a smooth cutting edge that may bear a few large, irregular teeth.

The shafts are usually fused, at least partially, and running parallel when free from each other, except for

Augeneria where the shafts are divergent so that the mandibles are roughly X-shaped. All jaw pieces are

I formed folds of the inner

by pharyngeal lining.

I 1ENSRConsultingand Engineering,89WaterSlreet,WoodsHole,Massachusetts02543

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The peristomium consists of two apodous and achaetous rings (not true segments) that dorsally m

resemble the following setigers. Ventrally the peristomium is developed into a pair of buccal cushions and the

posterior margin to the mouth. Except for some slight tapering toward the prostomium and the pygidium, the m

setigers are very similar. Small species reach their greatest body width in the pharyngeal region, but large l

individuals are of uniform width along the entire length of the body. Parapodia are subbiramous, with the

notopodium usually reduced to a small bundle of fine notoaciculae; large individuals may show an externally •

visible notopodial rudiment. The neuropodia have presetal lobes that are usually short and rounded and I

postsetal lobes that are longer than the presetal lobes. OnlyKuwaita andLysarete possess dorsal and ventral

cirri; palmate branchiae emerge from the back of the postsetal lobes ofNinoe. Vascularized postsetal lobes, •

functioning as gills, are found in anterior setigers of Paraninoe and in middle and posterior setigers of m

Lumbrineris, Eranno, and Scoletoma (Lumbrineris s.1.). Neuroaciculae are conspicuous, yellow to black,

often occurring in bundles of two to about six in a parapodium. Setae include simple limbate capillaries •

(Lumbricalus has compound limbate setae in addition) and hooded hooks that are either bidentate or bear a m

crest of small teeth above a main fang; they may be compound in some anterior setigers. Hooded hooks are

absent in Arabellonereis.

The pygidium bears one or two pairs of short anal cirri and a terminal anus; some species lack anal m

cirri, and the anus is dorsal.

Taxonomic History I

I

The morphological homogeneity of the lumbrinerids and the paucity of external features has made m

generic definitions difficult ever since the first lumbrinerids were described. Most of the lumbrinerid species m

were erected in the I9th century by Kinberg, Grebe, Mclntosh, Ehlers, andothers, and while the majority of m

the species are still valid, the genus has been changed more than once for many of them. The oldest species is 1

Scoletomafragilis (O.E Mfiller, 1776) described asLumbricusfragilis. Blainville (1828) erected Scoletoma

for this species. The genus was subsequently synonymized withLumbrineris by Audouin and Milne-Edwards 1

(1833 ), but was recently resurrected by Frame (t 992) to include specie s formerly assigned to Lumbrine ris

that are characterized by the presence of simple hooded hooks only. Eranno, erected in 1865 by Kinberg, has

a similar history. Kinberg considered the everted nuchal organ ofE. bifrons, the type species, as a diagnostic m

character; this proved to be invalid, and Eranno was synonymized with Lumbrineris by Hartman (1949).

Orensanz (1990) resurrected the genus with a different set of diagnostic characters, including details of the

jaw apparatus. Similarly, Augeneria Monro, 1930 was defined by the presence of an everted nuchal organ, m

but later emended by Orensanz (1973) based on characters of the mandible and maxillae. The emphasis in

lumbrinerid systematics has thus shifted in the last 25 years from the often rather ill-defined differences in

external morphology to more reliable, although less readily accessible characters of the jaws, a trend that can m

be observed in the systematics of all euniceform polychaetes. The generic revisions by Orensanz (1990) and

Frame (1992) mark the end point of about 30 years of aiming toward integration of fossil and extant records

of euniceform polychaetes to develop a more natural system for these families. Important milestones include •

publications by Kielan-Jawomwska (1966, 1968) on fossils and Hartman (1944) and Fauchald (1970) on

extant Lumbrineris.

!

l

I

280 m
I

Distribution and Biological Notes

I Most lumbrinerids live in soft bottoms the continental but found within

on shelf, some species are a

large depth range between the intertidal and abyssal zone. They are generally free living burrowers, although

some species construct temporary mucus tubes (Fauchald, 1977; Perch, 1986). Some species live on hard

I substratum such as reefs. Lumbrinerids have been variously reported to be carnivores, detritus

scavengers,

feeders, and deposit feeders. The strong musculature and the large, heavily sclerotized jaw apparatus suggest

that food is picked up by a grasping and sucking motion of the eversible pharynx. Petch (1986) observed

I feeding habits of two Australian populations of Lumbrineris cf. latreiUi and concluded that this species, at least in the study area, is a selective deposit feeder. The animals construct mucus-lined tubes down to about

11 cm into the sediment; from this main tube, they burrow horizontal side tunnels in which they feed.

I is little information on the reproduction of lumbrinerids; a summary can be found in Bhaud and

There

Cazaux (1987). The few studies available indicate that the eggs are deposited in gelatinous ribbons or masses

from which large lecithotrophic larvae with at least four setigers eventually emerge. The pelagic stage is very

I short, and lumbrinerid larvae found in The first the 2- 3-

ale seldomly plankton samples. setae appear at to

setiger stage; they are very broadly limbate and can still be seen injuveniles with about ten setigers (personal

observation), but are later replaced by limbate capillaries with moderate to narrow wings and long, fine tips.

I The jaw apparatus is visible at the 4-setiger (Bhaud and Cazaux, 1987).

stage

I Key to the Lumbrineridae

I genera areTheincluded key follows because the getheneric newlyconestablished cepts established genra bymayOrensanz not yet be(1990 widely ) andknown. Frame (1992). Genera that All known are not

represented in the Santa Mafia Basin and Santa Barbara Channel are marked with a footnote. Paraninoe and

I Abyssoninoe may be synonyms of Scoletoma.

1A. Parapodia with notopodial rudiments and dorsal cirri; prostomium with 3 antennae ..................... 2

I absent; notopodia usually reduced to small bundle of thin notoaciculae, not visible

lB. Dorsal cirri

externally; antennae absent .............................................................................................................. 3

I 2A. MI bifid ............................................................................................................................... Lysarete 2

2B. MI falcate, unidentate ......................................................................................................... Kuwaita 2

I

3A. All setae simple limbate capillaries; hooded hooks absent ....................................... Arabellonereis 2

I 3B. Hooded hooks present on at least middle and posterior parapodia ................................................... 4

4A. Prostomium much longer than wide, acutely pointed; hooded hooks bifid ...................................... 5

I 4B. Prostomium conical to rounded, about as long as wide; hooded hooks multidentate (worn hooks on

small species may look bidentate) .................................................................................................... 6

!

I 2GenusnotknownfromCalifornia.

I 281
!

5A. Mandibles fused entirely; MI bi- or tridentate, MIV with smooth cutting edge (also referred to as

unidentate), maxillary carriers very large and wide ................................................. Lumbrinerides 3 I

5B. Mandibles free along posterior part; MI falcate, unidentate; MIV multidentate; maxillary carriers

long and slender ..................................................................................................... Lumbrineriopsis 2

!

6A. Setae including simple limbate capillaries, limbate compound setae, and simple and compound hooded

hooks ............................................................................................................................ Lumbricalus z •

6B. Setae including simple limbate capillaries and simple hooded hooks, sometimes also compound hooded n

hooks in anterior setigers ................................................................................................................ 7

!

7A. Anterior notopodia with palmate gills; Mill and MIV, or MIV only, with multidentate cutting edge:

genus ........................................................................................................... genusNinoe ............... 8

7B. Gills absent or represented by vascularized postsetal lobes of anterior pampodia ....................... 12 I

8A. Hooded hooks present from setiger 1 to 3 ....................................................................................... 9 n

II

8B. Hooded hooks first present on or after setiger 7 ........................................................................... 11

9A. Gills present in setigers 2 to 17, with maximally 6 filaments; hoods of anteriormost hooks short and I

rounded; MII with 4 teeth, MIII and MIV multidentate ................................................. Ninoe sp. C

9B. Last gill present between setigers 30 and 50; hoods of anteriormost hooks long and slender ....... 10 I

10A. Gills fh'st present on setiger 2, with maximally 5 filaments; MII with 6 teeth, MIV multidentate ..... []

................................................................................................................................... Ninoe palmata |

10B. Gills first present on setiger 5 or 6, with maximally 4 fdaments; MII with 8 teeth, MHI and MIV

multidentate ................................................................................................................ Ninoe gemmea n

11A. Gills fn-stpresent on setiger 1or 2, with maximally 10 filaments; MII with 4 to 5 teeth, Mill and MIV

multidentate (?) .............................................................................................................. Ninoe sp. D

11B. Gills fin'st present on setiger 2 or 3, with maximally 11 filaments; MII with 8 teeth, MIII tridentate,

MIV multidentate .................................................................................................... Ninoe tridentata n

u

12A. Postsetal lobes of anterior parapodia vascularized; aciculae black; prostomium with two longitudinal

silts ................................................................................................................................... Paraninoe a n

12B. Vascularized lobes, if present, on posterior parapodia; aciculae yellow or black; prostomium without

dorsal slits ...................................................................................................................................... 13

!

13A. Setae including simple limbate capillaries and simple hooded hooks (may be absent or transitional in

anteriormost setigers) ..................................................................................................................... 14 •

13B. Setae including simple limbate capillaries and simple and compound hooded hooks ................... 17 II

(also referred to as lateral supports) fused with MIV; aciculae yellow ................. Abyssoninoe 2 n

14A. MV

3L. acutus occurs in California, but is not included in this Atlas. See Hartman (1968). I
!

14B. MV free or partially fused with MIV; aciculae yellow or black .................................................... 15

I 15A. MV free from MIV; MIII with smooth cutting edge or 1 to 3 teeth; MI and MII of subequal length,

connected by thin, threadlike basal ligament; aciculae yellow; posterior parapodia with elongate,

conical postsetal lobes; M/II bidentate; hooks from setiger 1,in middle and posterior setigers

present

with numerous small teeth and slightly larger main fang ..................................... Scoletoma tetraura

i 15B. MV partially fused to MIV; MIH with smooth cutting edge; MII to 1/2as long as MI, connected to MI by wide, ribbonlike, sclerotized ligament or additional jaw piece ............... genus Eranno ........... 16

I 16A. Acic digitiform, ulae blabout ack, 3asto 6 per parapodium; pre-and postsetal lobes of posterior parapodia elongate, long as setae ............................................................................ Eranno bicirrata

16B. Aciculae yellow to amber, I per parapodium; pre-and postsetal lobes of posterior parapodia elongate,

I digitiform, projecting about twice as far as setae Eranno lagunae

I 17A. Mandible rim; MV absent only slightly ............................................................................................................... longer than wide, roughly X-shaped; MIV large, with white centerAugeneria and dark2

17B. Mandible much longer than wide, roughly Y-shaped; MIV uniformly dark; MV present in most

I species, free from MIV ............................................................................... genus Lumbrineris ... 18

18A. Aciculae yellow .............................................................................................................................. 19

m 18B. Aciculae dark brown or black ........................................................................................................ 22

I 19A. At least postsetal lobes of posterior parapodia elongate ................................................................ 20

19B. Pre- and postsetal lobes short and rounded throughout ................................................................. 21

I 20A. Pre-and postsetal lobes of posterior parapodia elongate and digitiform; blades of compound hooks

short; hoods of shaft inflated and short, hoods of simple hooks of the same shape ...........................

I ....................................................................................................................... Lumbrineris cruzensis

20B. Only postsetal lobes of posterior parapodia elongate; blades of compound hooks about 4-5 times as

long as wide, all hoods slender ........................................................................ Lumbrineris limicola

I

21A. Prostomium conical; MIII with 2 teeth, MIV unidentate; compound hooks with slender blades, 6-7

I times as long as wide ......................................................................................... Lumbrineris latreilli 2lB. Prostomium globular; Mill with 3-5 teeth, MIV bidentate; compound hooks with short blades, about

3-5 times as long as wide .................................................................................... Lumbrineris inflata

!

22A. At least postsetal lobes of posterior parapodia elongate ............................................................... 23

I 22B. Pre- and postsetal lobes short and rounded throughout; MIII bidentate .......... Lumbrinerisjaponica

23A. Pre- and postsetal lobes of posterior parapodia digitiform and elongate; Mffl unidentate, maxillary

I carriers slender, about 3 times as long as wide Lumbrineris californiensis

23B. Only postsetal lobes of posterior parapodia elongate, conical; presetat lobes short and rounded

throughout; MIII bidentate, maxillary carriers less than twice as long as wide, very broad ..............

I .............................................................................................................................. Lumbrineris index

I 283
!

!

Description of Species

i

Two species of Eranno, one species of ScoIetoma, seven species of Lumbrineris s. str., and five

species of Ninoe were found in the Santa Maria Basin and Western Santa Barbara Channel. One species, m

Ninoe tridentata, is new to science. The following species are treated in this account: i

Eranno bicirrata (Treadwell, 1929) •

Eranno lagunae (Fauchald, 1970) 1

Lumbrineris californiensis Hartman, 1944

Lumbrineris cruzensis Hartman, 1944 i

Lumbrineris index (Moore, 1911) 1

Lumbrineris inflata Moore, 1911

Lumbrinerisjaponica (Marenzeller, 1879) I

Lumbrineris latreilli (Audouin & Milne-Edwards, 1834) 1

Lumbrineris limicola Hartman, 1944

i

Ninoe gemmea Moore, 1911

Ninoe palmata Moore, 1903 i

Ninoe tridentata Hilbig, new species

n

Ninoe sp. C i

Ninoe sp. D 1

1

Scoletoma tetraura (Moore, 1911)

1

Genus Eranno Kinberg, 1865, emended Orensanz, 1990 I

Type Species. Eranno bifrons Kinberg, 1865 I

Diagnosis. Setae including limbate capillaries and simple hooded hooks that may be very slender in 1

anterior setigers. MII only about half as long as MI, connected to MI with wide, sclerotized ligament or 1

additional jaw piece. MV present, partially fused to MIV or free.

u

Remarks. Orensanz (1990) provided the taxonomic history of this genus andemended itto encompass •

a number of species previously assigned toLumbrineris. His generic definition is followed here and slightly 1

modified to include species with free, rather than partially fused, MV.

1

Eranno bicirrata (Treadwell, 1929), new combination I

Figure 11.1 •

Lumbrinereis bicirrata Treadwell, 1929:1-3, figs.l-7.

Lumbrineris bicirrata: Hartman, 1944:156-158, pl.9, figs. 207-212; t968:745.--Fauchald, 1970:77-78, pl. 1

10, figs. e-g.--Lissner et al., 1986:D-11.--Hyland and Neff, 1988:A-2.--Hyland et al., 1990:F-1. 1

Lumbrineris nr. bicirrata: Lissner et al., 1986:D-12. 1

Material Examined. California: Santa Mafia Basin, off Point San Luis, Sta. R-1 (1); off Point I

Arguello, Sta. 58 (2), Sta. 67 (1); Western Santa Barbara Channel, Sta. 85 (1).

I

!
Figure 11.1. Eranno bicirrata: A, anterior end, dorsal view, peristomium slightly distorted because of everted

I B, dorsal view; C, and extended lateral proboscis; posterior end, prostomium partly proboscis,

view; D, maxillae, dorsal view, left Mill - MV omitted, right MIII - MV slightly displaced for

better visibility; E, mandibles; F-H, anterior, median, and posterior paxapodium, anterior view; I,

hooded hook, setiger 15; J, same, setiger 120 (fi'om Fauchald, 1970).

i 285
!

Description. Length to 200 mm, width to 7 ram, setigers to about 300. Body long, of equal width I

along greatest portion, widest in pharyngeal region, tapering toward pygidium within last 20 to 30 segments

(Fig. ll.IA, B). Color in alcohol pale. •

Prostomium conical, about as long as wide, distinctly narrower than first segments, slightly depressed

dorsoventrally (Fig. 11.1A, C). Maxillae: MI large, pointed, with squarish, flared base, distally very slender;

MII much smaller, only about one-third as long as MI, with 4 major teeth and 2 to 3 additional small ones I!

proximally, often developed as barely visible ridges; major teeth with white calcified tips. MIII elongate, very

hard, with prominent conical tooth; MIV large soft plate, tapering to single tooth; MV square, weakly sclerotized

plates. Maxillary carriers rounded, fused along distal half, darkly sclerotized but soft. MII attached to II

carriers by very broad sclerotized ligament. Pharyngeal wall alongside MI sclerotized, forming elongate, soft

lateral support pieces (Fig. 11.1D). Mandibles slightly longer than maxillae, light-colored but very hard;

handles fused along distal half, bearing flared cutting edge with concentric lines and several irregular teeth, •

very dark except for middle ones that are white and very hard (Fig. 11.1E).

Parapodia bluntly rounded in anterior segments, with low presetal lobe, slightly longer acicular lobe,

and asymmetrical, bluntly triangular postsetal lobe surpassing acicular lobe (Fig. 11.1F). By about setiger 1

75, postsetal lobes becoming more pointed, gradually developing into digitiform structure and increasing in

length; presetal lobes with papilliform projection directed upward, gradually developing into digitiform lobe

matching postsetal lobes; both lobes longer than setae where best developed, reaching about one-half body 1

width (Fig. 11.1 G, H). All setae simple, including limbate setae in anterior third of body and hooded hooks

starting from setiger 1 (young individuals) to about 20; hoods long and slender in anterior segments, gradually

shortening and widening; hooks with small heads bearing few very fine teeth in anterior setigers, much stouter 1

and with main fang surmounted by several smaller teeth thereafter (Fig. I1.1I, J). Aciculae numerous, up to

6 in anterior parapodia, 3 or 4 in middle and posterior parapodia; setae dark amber at bases, distally colorless.

Pygidium small, with two ventral cirri and two shorter dorsal ones. 1

mat

Remarks. Lumbrineris bicirrata was not included in Orensanz's list, but agrees with his generic

definition and is therefore referred to Eranno.

am

Type Locality and Type Specimens. Friday Harbor, Washington; holotype: AMNH 1692. !

Habitat. Eranno bicirrata lives in sand, silt, and mixed sediments.

Distribution. Puget Sound, Washington to western Mexico, shallow subtidal to 2500 m. !

Eranno lagunae (Fauehald, 1970) •

Figure 11.2

Lumbrineris lagunae Fauchald, 1970:92-94, pl. 15, figs. a-e.--Lissner et al., 1986:D-12.--Hyland et al., l

1990"F-1. IB

Lumbrineris bifilaris: Hart•an, 1944:153-155, pl. 9, figs. 196-206. Not Ehlers, 1901. I

Lumbrineris sp. A Lissner et al., 1986:D-12. l

Eranno lagunae: Orensanz, 1990:78.--Frame, 1992:195. i

Material Examined. California: Santa Maria Basin, off Point Buchon, Sta. 12 (1); off Point 1

m

Conception, Sta. 73 (1); off Point San Luis, Sta. R-1 (1).

Description. Length at least 85 mm, width to 3 mm, segments more than 150. Body robust, almost •

cylindrical, with parapodia inserted ventrolaterally; body widest behind pharynx, around setiger 20. Color in I

alcohol pale, cuticle iridescent.

!

286 |
I

I

, !

I

I

I v

JIJ V

I

no

I ::: , F

I °

postL

!

Figure 11.2. Eranno lagunae: A, anterior end, dorsal view; B, maxillae, dorsal view, in situ; C-F, anterior,

median, posterior,and far posterior parapodium, anterior view; G, hooded hook, anterior setiger,

I with detail of crest; H, same, posterior setiger.

I

i 287
!

Prostomium conical, about as long as wide (Fig. 11.2A). Maxillae with short, oval, laterally incised I

carriers and large, basally almost rectangular forceps; MII only half as long as MI, with 4 large and 0 to 2

much smaller teeth, large teeth with white calcified tips; MIII elongate, hard plates with single tooth, MIV •

large, rounded, soft plates with single tooth; MV large, roughly triangular, soft plates. MII attached to

carriers with long, broad ligaments and also thin ones; lateral supports long and slender (Fig. 11.2B).

Parapodia short in anterior setigers, with obliquely truncate pre- and postsetal lobes and distinct

notopodial rudiment (Fig. 11.2C); in middle setigers parapodial lobes reduced, with postsetal lobes only as

long as presetal lobes (Fig. 11.2D); posterior to about setiger 70 both lobes increasing and becoming digitiform,

about twice as long as parapodial stem and surpassing setal fascicles in far posterior setigers (Fig. 11.2E, F). R

Limbate setae present in anterior setigers; hooded hooks simple throughout, very slender and long-hooded in

anterior setigers, with hoods gradually shortening in middle setigers. Anterior hooks with very small crest

bearing 7 to 8 f'me teeth (Fig. ll.2G), middle and posterior hooks with main fang surmounted by 10 to 14 i

irregular teeth (Fig. 11.2H). Aciculae single in each parapodium, yellow to amber, darkest in posterior

parapodia of large specimens.

Remarks. E. lagunae is most easily distinguished from the co-ocurring E. bicirrata by the color of i

the aciculae (yellow inE. lagunae, black in E. bicirrata) and the proportions of the parapodia along the body.

Type Locality and Type Specimens. Laguna Beach, California; holotype: LACM-AHF Poly 000340. i

Habitat. Sand mixed with silt. Bl

Distribution. Western Canada to western Mexico, shallow subtidal to 110 m.

!

Genus Lumbrineris Blalnville, 1828 l

Type Species. Lumbrineris latreilli Audouin and Milne Edwards, 1834

Diagnosis. Setae including simple limbate capillaries, simple hooded hooks, and compound hooded i

hooks. MII about as long as MI, connected to MI with short, thin basal ligament, with aliform lateral

expansions; MIII and MIV with aliform expansions; MV present, free from MIV. •

Remarks. Frame (1992) provides an account of the taxonomic history ofLumbrineris and discusses 1

the problem of a missing type species of the restricted genus.

Lumbrineris californiensis Hartman, 1944 i

Figure 11.3 i

I

Lumbrineris californiensis Hartman, 1944:163-165, pl. 12, figs. 257-262; 1968:749.--Fauchald, 1970:78-

80, pl. 10, figs. a-d.--Lissner et al., 1986:D-12.--Hyland and Neff, 1988:A-2.--Hyland et al., •

1990:F- 1. 1

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. 21 (1); off Point Sal, •

Sta. R-8 (1); Tomales Bay, coll. 1957 (1), coll. 1958 (I).

Description. Length to 130 mm, width to 3 mm, segments to 200. Body robust, cylindrical to

slightly depressed, segments longest and widest in pharyngeal region, much shorter in posterior region (Fig. 1

11.3A, B), body tapering toward pygidium within last 20 or so segments. Color pale to purplish, with

iridescent cuticle.

I

288 I
I Figure californiensis: A, view; B, some posterior segments, view;

11.3. Lumbrineris anterior end, dorsal dorsal

C, maxillae, dorsal view, in situ; D, mandibles (after Hartman, 1968); E, anterior parapodium,

posterior view; F,same, anterior view; G, posterior parapodium, anterior view; H, compound hooded

m hooks, same fascicle, setiger 7; I, simple hooded hook (F, G, I after Fauchald, 1970).

m 289
!

Prostomium conical, slightly longer than wide, much narrower than first segment. Maxillae : MI m

short, rounded, falcate; MII as large as MI, with 3 or 4 very coarse teeth bearing white tips in large specimens;

MIII elongate, tapering to single tooth; MIV large, soft rectangular plates bearing single tooth; MV weakly •

sclerotized, squarish plates. Maxillary carriers shorter than forceps (MI), slightly incised laterally. MII l

attached to carriers by short, thin ligaments. Weakly sclerotized lateral supports along MI (Fig. I 1.3C).

Mandibles fused through most of their length, handles free inproximal third; cutting edge flared, with concentric •

rings and irregularly dentated anterior margin (Fig. 11.3D). l

Pampodia minute in first 5 to 15 setigers, gradually increasing to about one-third body width; anterior

parapodia broadly rounded, with short presetal and acicular lobes and somewhat longer postsetal lobe; notopodial m

rudiment conspicuous (Fig. 11.3E,F). Pre- and postsetal lobes gradually lengthening until about setiger 70 to

100, then becoming digitiform and projecting beyond setal fascicles (Fig. ll.3G). Up to 5 aciculae, dark

amber to black; setae yellow at base, clear and colorless distally. Anterior 20 to 30 setigers with composite •

hooded hooks; blades about 4 times as long as wide, with 6 to 8 apical teeth of equal or subequal size, with the

largest tooth in proximal position (Fig. I 1.3H). In further posterior setigers hooks becoming simple, more

stout than composite hooks, with short hood and several progressingly smaller teeth above main fang (Fig. •

11.31). Limbate setae present throughout anterior half of body, in superior and inferior position of fascicle. l

Pygidium with two lateral anal cirri; anus terminal.

Remarks. In her description of L. californiensis, Hartman (1944) stated that the aciculae were m

m

yellow, although in the key this species is among those that have black aciculae. The same discrepancy can be m

found in the Atlas of Polychaetous Annelids (Hartman, 1968). However, a specimen confirmed by Hartman m

(Tomales Bay) was examined here and was found to clearly possess black aciculae. Fauchald (1970) also m

states that L. californiensis has black aciculae. Hartman's statement is therefore regarded as erroneous.

m

Type Locality and Type Specimens. San Nicolas Island, California; holotype: LACM-AHF Poly

50. m

Habitat. L. californiensis is found in sandy and muddy sediments.

Distribution. Washington to western Mexico, in 10 - 1200 m. m

Lumbrineris cruzensis Hartman, 1944

Figure 11.4 I

Lumbrineris cruzensis Hartman, 1944:165-166, pl. 12, figs. 263-269; 1968:751.--Fauchald, 1970:83-84, m

pl. 12, figs. g-j.--Lissner et al., 1986:D-12.--Hyland and Neff, 1988:A-2.--Hyland et al., 1990:F1.

Lumbriconereis cruzensis: Uschakov and Wu, 1965:81, figs. 22A-E m

Lumbrinereis crucensis [sic]: Rioja, 1962:180.

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. 21 (2); off Point Sal, m

Sta. PJ-2 (2), Sta. R-8 (2); Western Santa Barbara Channel, Sta. 79 (4). m

Description. Length to 55 mm, width to 1mm, segments to 200. Body slender, gracile, with segments i

only about as wide as prostomium (Fig. 11.4A); color in alcohol very pale with iridescent cuticle, m1

Prostomium about as wide as long, bluntly conical. Maxillae: forceps (MI) short, rounded, falcate;

MII at least as long as forceps, with 4 large teeth. MIII hard, elongate plates with one tooth; MIV large, soft •

plates tapering to single tooth. Maxillary carriers about as long as forceps, incised laterally, tapering to fine |

tip proximally. MII attached to carriers with short, thin ligaments (Fig. 11.4B). Mandibles fused through

about two-thirds of their length, bearing crescent-shaped plate with irregularly dentate cutting edge (Fig. 11.4C). m

I
I

I

I 7

I A

I

IV

I

II

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I :i]:._,._........ ===_£-__.,-_]._,_!_._ W.i, =-

I " ": "' ' )'"(_""_" .,ii::_;i:(t...... _":";i_i::{_i.,:,.,_

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I

I Figure 11.4. l_zonbrineris cruzerLsis:A, anterior end, dorsal view (after Day, 1973); B, maxillae, dorsal view,/n

sire, fight _ and MIV after Hartman, 1944; C, mandibles; D-F, anterior, median, and posterior

parapodium, anterior view; G, simple hooded hooks, setiger 20; H, compound hooded hooks, sefiger

I 10 (C-F after Hartman, 1944).

i 291
!

Parapodia in anterior segments with bluntly rounded, short presetal and acicular lobes and longer, I

rounded postsetal lobes (Fig. 11.4D); from setiger 60 to 100 pre-and postsetal lobes increasing in length and

gradually turning digit•form, surpassing setal fascicles in posterior setigers (Fig. 11.4E, F). Setae of 3 k_nds: •

(1) in anterior 12 to 21 setigers compound hooded hooks with short blades, about 2 to 3 times longer than l

wide, and apex bearing either 6 to 8 subequal teeth or main fang surmounted by about 6 increasingly smaller

teeth (Fig. 11.4G); both types found in same fascicle; (2) simple hooded hooks, first appearing around setiger •

15, with short hoods and apex beating main fang, 1 to 2 coarse teeth and several much smaller ones (Fig. l

11.4H); (3) limbate capillaries, present in anterior hail of body, Usually 2 to 4 aciculae present; all setae

transparent and colorless, aciculae in juveniles clear, in older specimens yellow. •

Pygidium with 2 anal cirri. I

Remarks. The prolonged pre- and postsetal lobes in posterior setigers combined with yellow or clear

setae and aciculae make this species readily recognizable if whole specimens or anterior ends of at least 100 i

setigers are present; short anterior fragments encompassing only parapodia with short pre-and postsetal lobes

may be confused with L. lira•cola (see below) and should be identified with caution. Examination of the

compound hooded hooks may be helpful because the blades are short and bear relatively large teeth in L. i

cruzensis, but of moderate length with a crest of many equal, slender teeth in L. limicola.

Type Iamality and Type Specimens. Santa Cruz Island, California; holotype: LACM-AHF Poly 51. m

Biology. The species lives in silt and silt mixed with sand, shells, and rocks. In the Yellow Sea, i

gravid females were found from June through August (Uschakov and Wu, 1965).

III

Distribution. British Columbia to western Mexico; Yellow Sea; 4 - 200 m. i

Lumbrineris index (Moore, 1911) I

Figure 11.5 i

Lumbrineris japonica index Moore_ 1911:288-289, pl. 19, figs. 119-127. •

Lumbrineris index: Hart•an, 1944:162-163, pl. 12, figs. 254-256; 1968:755.--Lissner et al., 1968:D- 12.-- I

Hyland et al., 1990:F-1.

Material Examined. California: Santa Maria Basin, off Point Buchon, Sta. 13 (1); off Point Sal, I

Sta. 36 (1); off Purisima Point, Sta. R-4 (1).

Description. Length to 109 mm, width to 4 mm, segments over 200. Body robust, depressed, with i

prominent parapodia throughout. Color in alcohol tan with iridescent cuticle.

Prostomium bluntly conical, depressed, slightly longer than wide. Peristomium and subsequent setigers

not much wider than prostomial base, body width gradually increasing within pharyngeal region, greatest 1

width around setiger 15 (Fig. 11.5A), tapering toward pygidium within last 20 segments or so (Fig. 11.5B).

Maxillae with large, broadly rounded, weakly incised carriers and slender forceps; MII with 4 white-tipped

teeth, the distal two much larger than the proximal two; MIII elongate, hard plates with 2 distinct teeth, MIV 1

large, soft plates with single large tooth. MH attached to carriers with short, thin ligaments (Fig. 11.5C).

Mandibles in adults almost completely fused, roughly rectangular, with crescentic cutting edge bearing several

irregular calcified teeth (Fig. 11.5D); mandibles in juveniles with much slenderer shafts, fused along distal i

two-thirds, triangular, with cutting edge bearing about 7 to 8 regular, small black teeth (Fig. 11.5E); the latter

visible in adult mandibles as f'me subdistal line.

Parapodia with short presetal lobes and conspicuous postsetal lobes throughout; postsetal lobes broadly 1

triangular and vascularized (Fig. 11.5F, bv) in anterior setigers, narrowing around setiger 15, becoming i

digit•form by setiger 30, pointing laterally and slightly upward, slightly surpassing length of parapodial stem

!

292 I
i

I

! A j

I

I

, I

! - J

!

I °°

I

I

I

'

I Figure 11.5. l_zonbrineris index: A, anterior end, dorsal view; B, posterior end, dorsal view; C, maxillae, dorsal

view, in situ; D, mandibles, adult specimen; E, same, juvenile specimen; F-H, anterior, posterior,

and far posterior parapodium, anterior view; I, compound hooded hook, with detail of crest; J,

I simple hooded hook (G, H after Moore, 1911).

i 293
!

where best developed (Fig. I1.5F-H). Notopodial rudiment present throughout, supported by fascicle of i

several thin, translucent aciculae. Setae very numerous in anterior parapodia, fascicles consisting of about 12

limbate setae and 16 hooks. Hooded hooks in anterior 25 to 33 setigers composite, with long, slender blades •

and small apex carrying about 7 teeth of subequal size (Fig. 11.51). Hood of shaft short, narrow. Simple l

hooks with moderately long hood and main fang surmounted by about 4 small teeth (Fig. 11.5J). Limbate

setae present through setiger 50 to 70, sometimes single fine limbate seta present throughout entire body. •

Aciculae black, in fascicles of about 3 in anterior parapodia, single in posterior parapodia. l

Pygidium short, with four subequal anal cirri.

Remarks. Among species with black aciculae and compound hooded hooks, L. index is easily i

recognized by the postsetal lobes that are prominent throughout and digitiform inmiddle and posterior pampodia.

The dense setal fascicles in anterior setigers are also characteristic. Anterior ends can further be distinguished

from the co-occurring L. californiensis by the number of teeth on MEI (2 for L. index, 1 forL. californiensis), i

i

Type Locality and Type Specimens. California, Monterey Bay, off Point Pifios Lighthouse: holotype

(USNM 17201); off Santa Catalina Island: paratype (ANSP AD1991). i

Habitat. The species lives in silt and green mud in depths between 30 and 1400 m. H

Distribution. Off Southern and central California.

i

Lumbrineris inflata Moore, 1911 i

Figure 11.6 Bi

Lumbrineris inflata Moore, 1911:289-291, pl. 19, figs. 128-132, pl. 20, figs. 133-134.--Hartman, 1944:160;

1968:757.--Fauchald, 1970:89-91, pl. 14, figs. a-d.--Imajima and Higuchi, 1975:20-22, figs. 7a- i

1.--Uebelacker, 1984:41-37, figs. 41-33, 41-34a-g.

Lumbriconereis inflata: Uschakov and Wu, 1965:82-83.

Lumbrineris cervicalis Treadwell, 1922:176.--Uschakov, 1955:239, figs. 79A-D. Fide Hartman, 1959. •

Lumbrineris cingulata Treadwell, 1917:263, pl. 2, figs. 7-12. Fide Hartman, 1959.

Material Examined. California: Santa Maria Basin, offPoint Estero, Sta. 6 (1); off Point San Luis, 1

Sta. 27 (2).

Description. Length to 68 mm, width to 3 mm, segments at least 130. Body long, linear, tapering

toward pygidium (Fig. 11.6A, B). Color in alcohol pale with iridescent cuticle.

I

Prostomium about as wide as long, usually globular, sometimes oval, but not depressed. Maxillae

with strongly incised carriers and long, slender forceps; MII with 4 to 5 teeth, often asymmetrical; Mill with I

3 to 5 teeth, MIV with 2 teeth (Fig. 11.6C, D). Mandibles fused along distal half, with crescentic cutting edge i

bearing 2 irregular teeth on each side of median notch (Fig. 11.6E).

i

Parapodia similar throughout, with short, rounded presetal lobe and moderately long postsetal lobes, •

pointing lateral in anterior parapodia and slightly upward in middle and posterior parapodia (Fig. 11.6F-H). 1

Limbate setae present in first 30 to 40 segments; hooded hooks in first 22 to 26 parapodia compound, with

short blades bearing small main fang and 4 to 8 teeth decreasing in size; hood of blade continuous with that of •

shaft (Fig. 11.6I). Middle and posterior parapodia bearing simple hooks with main fang surmounted by 6 to |

11 fine teeth; hood short, inflated (Fig. 11.6J). Aciculae yellow to amber, usually 2 or 3 per parapodium.

Pygidium small, with four short, equal, conical anal cirri (Fig. 11.6B). i

Remarks. Although L. inflata is most readily recognized by the globular shape of its prostomium,

young specimens of this species are easily misidentified because their prostomium is often bluntly conical

rather than spherical. However, even very small individuals possess more than one tooth on both Mill and i

MIV.

l
I

i

' c

I A _ 4

,I

I

tl

I o

!

I

I _-'_'_-_-_'2.....,._.-.:

!

Figure 11.6. I_mbrineris inflata: A, anterior end, dorsal view; B, posterior end, dorsal view; C, MII, MIII, and

I MIV posterior , dorsal parapodium view; D,, MIantand eriormaxi vielw; laryI, ccompound arders, dorsal hoodviedewhook, ; E, mandibles; setiger 6; FJ,-Hsimpl , firset,hooded median,hook and

(A, B, F-H after Imajima and HJguchi, 1975; C-E, I, J from Imajima and Higuchi, 1975).

I

I 295
!

Type Locality and Type Specimens. Monterey Bay, California, offSanta Cruz Lighthouse: syntypes i

(USNM 16840, USNM 17205).

Habitat. L. inflata lives in mixed, gravelly, sandy, shelly or rocky sediments, i

Distribution. British Columbia to western Mexico; Gulf of Mexico; Bering Sea; Yellow Sea; Japan;

South Africa; intertidal to 130 m.

!

Lumbrinerisjaponica (Marenzeller, 1879)

Figure 11.7 I

Lumbriconereisjaponica Marenzeller, 1879:137-138, pl. 5, fig. 3-3D.--Izuka, 1912:139-140, pl. 14, figs.

Lumbriconereis 17-18. I IatreiUi: Uschakov and Wu, 1962:65-66; 1965:83. Not Audouin and Milne Edwards, 1834.

Lumbrineris latreilli japonica: Hartman, 1944:159-160.

Lumbrineris japonica: Imajima and Hartman, 1964:263-264.--Hartman, 1968:759.--Fauchald, 1970:9 I- 1

92, pl. 14, figs. e-f.--Lissner etal., 1986:D-11.

Material Examined. California: Santa Maria Basin, off Point Buchon, Sta. 13 (1); Tomales Bay, 1

coll. 1959 (2).

Description. Length to 130 mm, width to 6 mm, segments at least 115. Body long, slender, linear, I

almost cylindrical, gently tapering at both ends (Fig. 11.7A, B); segments appearing weakly biannutate due to 1

whitish ring of epidermal cells, perhaps glandular. Color in alcohol pale to tan, iridescent.

I

Prostomium bluntly conical, somewhat depressed, almost as wide as long. Maxillae with laterally •

incised carriers and long, sharply falcate forceps. MII with 4 to 6 teeth, MIII elongate plates with 2 teeth l

(sometimes only a boss on lower edge), MIV large plates with one tooth (Fig. 11.7C, D). MII attached to

carriers with short, thin ligaments. Mandibles fused for almost entire length, with crescentic cutting edge •

bearing a calcified tooth on either side (Fig. ll.7E). 1

Parapodia similar throughout, with short presetal and somewhat elongate postsetal lobes (Fig. 11.7F,

G). Limbate setae present in first 40 to 50 setigers, few per fascicle (Fig. ll.7H); hooks in first 15 to 33 •

setigers compound, with medium long blades (length/width ratio about 3) bearing 8 to 9 f'me teeth above main II

fang; hoods of blades clearly separated from those of shafts (Fig. 11.71). In subsequent setigers hooks becoming

simple, with short hoods and large main fang surmounted by crest of 4 to 8 small teeth (Fig. 11.7J). Aciculae •

dark amber to black, in fascicles of 2 to 4 in anterior parapodia, single in posterior parapodia. 1

Remarks. This species is characterized by having black aciculae and parapodia of similar shape

throughout the entire body; the postsetal lobes are somewhat elongate and nearly reach the length of the •

parapodial stem. Anterior ends can be distinguished from co-occurring L. californiensis by the number of

teeth on MIll (2 for L.japonica, 1 for L. californiensis.)

Type Locality. Eno-Sima, Japan. 1

i

Biology. Gravid females were found between May and October in the Yellow Sea (Uschakov and

Wu, 1965). The species lives in mixed sediments including sand, shells, rocks, andeelgrass roots. i

Distribution. Japan; Indopacific; western Canada to western Mexico; Yellow Sea; intertidal to 75 m. B

!

!

296 I
I

I

C

I

....

I __ i

I

,

I

I

I

I

I

I

I

Eigure 11.7. Lumbrineris japonica: A, anterior end, dorsal view; B, posterior end, dorsal view; C, MII, MIII,

and MIV, dorsal view; D, MI and maxillary carriers, dorsal view; E, mandibles; F, first parapodium,

I anterior view; G, posterior parapodium, anterior view; H, limbate capillary seta, setiger 5; I, compound hooded hook, setiger 1; J, simple hooded hook, setiger 60 (A, B, F, G after Imajima and Higuchi,

1975; C-E, H-J from Imajima and Higuchi, 1975).

I

i 297
!

!

Lumbrineris latreilli Audouin and Milne-Edwards, 1834 !

Figure 11.8

Lumbrineris latreiUi Audouin and Milne Edwards, 1834:168, pl. 38, figs. 13-15.--Hartman, 1944:158-159, I

pl. 9, figs. 213-216; 1968:761 .--Pettibone, 1963:258-260, figs. 67 a-c.--Fauchald, 1970:94-97, pl.

15,figs. f-h.---Orensanz, 1973:359-361, pl. 6.--Imajima and Higuchi, 1975:32-36, fig. 13.--Ramos, •

1976:121-124, figs. 16-18. I

Lumbrinconereis latreilli: Fauvel, 1923:431, figs. 171 m-r; 1953:266, figs. 134 m-r.--Okuda, 1946:120, pls.

12-13. I

Lumbrinereis latreilli: Berkeley and Berkeley, 1948:98, figs. 154-156.

Description. Length to 300 ram, width to 5 mm, segments at least 120. Body robust; color in alcohol •

pale to brown with iridescent cuticle.

Prostomium conical, slightly longer than wide, depressed (Fig. 11.8A). Maxillae with broad, laterally

incised carriers and strong forceps; MII with 4 to 6 teeth, often asymmetrical, MIII with 2 teeth, MIV with

single tooth (Fig. 11.8B, C). Mandibles large, roughly rectangular, fused except for small proximal part, with

crescentic cutting edge (Fig. 11.8D).

Parapodia similar throughout, with low presetal lobes and conical postsetal lobes, in posterior setigers I

directed slightly upward (Fig. 11.8E-G). Limbate setae present in anterior and middle segments, sometimes Ill

singly in posterior segments (Fig. ll.8H); hooded hooks in first 11 to 24 segments composite, with blades I

about 5 times as long as wide, bearing small main fang surmounted by 6 to 9 fine teeth; hood of blade separate

from that of shaft (Fig. 11.8I). Hooded hooks in subsequent setigers simple, much stouter than compound ID

hooks, with large main fang and 9 to 10 smaller teeth, hoods short (Fig. 11.8J). Aciculae yellow to amber, in

fascicles of 3 anteriorly, only 2 in posterior segments; aciculae occasionally occurring singly in a parapodium. I

n

Pygidium with 4 equal, short anal cirri (Fig. 11.8K).

Remarks. Within the genus Lumbrineris, L. latreiUi appears to be one of the most variable species. •

Ramos (1976) explored the ranges of variability for several characters, such as the number of teeth on MII |

and the length of the blades of composite hooks relative to the blade width and the length of the shaft. She

compared two collections from the Catalonian coast (Mediterranean Sea) and concluded that the species •

could not b'e split into two even though slight differences between the two populations were found. |

Type Locality. France.

Biology. Hartman (1944) found gravid females tightly packed with salmon colored eggs about 220

im in diameter present in all segments but the fu'st and last 20 to 30. Eggs are laid in gelatinous masses and

attached to macroalgae. Development is direct; larvae leave the egg masses at the 4- to 7-setiger stage

(Pettibone, 1963). L. latreilli occurs in mud and sand, among eelgrass roots, and under rocks.

Distribution. Cosmopolitan; Greenland to France; Mediterranean Sea; eastern Canada to North

Carolina; western Canada to California; Mexico, Peru; Galapagos Islands; Gulf of Mexico to Panama; Japan;

Indian Ocean, Red Sea, Persian Gulf; West and South Africa; intertidal to 2300 m.

!

!

I

Lumbrineris limicola Hartman, 1944 I

Figure 11.9

Lumbrineris timicola Hartman, 1944:161-162, pl. 11, figs. 230-237; 1968:765.--Fauchald, 1970:97-98, pl. m

16, figs. a-d.--Hyland et al., 1990:F-1. []

Material Examined. California: Santa Maria Basin, off Point Sal, Sta. R-8 (1). m

Description. Length to 84 ram, width to 6 mm, segments to 81. Body robust; color in alcohol pale m

with iridescent cuticle. I1

I

Prostomium bluntly conical, as long as wide, depressed (Fig. 11.9A). Maxillae with large, laterally

incised maxillary carriers about as long as MI; MII with 4 teeth, MIH unidentate, sometimes with small boss •

on cutting edge resembling a reduced second tooth; MIV with single heavy tooth (Fig. 11.9B). Mandibles 1

fused along anterior half, shafts posteriorly diverging, wide, approaching shape typical for Augeneria (Fig.

11.9C). []

Parapodia conspicuous from setiger 1, with notopodial rudiments; presetal lobes low, rounded n

throughout, postsetal lobes becoming elongate and pointing dorsad in posterior setigers (Fig. 11.9D-F).

Compound hooded hooks and simple limbate setae present through 25-40 setigers in adult specimens; shafts •

of hooks 3-4 times as long as wide, with crest of about 8 subequal teeth (Fig. 11.9G). Simple hooks stouter 1

than composite ones, with main fang surmounted by about 8 small teeth (Fig. 11.9H). Aciculae pale yellow.

Remarks. This species may be much more common than suggested by the small number of records m

off California. It is likely that anterior ends have been misidentified asL. latreilli, a "cosmopolitan" species

with few unmistakable diagnostic characters. It is suggested that only specimens with a distinctly bidentate

MIII be assigned toL. latreilli, whereas individuals with a unidentate or indistinctly bidentate MHI should be m

identified as L. limicola.

Type Locality and Type Specimens. California, off Point Fermin, California: holotype (LACMAHF Poly 49). n

1 Habitat. Mud and silt.

Distribution. Central California to western Mexico, shallow subtidal to about 190 m. m

m

Genus Ninoe Kinberg, 1865

1

Type Species. Ninoe chilensis Kinberg, 1865 m

Diagnosis. Prostomium conical, usually with longitudinal slits or folds. Parapodia with branchiae m

arising from postsetal lobe, with 2 to about 15 digitiform lobes. Setae including simple limbate capillaries and

simple hooded, multidentate hooks. Aciculae dark amber to jetblack, setae usually with dark bases and

amber-colored to clear tips. MIV and often also MHI with 1 to 3 teeth and serrated cutting edge; MV absent. 1

I

Remarks. Kinberg (1865) established the genus (and a family Ninoidea) to accommodate eunicemorph

forms with no prostomial appendages or tentacular cirri, but with branchiae arising from the postsetal lobes 1

of a limited number of anterior parapodia. He included three new species from off Chile and Brazil: N. 1

chilensis, N. brasiIiensis, and N. oculata. Fauchald (1970) emended the genus to include not only species II

with gills on postsetal lobes of anterior parapodia, but also those with gills on posterior parapodia and the

adjacent body wall. This emendation was rejected by Orensanz (1973, 1990) because these different branchial m

structures are not considered homologous. The definition of Ninoe, as originally stated by Kinberg, is followed 1

here. The presence of serrations on the cutting edges of MIV, and often MIII, is an additional reliable

character for discrimination of this genus, n

300 m
Figure 11.9. Lumbrineris limicola: A, anterior end, dorsal view; B, maxillae, dorsal view, in situ;

C, mandibles; D-F, anterior, median, and posterior parapodium, anterior view; G, simple

hooded hook; H, compound hooded hook, with detail of crest (C-F after Hartman,

1944; G from Fauchald, 1970).

i 301
!

!

II

Ninoe gemmea Moore, 1911 •

Figure 11.10

Ninoe gemmea Moore, 1911:283-285, pl. 19, figs. 101-109.--Lissner et al., 1986:D-12. i

Material Monterey Bay, Sta. 4450 (holotype); Santa Mafia Basin, off Point gExamined. California:

Sal, Sta. 38 (1); off Purisima Point, Sta. 42 (1).

i

Description. Length to 104 ram, width to 2.5 mm excluding parapodia, segments to 146. Body i

slender, linear, slightly wider in branchial region than in postbranchial region. Color in alcohol tan to dark l

grey, with highly iridescent cuticle.

Prostomium slightly longer than wide, conical, somewhat depressed dorsoventrally, with 2 distinct •

nuchal organs at posterior margin (Fig. 11.10A); dorsal surface smooth, ventral side with 2 longitudinal

furrows stretching from the insertion of the large palps to nearly tip of prostomium. Two peristomial rings

developed dorsally, partly fused ventrally to form crenulated lower tip (Fig. 11.10B). Maxillae with slender, i

laterally incised carriers about as long as MI; MII with 8 teeth, MHI and MIV unidentate with serrated cutting I!

edges (Fig. 11.10C). Mandibles slender, proximally fused, with irregularly serrated cutting edge (Fig. 11.10D).

Parapodia small in first five setigers, then increasing; presetal lobe obliquely truncate, postsetal lobe i

much longer, conical to foliaceous (Fig. 11.10E-G). Gills first present on setigers 4 (holotype) to 6, starting

as 2 nearly equal halves of the postsetal lobe (Fig. 11.10H); upper branch increasing in length and pointing

slightly upward in subsequent setigers, lower branch dividing into maximally 4 digitiform branches that are i

somewhat shorter and narrower than the upper lobe (Fig. 11.10I, J). Number of branches decreasing gradually

after setiger 39 to 46, single gill after setiger 50, rudimentary process on postsetal lobe through about 10

additional setigers. Setae including limbate capillaries and simple hooded hooks throughout entire length of •

body; hooks slender with very small apex and long hood in anterior setigers, wider with distinct apex bearing []

crest of several teeth over main fang and short, rounded hood in posterior setigers (Fig. 11.10K-M). Aciculae

black, setae clear with dark bases, i

[] Remarks. Moore, in his original description, reported the gills starting from setiger 5 or 6; however,

an examination of the holotype revealed that this specimen has gills from setiger 4. Hartman's (1968) notion

as to where the gills fit-stappear is incomplete and easily misunderstood, because she only mentioned the first

appearance of the second branchial filament (setiger 8),but not of the first filament (setiger 4-6). The distribution

of the gills along the body should be used with caution for species discrimination because it may have a wider

range of variability than originally thought; however, slow increase of the number of branchial filaments []

along the length of the branchial region distinguishes N. gemmea from the co-occurring N. palmata (see

below).

Type Locality and Type Specimens. California, Monterey Bay, off Point Pifios Lighthouse: holotype i

(USNM 17348).

Habitat. Soft green or dark mud and sand.

Distribution. Central and southern California, 79 - 1272 m. i

!

!
i Figure 11.10. Ninoe situ; D,gemmea mandibles; : A, E-I, anterior first enthrough d, dorsal fifthview; parapodium, B, same, anterior ventral view; view; J,C,25th maxill parapodium, ae, dorsal view, anteriorin

view; K, hooded hook, setiger 25; L, same, setiger 100; M, limbate capillary seta, setiger 25 (A, B,

i E-I from holotype; D, K-M after Moore, 1911).

i 303
!

!

Ninoe palmata Moore, 1903 1i

Figure 11.11

Ninoe palmata Moore, 1903:456-457, pl. XXVI, figs. 69-71 .--Izuka, 1912:137-139.--Imajima and Hartman, N

1964:264.--Imajima, 1968:141.--Uschakov and Wu, 1965:83-84, fig. 21H-M.--Imajima and

Higuchi, 1975:10-12, fig. 3a-m.--Lissner etal., 1986:D-12. am

Ninoe nr. palmata: Lissner et al., 1986:D-12. |

Ninoe nr. gemmea: Lissner et al., 1986:D-12.

Material Examined. Japan: Sendal Bay, off Honshu Island, Sta. 3767 (holotype). California: Santa I

Mafia Basin, off Purisima Point, Sta. 42 (1), Sta. 43 (1), Sta. 53 (1); off Point Arguello, Sta. 67 (1).

Description. Length to 51 mm, width to 2 mm excluding parapodia, segments to 158. Body slender, !

widest in branchial region, gently tapering at both ends. Color in alcohol light tan. |

Prostomium conical, slightly longer than wide, with paired nuchai organ at posterior margin (Fig.

ll.llA). Maxillae with slender, laterally incised carders about as long as MI; MII with 6 or 7 teeth, the m

subterminal one tiny; MIII and MIV unidentate; MIII with whitish center, MIV with serrated cutting edge

(Fig. 11.11B, C). Mandibles poorly sclerotized, delicate, fused along proximal half, with 2 large, irregular

teeth on cutting edge (Fig. 11.11D). I

1

Parapodia minute in first 2 setigers, with obliquely truncate presetal lobe and conical to foliaceous

postsetal lobe, increasing in length in subsequent setigers (Fig. 11.11E). Gills starting on setigers 2 to 4, first lit

developed as 2 subequal halves of postsetal lobe; lower half splitting into additional digitiform branches, el

number of branches increasing quickly to 6 within about 15 setigers; upper half increasing in size, becoming m

about twice as long and distinctly thicker than lower branches where best developed, pointing dorsad and I

curved across dorsum (Fig. 11.11F). Last gill between setigers 30 and 43; postsetal lobes in postbranchial •

region pointed, conical, distincly longer than presetal lobes (Fig. 11.11G). Setae including limbate capillaries mar

and hooded hooks throughout; anterior hooks slender, with small crest of about 5 teeth and long, narrow hood

(Fig. 1011H, I). Posterior hooks stouter, with distinct crest of about 8 teeth over small main fang and shorter, n

wider hood (Fig. 11.11J). Aciculae dark amber, setae clear, sometimes with amber colored bases. BII

Pygidium with 2 ventrally inserted anal cirri (Fig. 11.11K). all

Remarks. Moore's description ofN. palmata is unusually short, especially with regard to the branchial 1

development, and examination of the holotype revealed that it is also incorrect. He states that the gills start

"as far forward as V," whereas they were found to first appear on setiger 2 on one side of the holotype and on li

setiger 3 on the other side; the maximum number of filaments is 6 ratherthan 5 as stated by Moore. Specimens n

originally identified as Ninoe nr. gemmea and N. nr. palmata have been included in N. palmata even though

the gills start in setigers 4 or 5 rather than 2 or 3. It seems that the first appearance of gills usually varies by II

about two or three in other species of this genus, so that the assignment to a new species seems unjustified. All 1

other characters agree well with N. palmata.

Type Locality and Type Specimens. Japan, Sendal Bay, off Honshu Island, 30-35 m: holotype •

(USNM 15735). 1

Habitat. Sandy silt and shells, mixed sand and silt.

Distribution. Japan; Yellow Sea; central California; 30 - 1450 m. n

!
I

!

I

I __ _°_

| _ _ l

!

"

I

Figure 11.11. Ninoe palmata: A, anterior end, dorsal view; B, MI and maxillary carriers, dorsal view; C, MII,

I Mill, and MIV, dorsal view; D, mandibles; E-G, first, 16th, and 75th parapodium, anterior view; H,

limbate capillary seta, setiger 75; I, hooded hook, anterior setiger; J, same, setiger 75 (all from

Imajima and Higuchi, 1975).

!

I

I 305
!

Ninoe tridentata Hilbig, new species I

Figure 11.12

i

Ninoe sp. A Lissner et al., 1986:D-12.--Hyland et al., 1990:F-1. I

Material Examined. California: Santa Maria Basin, off Point Sal,Sta. R-8, 34°55.30'N, 120°45.87'W, i

90 m: holotype (USNM 170425) and 3 paratypes (USNM 170426); Sta. 30, 34°54.19'N, 120°47.0TW, 98 m:

1 paratype (SBMNH 142696); off Point San Luis, Sta. 21, 35°06.11'N, 120°44.82'W, 49 m: 1 paratype !

(LACM-AHF Poly 1658). !1

Description. Length to 13 mm, width to 1 ram, setigers to 96. Body slender, gently tapering at both

ends, widest in pharyngeal region. Color in alcohol light tan. II I

Prostomium conical, almost twice as long as wide, with faint dorsal longitudinal folds, much narrower

than peristomium (Fig. 11.12A). Maxillae with laterally incised carriers, shorter than MI; MII with 7 theeth,

MIII with three conical teeth, MIV unidentate with finely serrated cutting edge (Fig. 11.12B). Mandibles I

delicate, weakly sclerotized. II

Parapodia in first two setigers very small, with short, rounded presetal and longer conical, pointed /

postsetal lobes (Fig. 11.12C, D); subsequent parapodia increasing in size, bearing branchiae from setiger 3 to II

about 45, with maximally 11 filaments (holotype with up to 7 filaments) and foliaceous upper filament of II

slightly greater length than lower filaments (Fig. 11.12E-I). Simple capillaries with wide wings and long

filiform tip, in postbranchial setigers gradually turning more slender with wing tapering more gradually, 1

absent from far posterior setigers (Fig. 11.12.1, K). Hooks fu'st present from setigers 4 to 14, in anterior 1

setigers slender, with medium long hood and small apex bearing crest of about four teeth above main fang;

hoods gradually shortening and widening, hooks becoming stouter, bearing larger main fang and crest of five I

subequal teeth (Fig. 11.12L, M). Aciculae dark amber, setae clear with amber colored bases.

Remarks. Ninoe tridentata is easily recognized by the elongate prostomium and the absence of /

hooded hooks from at least three anterior parapodia. The species resembles N. palmata in the distribution and I

morphology of the gills, but can be distinguished by the tridentate, rather than unidentate, MIII.

u

Etymology. The epithet refers to the tirdentate thrird maxillae, iI

Habitat. Sand and sand mixed with silt. II

Distribution. Presently known only from the Santa Maria Basin.

Ninoe sp. C II

Ninoe sp. C Lissner et al., 1986:D-12. .i

Material Examined. California: Santa Maria Basin, off Point Arguello, Sta. 69 (8). I

1

Description. All available specimens juvenile; length to 10 mm, width to 0.5 mm, setigers to 45.

Body very slender, with prominent parapodia and gills; color in alcohol whitish. Prostomium conical, slightly I

longer than wide, with indistinct nuchal organs. Maxillae soft, with laterally incised carriers about as long as II

MI; MII with 4 or 5 teeth, MIII and MIV with serrated cutting edge and one or two teeth (jaws too small for

accurate observation). Parapodia conspicuous throughout, with short, bluntly rounded presetal lobes and

digitiform, much longer postsetal lobes; branchiae present from setiger 2 to maximally 17 (usually 15), with II

up to 6 filaments and digitiform to foliaceous upper filament, up to twice aslong as lower f'daments where best

m

developed. Setae including limbate capillaries and hooded hooks; capillaries with broad wings in anterior

I
l

I

I

tl'I

I B1 ', /

I _\iI ,,, i_i'

•I _ !i/ _Zi

i:"i""": : ...... ' K

l _,,,_ ,__L_

, ..., ,:

I D ,".__\ :i:!" '/ __ P_'OSt L H

III E__,, -,,_-?i _ G _ I

!

Figure 11.12. Nmoe _n_ Itilbig, new species: A, anterior end, dorsal view; B, maxillae, dorsal view, in

sin,; C-G, In'st, second, third, 10t.h, and 25_ parapodium, anterior view; H-I, second-to-last and

I gill-bearing parapodium, view; J, capiUary seta, setiger; K, same,

last anterior limbate anterior

posterior setiger; L, hooded hook, anterior setiger; M, same, posterior sefiger. (A, C-I: paratype

USNM 170426; B, J-M: holotype).

!

i 307
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!

setigers, much more slender and with long, narrow wings in postbranchial setigers, absent after setiger 35 to

40; hooks present from setiger 1, very slender with moderately long hoods and small apex in anterior setigers, II

stouter, with short hoods and large main fangs in posterior setigers. Crest of about 5 teeth forming sharp

m

angle with shaft in hooks from posterior setigers, resembling hooks ofLumbrineris latreilli. m

Remarks. Specimens assigned toN•hoe sp. C are most likely juveniles of N. long•branchia Fauchald, i

1972, a species found in depths between 1800 and 2600 m off western Mexico. The station where N. sp. C

was found is about 1000 m deep and may be at the shallow end of the distribution of N. long•branchia. The i

only difference betweenN, long•branchia andN. sp. C is the number of teeth on MII (4 or 5 forN. sp. C, 6for _l

N. long•branchia) and possibly the shape of the maxillary carders that are reported to be straight in N.

long•branchia, but laterally incised in N. sp. C. Examination of the holotype of N. long•branchia will be m

necessary to confirm the assignment ofN. sp. C. []II

Ninoe sp. D I

Ninoe sp. D Lissner etal., 1986:D-12.

I

Material Examined. California: Santa Maria Basin, off Point Arguello, Sta. 58 (1).

Description. Only one incomplete specimen found during entire program in the Santa Maria Basin E

and Santa Barbara Channel. Length 19 mm, width 1 mm, 86 setigers; color in alcohol whitish. Prostomium J

conical, pointed, longer than wide, distinctly narrower than peristomium. Maxillae with very slender, laterally

incised carriers, longer than MI; MII with 5 teeth, MIII with 1 tooth and rounded boss, MIV unidentate with I_

serrated cutting edge. Branchiae present from setiger 3, maximally 6 Fdaments plus superior lobe, the latter 1

foliacous and somewhat longer than lower filaments. Last branchia on setiger 41. Limbate capilIaries present

throughout length of fragment, with broad wings in anterior setigers and narrow wings in postbranchial •

setigers; hooded hooks from setiger 15, narrow, with moderately long hoods and very small apex bearing I

about 6 f'me,subequal teeth, gradually becoming stouter with hoods shortening in posterior setigers of fragment.

Aciculae dark brown, setae clear with dusky bases. I

Rema | rks. Based on the softness of the maxillary plates, it can be assumed that this specimen is a

juvenile; no final assignment ofNinoe sp. D, either to a known or a new species, could be made.

Genus Scoletoma Blainville, 1828, emended Frame, 1992 I_

Type Species. Lumbricusfragilis O.E MiiUer, 1776 I

Diagnosis. Prostomium conical to globular. Setae including simple limbate capillaries and simple, []

multidentate hooded hooks. Aciculae yellow or black. MII about as long as MI, linked to MI with threadlike II

ligament. MV, if present, free from MIV.

Remarks. Blainville erected Scoletoma for Lumbricus fragilis O.E Miiller without stating which l

characters he considered diagnostic for his new genus. Frame (1992) resurrected Scoletoma and provided a

generic definition based on the above characters. Although the jaw apparatus is very much like that of

Lumbrineris, the separation of the two genera based on setal morphology is acceptable because the presence l

or absence of compound hooks is consistent within all other lumbrinerid genera.

!

308 |
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I Scoletoma tetraura (Schmarda, 1861)

I Figure t 1.13

i NLumbriconereis otocirrus tetraurus tetrauraSchmarda, : Ehlers, 1861:1 1901:137-139, 17, 6 figs.pL 17, figs. 1-10 (in part).

Lumbrinereis tetraura: Day, 1953:435-436.

i Lumbrineris tetraura: Moore, 1911:291 .--Augener, 1933:61-62.--Hartman, 1944:147-149, pl. 8, figs. 175, 190, 191, pl. 9, figs. 192-195.

Lumbrineris brevicirra Monro, 1933:83 (in part).

I Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. 21 (1), Sta. 23 (1), Sta.

R-2 (19); Western Santa Barbara Channel, Sta. 79 (2); Tomales Bay, coll. 1958 (1).

! B Description. Length to 250 mm, width to 5 ram, segments to 280. Body slender, with relatively long

parapodia in posterior half; color in alcohol pale, cuticle iridescent.

Prostomium conical, about as long as wide, rounded anteriorly (Fig. 11.13A). Maxillae with short,

I wide, laterally incised maxillary carriers; MII with 4-5 teeth and large aliform expansion, M//I bidentate, MIV large, roughly triangular, unidentate plates, MV large, squarish plates, free from MIV (Fig. II.13B).

Mandibles relatively short, fused for most of their length (Fig. 11.13C).

I Parapodia conspicuous from setiger 1, with short, truncate presetal lobes and conical postsetal lobes;

posterior parapodia much longer than anterior ones, with elongate, erect postsetal lobes pointing dorsad, at

least three times as long as presetal lobes, often irregularly wrinkled (Fig. 11.13D, E). Simple hooded hooks

I from 1 anterior hooks with hoods and small of several present setiger slender, long crest free teeth, posterior

hooks stouter, with short rounded hoods and crest of numerous small, subequal teeth above a slightly larger

main fang (Fig. 11.13F, G). Limbate setae present through about 40 to 60 setigers. Aciculae yellow to amber,

I usually in fascicles of 2 or 3 parapodium.

present per

Pygidium small, with 2 pairs of short anal cirri.

I hooks that Remaallowrks. forScoletoma identification tetraurawithout is oneexamining of the fewthespecies jaws. that Thispocssess haracterverymaycharabecteristi helpful c hoo for dedthe

identification of small specimens where dissection of the jaws is difficult. It also seems that the second tooth

I on MIII Type is not Locality present in small individuals (2 to 3 cm length). . Chile.

Biology. Orensanz (1973) reported that gut contents consisted of a variety of algae, detritus, sediment,

I small crustaceans and pelecypods, and diatoms, suggesting an omnivorous life style. Scoletoma tetraura

appears to be equally flexible with regard to substatum; it has been reported from mud to sand, mixed sediments,

and rocks.

I Distribution. Central California intertidal 220

to Chile, m.

!

!

| 309
I

I

I

I

ii '

!

I.

I

i,_ I

; I

I

I

Figure 11.13. Scoletoma tetraura: A, anterior end, dorsal view; B, maxillae, dorsal view; in situ; C, mandibles; i

D, 10th parapodium, anterior view; E, posterior parapodium, anterior view; F, hooded hook, anterior

setiger, with detail of crest; G, same, posterior setiger (C-G after Orensanz, 1973). i

i

31o |
!

!

Literature Cited

!

Audouin, J.V. and H. Milne Edwards. 1833. Classification des Ann_lides, et d6scription de celles qui habitent

1 les c6tes de la France. Annales de Sciences naturelles Paris, s6rie 1, vols. 28-30.

I Audouin, J.V. and H. Milne Edwards. 1834. Recherches pour servir _ i'histoire natureUe du littoral de la

France, ou Recueil de m_moires sur l'anatomie, ta physiologie, la classification et les mocurs des

I anim pt. Paris, aux de290nopp., s crtes; 8pls.ouvrage accompagn6 de planches faites de apr_s nature. Vol. 2. Annrlides. 1

i Augener, H. 1933. Polychaeten von den Galapagos-Inseln. The Norwegian Zoological Expedition to the Galapagos Islands 1925, conducted by Alf Wollebaek. VI. Nytt Magasin for Naturvidenskapene

Oslo 73:55-66, 1 fig.

I Berkeley,Fishe E. anriesd C. Research Berkeley. Board1948. Canada: Annel1-100, ida, Polych 160 aeta errantia. Canadian Pacific Fauna, No. 9b (1). figs.

Bhaud, M. and C. Cazaux. 1987. Description and identification of polychaete larvae; their implications in

I current biological problems. Oceanis 13(6):596-753.

Blainville, H. de. 1828. Dictionaire des sciences naturelles. Vol. 57:368-501.

I Day, J.H. 1953. The polychaet fauna of South Africa. Part 2. Errant species from Cape shores and estuaries.

Annals of the Natal Museum 12:297-441.

I Ehlers, E.Festschrift 1901. DiezurPolychaeten Feier des hundertj_hrigen des magellanischen Bestehens und chilenischen der k6niglichenStrandes. Gesellschaft Ein faunistischer der Wissensch Versuch. aften

zu Grttingen. (Abh. Math.-Phys.). Berlin. pp. 1-232, pls. 1-25.

I Fauchald, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae,

Lysaretidae and Dorvilleidae from western Mexico. Allan Hancock Monographs in Marine Biology

5:1-335.

I Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adjacent areas

in the eastern Pacific Ocean. Allan Hancock Monographs in Marine Biology 7:1-575.

I Fauchald, polychaete worms, keys to orders, and genera. Natural

K. 1977. The Definitions and the families

History Museum of Los Angeles County, Science Series 28:1-190.

I Frame, A.wiBth. 1de99sc2.ripThe tionslumbr of ainneweridsge(nAnn us eanliddat:woPolynewchaespecie ta) coll s. ecPtredoceedin tinwgosnoofrthwestern the BiologAtlan ical ticSocie survtyeysof

Washington 105(2): 185-218.

I Fauvel, P. 1923. errantes. Faune de France, Paris, vol. 5:1-488. Polych_tes

Fauvel, P. 1953. Annelida Polychaeta. The Fauna of India, 507 pp.

I O. 1944. annelids. Part V.Eunicea. Allan Hancock Pacific 10:1-237.

Hartman, Polychaetous Expeditions

Hartman, O. 1949. A new marine annelid from Florida. Proceedings of the U.S. National Museum 99:503-

il i 508.

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Hartman, O. 1968. Atlas of theErrantiate Polychaetous Annelids from California. Allan Hancock Foundation, I

University of Southern California, Los Angeles. 828 pp.

Hyland, J. and J. Neff. 1988. California OCS Phase II Monitoring Program. Year-One Annual Report. Vols. n

i and II.Prepared for the U.S. Department of the Interior, Minerals Management Service, Los Angeles,

CA. OCS Study, MMS 87-0115.

m

Hyland, J., E. Baptiste, J. Kennedy, J. Campbell, R. Kropp, C. Robinson, and S. Williams. 1990. Macroinfaunal I

Assemblages in the Santa Mafia Basin offthe Coast of Southern California. Chapter 7 In: California

OCS Phase II Monitoring Program. Year-Three Annual Report (M. Steinhauer and E. Imamura, n

eds.). Prepared for the U.S. Department of the Interior, Minerals Management Service, Pacific OCS II

Region, under contract No. 14-12-0001-30262.

Imajima, M. 1968. Potychaetous annelids from Hayama, Miura Peninsula. Science Reports of the Yokosuka I

City Museum 14:20-41, 6 pls. [in Japanese]. in

Imajima, M and O. Hartman. 1964. The polychaetous annelids of Japan. Part 2. Allan Hancock Foundation m

Occasional Papers 26:239-452, pls. 36-38. II

Imajima, M. and M. Higuchi. 1975. Lumbrineridae of polychaetous annelids from Japan, with descriptions

of six new species. Bulletin of the National Science Museum Tokyo, Series A i (1):5-37. n

n,

Izuka, A. 1912. The errantiate Polychaeta of Japan. Journal of the College of Sciences, Tokyo University

30(2):1-262, 24 pls. n

II Kielan-Jaworowska, S. 1966. Polychaete jaw apparatuses from the Ordovician and Silurian of Poland and

a comparison with modern forms. Paleontologia Polonica 16:1-152.

m

Kielan-Jaworowska, S. 1968. Scolecodonts versus jaw apparatuses. Lethaia 1:39-49. I

Kinberg, J.G.H. 1865. Annulata nova. Ofversigt af K. Vetenskaps-Akademiens Forhandlingar 21:559-574.

m

Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bernstein, R. Cimberg, T. Kauwling, and W. Anikouchine. I

1986. Assessment of long-term changes in biological communities of the Santa Maria Basin and

Western Santa Barbara Channel - Phase I. Report prepared for the U.S. Department of the Interior, m

Minerals Management Service, Pacific OCS Region, under contract No. 14-12-0001-30032. l

Marenzeller, E. v. 1879. Siidjapanische Armeliden. Iund II. Denkschrift der Akademie der Wissenschaft zu

Wien 41(2):109-152, pls. 1-6. N

Monro, C.C.A. 1933. The Polychaeta Errantia collected by Dr. C. Crossland at Col6n in the Panama region

and the Galapagos Islands during the expedition of the S.Y.St. George. Proceedings of the Zoological B

Society of London pt. 1:1-96, 36 figs. |

Moore, J.P. 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. Proceedings

of the Academy of Natural Sciences Philadelphia 55:401-490, pls. 23-27. m

Moore, j.P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of Southern

California in 1904. 3. Euphrosynidae to Goniadidae. Proceedings of the Academy of Natural Sciences n

Philadelphia 63:234-318, pls. XV-XXI. |

Okuda, S. 1946. Studies on the development of the Annelida Polychaeta. I. Journal of the Faculty of

Sciences, Hokkaido Imperial University, new series, 9(2): 115-219, 17 pls., 33 figs. i

Orensanz, J.M. 1973. LOs anelidos poliquetos de la provicia biogeografica Argentina. IV. Lumbrineridae.

Physis Sec. A 32(85):343-393. m

312 m
!

I Orensanz, J.M. 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas. With

addenda to the Eunicemorpha of Argentina, Chile, New Zealand, Australia, and the southern Indian

i Ocean. Biology of the Antarctic Seas XXI. Antarctic Res. Ser. 52:1-183. Perch, D.A. 1986. Selective deposit-feeding byLumbrineris cf. latreilli (Polychaeta: Lumbrineridae), with

a new method for assessing selectivity by deposit-feeding organisms. Marine Biology 93:443-448.

I Pettibone, M. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through

Trochochaetidae. United States National Museum Bulletin 227(1 ): 1-356.

I Ramos, J.M. 1976. Lumbrineridae (Polych_tes errantes) de Mgditerran6e. Annales de l'Institut OcEanographique 52(1 ):103-137.

i Rioja, E.Mexico. 1962. Estudios Anales anelidologicos XXVI. Algunos Anelidos Poliquetos de las costas del Pacifico de del Instituto Biologia de la Universidad Nacional de Mexico 33(1-2):131-229.

Schmarda, L.K. 1861. Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853

I bis 1857. Leipzig, Vol. 1. Turbellarien, Rotatorien und Anneliden. Pt. 2:1-164, 22 pls.

Treadwell, A.L. 1917. Polychaetous annelids from Florida, Porto Rico, Bermuda, and the Bahamas.

i Publications of the Carnegie Institute Washington 251:255-272, 3 pls. Treadwell, A.L. 1922. Polychaetous annelids collected at Friday Harbor, State of Washington, in February

and March, 1920. Publications of the Carnegie Institute Washington 312:171 -181, 37 figs.

I Treadwell, A.L. 1929. Lumbrineris bicirrata, a new polychaetous annelid from Puget Sound. American

Museum Novitates 338:1-3, 7 figs.

I Uebelacker, J.M. 1984. Family Lumbrineridae Malmgren, 1867. Chapter 41 In: J.M. Uebelacker and P.G. Johnson (eds.): Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Volume VI.

Barry A. Vittor & Associates, Inc. 7 vols.

I Uschakov, P.V. 1955. Polychaeta of the Far Eastern Seas of the SSSR. Akademia Nauk SSSR 56:1-445,

figs. 1-164. [in Russian]. (Translated by Israel Program for Scientific Translation, Jerusalem, N.

i Uschakov,1259, P.V.419andpp., B.L.1965. Wu. NTIS 1965.No.Polychaeta TI" 65-50020). Errantia of the Yellow Sea. Issledovaniya Fauna Morei

3(11):I45-258. [in Russian] (Translated for the Smithsonian Institution and the National Science

I Foundation by Amerind Publishing Co. Ltd., New Delhi. viii + 237 pp, 1979.)

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I 12. FAMILYOENONIDAEKINBERG, 1865,

 lvmSOWO rUNSANZ1990 ,

I (including Arabellidae Hartman, 1944)

I by

I Brigitte Hilbig'

I Introduction

I The oenonids superficially resemble the lumberinerids, considerably characters,

but differ in several

some of which are easily observed. The most obvious difference is the absence of hooded hooks in oenonids,

so that only one type of seta is present (limbate capillaries). In many species, these limbate setae are strongly

I and serrated covered with surficial oenonids have characteristically thin and geniculate or

spines. Many

a

wiry habitus; the cuticle is very rigid and often strikingly iridescent. The prostomium of oenonids is generally

dorsoventrally depressed, whereas the typical lumbrinerid prostomium is conical or even globular, although it

I flatten, in individuals, when the is everted. The most difference ITlay especially large proboscis important

between lumbrinedds and oenonids, from an evolutionary and systematic point of view, is the arrangement of

the jaw apparatus (see below). Oenonids are distributed worldwide, but never abundant; some species are

I parasitic, at least through part of their life history, in other polychaetes, bivalves, and echiurans.

I Morphology

I The prostomium is conical, usually dorsoventrally flattened, and without any appendages except for

Oenone which has three minute antennae atthe posterior margin of the prostomium. Eyes may be present at

the posterior prostomial margin as well. The eversible pharynx is armed with five pairs of Jmaxillary plates

I and mandibles that often small in relation to the maxillae, carriers much paired ale very Maxillary are longer

than the maxillary plates and consist of a pair of slender dorsal pieces and a usually shorter and broader single

ventral piece. The maxillary plates (MI through MV) may be symmetrical, i.e., the corresponding plates are

I of equal size and have similar numbers of teeth, or they be asymmetrical, i.e., corresponding plates

may may

be of very different sizes and have clearly different numbers of teeth. An excellent explanation of the origin

and evolution of the different maxillary arrangements can be found in Orensanz (1990). The first maxillae are

I variable in shape; they may be falcate and proximally smooth, much like MI in lumbrinerids, or proximally

dentate; some genera have MI that are entirely dentate and have a similar shape to the MII. Intermediate types

of MI can be found that have a dentate border over most of their length, but a distinctly larger and hookI shaped distal tooth. The second maxillae axe similar in size to the MI and usually bear about 3 to 10 teeth; they are located ventral of the MI and free from the latter (not attached with ligaments as in lumbrinerids). In

asymmetrical jaw apparatuses, a smaller MI on one side is accompanied by a larger MII. The third, fourth,

I

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i 315
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and fifth maxillae are smaller than the first and second ones and located in front of the MI. They are developed I

as multi- or unidentate plates and usually well sclerotized; MV may be absent in some species. Parasitic

species have often greatly reduced maxillae; Biborin has no maxillae at all. The mandibles are comparatively n

small, rod- to diamond-shaped, simple plates that are free from one another. Mandibles are absent in one l

species of Drilonereis.

Because of the paucity of external features, the arrangement of the jaw pieces is one of the most I

important taxonomic characters in this family. For examination, the pharynx should always be dissected from

the dorsum. It may often be useful to dissect out the whole pharyngeal bulb and place it on a slide to view the

more delicate parts, such as the unpaired carrier or the mandibles. However, the morphological details il i

especially of MI and MII are size-dependent, and juveniles should be assigned to a species with caution I

(Colbath, 1987).

The peristomium consists of one or two apodous rings, with the anterior one forming the lower lip on 1

the ventral side. All setigers have essentially the same width as the peristomium; the body tapers only within U

the last few segments. Parapodia are subbiramous, with the notopodium reduced to a bundle of slender

notoaciculae and a minute papilla, except for Oenone where the notopodium is developed into a foliaceous n

lobe extending well beyond the neuropodium. The neuropodium has a short presetal and a typically conical, n

longer postsetat lobe; itis supported by one to several neuroaciculae and bears a fascicle of timbate, geniculate

setae and in some genera single large spines, with or without hood, or bidentate hooks. Dorsal and ventral I

cirri are absent. Most species of Drilonereis have minute parapodia, at least in the anterior part of their body.

Generally, the parapodia are about as large as the segmental length. Changes in proportion of the parapodia

constitute one of the few external characters useful for species discrimination. II

The pygidium bears two or four short anal cirri or two swollen pads; the anus is terminal. Colbath

(1989) considered the presence and number of anal cirri an important species specific character, although he

acknowledged that obtaining whole specimens is fairly difficult. I

i

Taxonoc History I i

The family Oenonidae was established by Kinberg (I 865) and was defined by ajaw apparatus with iI

long, slender maxillary carriers and mandibles that were shorter than the maxillae. The family included four

genera, two of which were subsequently synonymized with Oenone Kinberg. The oenonids were distinguished

from the closely related Laidea by the presence of foliaceous gills. Hartman (1944) noted that Kinberg's _l

family Laidea did not include the genus Arabella Grube, 1851, nor did any of Kinberg's other eunicemorph It

families. She therefore established the Arabellidae, consisting of Arabella and the genera assigned by Kinberg

to the Laidea. Because of the presence of superficially similar foliaceous notopodial lobes, the genus Oenone m

was added to the family Lysaretidae, also established by Kinberg (1865), but originally consisting only of the I

genus Lysarete. With the addition of Oenone, the Lysaretidae became less well-defined than intended by

Kinberg, who had positioned the two genera in two different major groups of genera due to the differences in •

the jaw apparatus. Colbath (I 986, 1989) proposed the restriction of the Lysaretidae to their original definition II

and recognition of the family Oenonidae to include Oenone and two other related genera. He based the

definition of those two families not only on morphological details of the jaw apparatuses, but also on the •

coinciding different chemical composition ofthejaw pieces. Lysaretids, like lumbrinerids, have jaws mineralized il

with calcite crystals, whereas the jaws of oenonids, like those of arabellids, consist only of scleroproteins and

lack crystals. Orensanz (1990), in his revision of the eunicemorph polychaete families, subsequently transferred 1

Lysarete to the Lumbrineridae because of similarities in the structure of the maxillary apparatus. He also |

merged the Arabellidae with the Oenonidae, the latter being the older name. The families Lysaretidae and

Arabellidae axe therefore no longer recognized. I

316 I
!

I The history of the oenonid genera is as confusing as that of the family, probably because oenonids are

never found inabundance. Most of the genera originally assigned tothe Oenonidae by Kinberg have subsequently

i been synonymized, others are indeterminable; within the free-living genera originally assigned to the Arabellidae, the weight of diagnostic characters is not equal. Fauchald (1970) suggested to either splitArabeIla into two

genera to separate species with a proximally dentate MI (Notopsilus) from those with proximally smooth MI

i (Arabella), or to combine Drilonereis and Notocirrus, which differ only by the dentate vs. falcate MI. The latter may be more practical because some species of Notocirrus have MI with falcate distal teeth. This

continuous range in shape of the MI was already noted by CrossIand (1924) who, for the first time, attempted

I to include individual variability of taxonomic characters such as setae and jaw pieces into the species descriptions, especially for lumbrinerids (then includingArabella and Drilonereis). According to Orensanz

(1990), there are five parasitic genera (Biborin Chamberlin, Drilognathus Day, Haematocleptes Wiren,

am Labrorostratus Saint-Joseph, and Oligognathus Spengel) and six mostly free-living genera ifNotocirrus and

l Drilonereis are kept separate (Oenone Savigny, HaUa Costa, Tainokia Knox and Green, Arabella Grube,

Drilonereis Clapar_de, and Notocirrus Schmarda). One additional parasitic genus, Pholadiphila, has since

been described by Dean (1992). !

I Distribution and Biological Notes

I Oenonids are widely distributed in all oceans from intertidal to abyssal depths. Free-living forms burrow in sand and mud, and are carnivorous or highly selective deposit feeders (Fauchald and Jumars,

1979). They do not build tubes, but appear to line their burrows with mucus. Parasitic species live in the

i body cavity of other polychaetes, such as eunicids, onuphids, spionids, cirratulids, and tricho-branchids, the echiuran Bonellia, (Pettibone, 1957) and pholadid bivalves (Dean, 1992). Not much is known about the

biology of these parasites; apparently not all of them are parasitic for their entire life, but leave their host's

body to become sexually mature as free-living forms. How they enter their host is not known; if they live

i singly in a host, they often reach a considerable size, occupying the entire body cavity of the host organism. w

I Other species are of small size and may occur in great numbers in a host (Pettibone, 1957). Parasites are not

included in the key below; Fauchald (1977) should be used to identify these forms to genus, and Pettibone

I (1957) is the most reliable reference for identification to but taxa have been added since that species, more

time (see Emerson, 1974 and Dean, 1992 for more detail). Uebelacker (1978) can be consulted for the

parasitic genus Labrorostratus.

!

i Key to the Oenonidae

1A. Parapodia with conspicuous, foliaceous notopodia; prostomium with three antennae along posterior

I lB. Notopodia margin ......................................................................................................................... absent or reduced to small papilla, sometimes with fascicle of delicate internal genus aciculae; Oenon_

prostomium without appendages ...................................................................................................... 2

I

2A. Setae including limbate capillaries and thick acicular spines, atleast in middle and posterior parapodia

.............. ................... ...... ................... .................. ................ ................. ............................................ 3

I 2B. All setae lirnbate, thick acicular spines absent ................................................................................. 8

i 2GenusnotrepresentedinthisArias

i 317
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3A. First maxillae falcate; small denticles if present restricted to basal part ..... genus Drilonereis ...... 4 M

3B. First maxillae dentate along entire cutting edge ..................................................... genus Notocirrus 2

I

4A. Acicular spines present from setiger 1; anterior parapodia minute, with short pre- and postsetal lobes;

postsetal lobes gradually turning digitiform in middle setigers, presetal lobes turning digitiform in I

posterior setigers and becoming subequal to postsetal lobes, parapodia thus distinctly bilabiate in •

posterior third of body; MI with 1-5 basal teeth, MII with 3-5 teeth, MIII and MIV with 1 tooth and

sometimes additional minute dentitions .................................................................. Drilonereis longa Im

4B. Acicular spines first present in setiger 10 or later; parapodia not distinctly bilabiate in posterior l

setigers 5

5A. Postsetal lobes twice as long as parapodial stem after setiger 70, cirriform, pointing dorsad; all setae I

with narrow, fmety serrated wings; MI with 6-7 basal teeth, MII 8+6, MIII 5-7, MIV 1, MV 1;

mandibles large, together forming a horseshoe shape .................................... Drilonereis spectabilis I

m

5B. Postsetal lobes papilliform or conical, as long as parapodial stem at most ..................................... 6

6A. Mandibles present; parapodia smaller inanterior setigers than in posterior, but proportions of parapodial I

lobes not changing; acicular spines first present in setiger 10 to 13; MI with 3-4 basal teeth (large

specimens up to 7), MII 6-9, MIII 1-5, MIV 1-2, MV 1 (occasionally absent); mandibles large ...... I

............................................................................................................................. Drilonereis falcata

6B. Mandibles absent ............................................................................................................................. 7

I

7A. Parapodia minute in anterior setigers, with papilliform postsetal lobes in posterior setigers; acicular

spines very large, projecting far beyond postsetal lobes; MI with 4-5 basal teeth ............................. I

|

......................................................................................................................... Drilonereis mexicana

7B. Parapodia conspicuous throughout, with digitiform postsetal lobes in posterior setigers; acicular

spines first present in setiger 25-30, projecting about as far as postsetal lobes; MI basally smooth. •

................................................................................................................................ Drilonereis nuda

Ii

8A. Postsetal lobes in posterior setigers digitiform, very long and pointing dorsad; upper supra-acicular I

and subacicular setae smooth, lower supra-acicular setae with serrated wings .................................

...................................................................................................................... Arabella semimaculata •

8B. Postsetal lobes prominent, but not digitiform; surficial structures of setae different ....................... 9 |

9A. All setae with serrated wings; MI with unidentate tips on both sides, MI/I and MIV with about 4-5 1

conical teeth, alternating large and small ................................................................ Arabella iricolor

9B. Some setae with smooth wings; left MI with bidentate tip ............................................................. 10 I

i

il

10A. Subacicular setae smooth in middle and posterior setigers, supraacicular setae serrated; MIII and

MIV with about 4-5 conical teeth, alternating large and small ....................... Arabella protomutans 1

I

10B. All setae smooth in anterior setigers, serrated in middle and posterior setigers; MIII and MIV with

about 10 needlelike equal teeth, giving comblike shape to maxillae .................... Arabella pectinata !

318 I
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I Description of Species

I

The Oenonldae are a relatively small family with about 80 species belonging to 12 genera. Only three

of these genera, Arabella, Drilonereis, and Notocirms consist of ten or more species, while most of the

I parasi the onlytic oenonids genera arefound represen belong ted byto Drilonereis only one spe, ciincluding es. In thone e Santa species Marinewa Bastoinscand ience, SantabutBarbara it is likelyChann thatel,at

least some of the species of Arabella described below occur in this area, especially in shallower depths. The

I following species are treated:

i Ara Ar bella (Arabella) iricolor (Montagu, 1804)

abelIa (ArabeUa) pectinata Fauchald, 1970

Arabella (Arabella ) protomutans Orensanz, 1990

i ADrilonereisfalcata rabella (Arabella) Moore, semimaculata (Moore, 1911) 1911

Drilonereis longa Webster, 1879

i Drilonereis mexicana Fauchald, 1970 Drilonereis nuda Moore, 1909

Drilonereis spectabilis Hilbig, new species

I Genus Arabella Grube, 1850

I Type Species. Nereis iricolor Montagu, 1804

Diagnosis. Prostomium spatulate to conical, without antennae, usually with four eyes along posterior

I margin; no ventral palps. Peristomium with two rings, separate from one another dorsally and ventrally, anterior one forming a simple lower lip. Mandibles simple, large plates; maxillae usually asymmetrical, MI

falcate, with or without basal dentition, or dentate along entire cutting edge; MII, MIII, and MIV multidentate,

I MV usually unidentate. Parapodia subbiramous with papilliform notopodial rudiment; postsetal lobes longer than presetal lobes. Setae all simple, limbate, with smooth or serrated wings; ventralmost seta modified in

some species to blunt-tipped spine with terminal winglike hood; thick acicular spines absent.

I Remarks. The genusArabella is somewhat ill-defined within the family Oenonidae because characters that are considered diagnostic for other genera, such as the presence or absence of dentitions on the first

maxillae, are variable inArabella. Additionally, the presence of modified setae in some species is unique for

i the genus Arabella; all other genera have very uniform setae. It therefore seems useful to keep some of the

genera previously synonymized withArabella as subgenera, as has been done by Orensanz (1976, 1990) and

Perkins (1979), even though a cladistic analysis of the oenonids may prove these subgenera, and likely the

I genusArabella, invalid (Fauchald, personal communication). The subgenera are defined as follows:Arabella (Arabella) has falcate MI and unmodified setae; A. (Cenothrix) has falcate MI and modified setae, and A.

(NotopsiIus) has dentate MI and unmodified setae.

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Arabella (Arabella) iricolor (Montagu, 1804) I

n

Figure 12.1

Nereis iricolor Montagu, 1804:82. I

Lumbriconereis opalina Verrill and Smith, 1874:26,38, etc., pt. 13, figs. 69-70.

Arabella opaIina: Webster, 1879:242; 1884:321; 1886:145.--VerriI1, 1881:291,301, etc., pl. 4, fig. 4.-- m

Webster and Benedict, 1884:721. II

u

Arabella iricolor: Mclntosh, 1910:395, pl. 54, fig. 4, pl. 62, figs. 8-8c, pl. 74, figs. 5-5c, pl. 83, figs. 2-2a.--

Hartman, 1944:173 (inpart); 1968:789. Pettibone, 1963:269-271, figs. 71a-e.--Fauchald, 1970:125-

128, pl. 20, figs. a-d.----Orensanz, 1974:384-386, pl. 1.--Blake, 1975:205, fig. 194.--Uebelacker, l

1984:42-5, figs. 42-1, 42-2a-f.

Material Examined. California: Tomales Bay, det. Olga Hartman (2); Dillon Beach, intertidal, coll. I

K. Woodwick, 19 June, 1962 (4). II

Description. Length to 600 ram, width to 5 ram, segments to 500. Body cylindrical, wiry, of equal I

width except for short prepygidial region. Color in alcohol pale to reddish brown with iridescent cuticle, head 1

usually the least pigmented.

Prostomium bluntly conical, longer than wide when fully extended, with four subequal eyes in straight I

line along posterior margin, sometimes obscured by peristomium when prostomium retracted (Fig. 12.1A). II

Proboscis with large, oval to triangular mandibles (Fig. 12.1B) and five pairs of maxillae; all jaw pieces

black, very hard, heavily sclerotized. Paired maxillary carriers very long and slender, free from one another; m

unpaired piece about two-thirds as long, elongate-oval. First and second maxillae asymmetrical; right MI 1

smallest, with 5 basal teeth, left MI with 10 teeth, both sides strongly falcate; MII curved over distal ends of

MI; right MII largest, with 10-16 teeth, left MII with 6-9 teeth. MIIJ through MV symmetrical; MIII and •

MIV large, rounded, homy plates with very hard cutting edge bearing 4-7 teeth, large and small ones alternating; |

MV single, long, sharply pointed tooth arising from very small base (Fig. 12.1B). Peristomium with two

rings, the anterior one forming lower lip, both equal to following setigers. II

Parapodia similar throughout, with short presetal and conical to digitiform postsetal lobes, the latter I

longest in posterior setigers (Fig. 12.1D, E). Setae all of one type, geniculate, limbate, with finely to coarsely

serrated wings (Fig. 12.1F, G); aciculae in fascicles of about 3 per parapodium, dark amber, with long m

filiform tips breaking through integument. |

Pygidium with 2 to 4 short anal cirri.

Remarks. Arabella iricoIor is one of the most familiar and widely reported species of this genus, but n

1

may have been confused with other species such as A. pectinata and A. protomutans that have an identical U

habitus, but differ in details of the setal surface structures and dentition of the maxillae (see below). To

positively identify A. iricolor, at least the setae should be examined on a slide, preferably those from a 1

posterior parapodium. The first and second maxillae are reported to be asymmetrical by most authors, II

although there appears to be some variability in this character; for example, Mclntosh (1910) described the

maxillae as being symmetrical. 1

II

Type Locality. Devonshire, England.

Habitat. A. iricolor burrows deeply into mud, sand and gravel; itis also found under rocks, in oyster II

and mussel beds, among Zostera boldfasts, among bryozoans and other colonial animals. II

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320 I
I Figure 12.1. Arabella (A.) iricolor: A, anterior end, dorsal view (from Blake, 1975); B, maxillae (after Orensanz,

1974); C, mandibles (after Pettibone, 1963); D-E, anterior and posterior parapodium, anterior view

I (after Fauchald, 1970); F, smooth limbate seta (after Pettibone, 1963); G, serrated limbate seta.

1 I

I Distribution. Cosmopolitan in and subtropical France; Mediterranean

temperate waters; England,

Sea; Massachusetts to Florida, Gulf of Mexico, Colombia and Venezuela; Vancouver Island to California,

Mexico, Argentina; Japan, China, Persian Gulf, Red Sea, Indian Ocean; Straits of Magellan; West and South

I Africa; intertidal to 90 m.

I

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Arabella (Arabella) pectinata Fauehald, 1970

Figure 12.2 I

Arabella pectinata Fauchald, 1970:130-132, pl. 22, figs. a-f. I

U

Material Examined. California: Tomales Bay (1).

Description. Length at least 182 mm, width to 3.5 mm, segments at least 355. Body cylindrical,

relatively flexible, increasing in width through pharyngeal region, rest of body of equal width (Fig. 12.2A, B). I

Color in alcohol copper to purplish brown, highly iridescent.

Prostomium bluntly conical, about as wide as long, less pigmented than rest of body. Peristomium i

with two equal rings, slightly wider than prostomial base (Fig. 12.2A). Proboscis with mandibles and well- m

developed maxillae. Mandibles paddle-shaped, fused anteriorly, diverging posteriorly (Fig. 12.2C); maxillae

with slender paired carders, free through most of length, fused anteriorly above insertion of slender unpaired

piece. First and second maxillae asymmetrical; both MI of subequal size, falcate, with 7-11 basal teeth, left I

side with bifid tip; MII curving over distal ends of MI; left MII smallest, with 8-12 teeth, right MII with 14-

20 teeth; MIU through MV symmetrical; MIII and MIV large, rounded, horny plates with heavily sclerotized

cutting edge beating 6-11 needlelike teeth, all of equal size, cutting edges looking like combs; MV single long,

acutely pointed tooth arising from small base (Fig. 12.2D).

Parapodia similar throughout, with short, rounded presetal lobes and digitiform postsetal lobes about i

twice as long as presetal ones, thickest in anterior setigers; notopodial rudiments present (Fig. 12.2E, F).

Setae of two kinds; anterior parapodia with smooth to very finely serrated, nearly straight limbate setae (Fig.

1

2.2G), middle and posterior parapodia with very finely serrated setae in uppermost and lowermost position

and strongly geniculate, coarsely serrated setae in middle of fascicle (Fig. 12.2H-J).

u

Remarks. ArabeUa pectinata has most likely been confused with A. iricolor, but differs markedly II

from the latter in the following characters: the setae are all serrated in A. iricolor, but smooth at least in I

uppermost and lowermost position in each fascicle inA. pectinata; the third and fourth maxillae have about 5

conical teeth of alternating size inA. iricolor, but about 10 needle-like teeth of equal size inA. pectinata. The m

fine structures of the serrated setae that Fauchald (1970) thought to be unique forA. pectinata are common to |

allArabella setae with coarse serrations; the posterior connection of the maxillary carriers were not observed

in the examined specimen, but a U-shaped transparent ligament, rather than a rounded plate, was seen that is mt

most likely the same structure. This record is the f'n-stone for California.

Type Locality and Type Specimens. Mexico, E1 Descanso: holotype and 3 paratypes (LACMAHF).

Distribution. Central California to Mexico (Baja California), intertidal to shallow subtidal depths. M

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I Figure 12.2. ArabeUa (Arabella) pectinata: A, anterior end, dorsal view; B, middle setigers, dorsal view; C,

mandibles; D, maxillae; E-F, anterior and posterior parapodium, anterior view; G, finely serrated

seta, anterior parapodium; H-J, upper, middle and lower seta from posterior parapodium. 03, E, F

I after Fauehald, t970).

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Arabella (Arabella) protomutans Orensanz, 1990

Figure 12.3 I

Arabella protomutans Orensanz, 1990:102-104, pl. 28.

Arabella novecrinita var. asymmetrica Crossland, 1924:80-82, figs.103-104 (in part).--Not Arabella II

(Cenothrix ) asymmetrica Orensanz, 1974.

Material Examined. California: Tomales Bay, (1), Tomales Point (1); Avila, intertidal, coll. K. 1

Woodwick, 3 March, 1963 (1); Dillon Beach, intertidal, coll. K. Woodwick, 19 June, 1962 (2). B

Description. Length to 350 mm, width to 2 mm excluding parapodia, segments to 600. Body m

cylindrical, wiry, with relatively long segments and small but conspicuous parapodia. Color in alcohol pale to 1

pinkish brown, most segments of Californian specimens with broad band of reddish pigment, head least

==

pigmented. I' Prostomium conical, as long as wide, with four eyes in a row along posterior margin (Fig. 12.3A). '

Proboscis with mandibles and five pairs of maxillae. Mandibles large, paddle-shaped, fused along distal haft

(Fig. 12.3C). Maxillary carders with long, slender paired pieces free from one another and ventral unpaired •

piece of similar shape and almost as long. First and second maxillae asymmetrical; left MI with bifid tip, I

slightly larger than right MI, both with about 8 basal teeth; left MII much smaller than right MII, with 6-7

unequal teeth; right MII with about 14 unequal teeth. MIH, MIV, and MV symmetrical; MHI and MIV large II

plates with very hard cutting edges each bearing 4 to 5 unequal teeth, large and small ones alternating; MV I

single, sharply pointed tooth arising from very small base (Fig. 12.3B).

Parapodia similar throughout, with papilliform notopodial rudiments, low presetal lobes, and conical, •

elongated postsetal lobes (Fig. 12.3D, E); setae geniculate, with f'mely to coarsely serrated wings (Fig. 12.3F); II

subacicular fascicles of posterior parapodia with smooth setae (Fig. 12.3G).

Pygidium with two lateral pads, no anal cirri, anus slitlike.

Remarks. Arabellaprotomutans is closely related and intermediate to bothA, iricolor andA. pectinata

(see above), with MI resembling that ofA. pectinata and the irregular dentition of MIH and MIV being similar

to A. iricolar. It is most easily recognized by the smooth subacicular setae in posterior parapodia; if only I

anterior fragments are available, the pharynx should be dissected to examine the first and third maxillae. This

record is the first from outside the Magellanic biogeographic province. The Californian specimens are

considerably larger than the type material. I

U

Type Locality and Type Specimens. Southwestern Chile, Eltanin Sta. 958, 92-101 m: holotype

(USNM 98099) and 1 paratype (USNM 98100). m

Habitat. This species was found in sandy sediments. I

Distribution. Strait of Magellan; central California; intertidal to 118 m.

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324 I
I

I

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I

I

I

F_ 12_3. Arabella (A.) protomutans: A, anterior end, dorsal view; B, maxillae; (2, mandibles; D-E, middle

and posterior parapodium, anterior view; F, serrated supra-acicular setae, posterior parapodium; G,

I ventraimost setae, posterior parapodium. (all after Orensanz, 1990 except for left seta in F and right seta in G).

I

I
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l

Arabella (Arabella) semimaculata (Moore, 1911) •

Figure 12.4

Aracoda semimaculata Moore, 1911:295-297, pl. 20, figs. 143-149. II

ArabeUa semimaculata: Hartman, 1944:173; I968:793.--Fauchald, 1970:132-133, pl. 20, figs. e-g.--Blake,

1975:205, fig. 197. I

Material Examined. California: Dillon Beach, intertidal, coll. K. Woodwick, 19 June, 1962 (1). II

Description. Length to 165 mm, width to 2.8 mm, segments to 278. Body slender, cylindrical, B

appearing widened in middle and posterior setigers because of very prominent parapodia (Fig. 12.4A). Color

in alcohol pale greyish, some specimens with three dorsal series of bluish dusky spots and deeply pigmented II

peristomium. m

Prostomium as long as wide or slightly longer, conical, slightly depressed dorsoventralty, with four

eyes, may be obscured in very large specimens. Jaw apparatus with large, anteriorly flared mandibles, fused •

subdistally, divergent proximally and distally. Maxillary carriers with three subequal, long, slender pieces. l

First and second maxillae asymmetrical; left MI somewhat larger than right one, with 8 to 11 basal teeth, right

MI with about 7 to 11 smaller teeth. Left MII much smaller than right one, with 6 to 9 teeth; right MII with •

about 13 to 17 teeth. MIII, MIV, and MV symmetrical; Mill and MIV rounded plates with cutting edges I

bearing 4 to 7 unequal teeth; MV with single tooth (Fig. 12.4B, C). Accessory elongate, soft plates present

alongside MII.

Parapodia small and inconspicuous anteriorly, with papilliform notopodial rudiments, rounded presetal U

lobes, and conical postsetal lobes pointing upward and outward (Fig. 12.4D). Parapodia increasing in size

after first few setigers; postsetal lobes increasing in length, turning digitiform, pointing dorsad (Fig. 12.4E). II

Setae few, geniculate, smooth to finely serrated in superior and inferior position of fascicle, distinctly serrated l

in median position of fascicle (Fig. 12.4 F-H). Notoaciculae delicate, clear, in fascicles of 2 or 3; neuroaciculae

stouter, pale, also in fascicles of 2 or 3, with filiform tips breaking through integument, i

Pygidium with 2 or 4 very small anal cirri.

Remarks. This species was reported to be the most common of the genus in Mexico by Fauchald

(I970). i

Type Locality and Type Specimens. California, Monterey Bay, off Santa Cruz Lighthouse: holotype

(USNM 16918), 3 paratypes (USNM 17272), 3 paratypes (ANSP 2985).

Habitat. Sand and rocks, mixed sediments, i

Distribution. Mexico to central California, intertidal to 80 m.

Genus Drilonereis Clapar_de, 1870 U

Type Species. Lumbriconereisfilum Ciapar_de, 1868 I

Diagnosis. Prostomium spatulate, without eyes or antennae. Peristomium with two rings, separate I

from one another, the anterior one forming simple lower lip. Setae including limbate capillaries and thick, n

acicular spines in most setigers; maxillae with falcate, basally smooth or dentate MI. U

Remarks. Some species of Drilonereis are parasitic; inthese forms,jaws, parapodia, and setae may

be reduced, i

I

326 i
I

I

I _ ::

I I__

I __

I oooo"', , /

I • \ ,.,: 'i_

I

I ..::._ .... .._

:!""""":"_:?::; ::?i" :"!_!.. ,_!!_!!:!':° /

ii E..:..::..: .."_ ..i.:_i:: -:....::::.,

I Figure 12.4. Arabella (A.) semimaculata: A, anterior end, dorsal view; B, maxillae; C, mandibles (after Moore,

1911); D-E, anterior and posterior parapodium, anterior view (after Fauchald, 1970); F, finely

I serrated seta; G-H, coarsely serrated seta, lateral and frontal view (detail).

I

i 327
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DriIonereisfalcata Moore, 1911 IJ

Figure 12.5

m

DrilonereisfalcataMoore, 1911: 289-299, pl. 20, figs. 150-154.--Hartman, 1944:179; 1968:797.--Fauchald, i

1970:135-136, pI. 21, fig. g.--Orensanz, 1974:397-399, pl. VI/.--Lissner et al., 1986:D-12.--

Steinhauer and Imamura, 1990:F- 1. 1

tim

Material Examined, California: Santa Mafia Basin, off Point Buchon, Sta. 12 (1); off Point San

Luis, Sta. 21 (1); off Point Sal, Sta. R-8 (1). i

Description. Length atleast 120 mm, width to 1.2mm, segments to 230. Body thin, wiry, cylindrical,

with anteriorly small but conspicuous parapodia throughout; anterior segments about twice as long as parapodial

width, middle and posterior segments much more crowded, only as long asparapodial width (Fig. 12.5A, B). i

Color in alcohol light tan to purplish brown.

Prostomium spatulate, about as long as wide, with median dorsal furrow; nuchal organs inconspicuous I

(Fig. 12.5A, C). Mandibles well sclerotized, roughly triangular, about half as long as MI, fused along short i

distal area (Fig. 12.5D); maxillae with long, slender carriers and lanceolate unpaired ventral piece; MI large,

strongly falcate, with 3 or 4 large basal teeth; MII with 6 to 9 teeth, subdistal one smallest; MI]/with 3 teeth,

1 large and 2 small; MIV andMV each with one pointed tooth, MIV occasionally with additional serrafions i

along cutting edge; MV may be absent (Fig. 12.5 E). Peristomium consisting of two rings, each as long as

following setigers and sligthly wider than prostomial base. All setigers uniformexcept forgradually decreasing

length within anterior part of body; parapodia with minute notopodial papilla supported by bundle of slender 1

notoaeiculae, neuropodia with short presetal and longer, conical postsetal lobes throughout; length of parapodial

stems doubling or tripling between anteriormost and posterior setigers, proportions of pre- and postsetal lobes

practically unchanged (Fig. 12.5F). Limbate setae with narrow wings, covered with minute surficial spines, i

finely serrated in side view (Fig. 12.5 G, H). Single thick, acicular spines present from about setiger 10,

protruding furthest in posterior setigers, almost reaching length of postsetal lobes (Fig. 12.5 I).

Pygidium with 4 anal cirri, i

UB

Remarks. The unchanging proportions of the parapodial lobes along the body and the basal teeth on

MI are diagnostic for this species. on

Type Locality and Type Specimens. California: Monterey Bay, off Point Pifios lighthouse: holotype I

(USNM 17283), 1 paratype (USNM 17282), 2 paratypes (ANSP 2984).

Habitat. DrilonereisfaIcata is free-living and occurs in mud, sand, and gravel and in mixed sediments, i

Distribution. Central California to Argentina; South Africa; intertidal to 350 m, possibly cosmopolitan.

I

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wob

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I /;

I r!'

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I

I Figure 12.5. Dlatrileronalereis viewfal; caD,tam: andA,iblesant; erE, iormax endil, laedor; sF, al midd view;leB,paramidpoddleium, setigers, anteriordorsviewal v;iew; G-H,C,seta anteirniorlateral end,

and frontal view; I, spine. (D-F after Orensanz, 1974).

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i 329
I

Drilonereis longa Webster, 1879 I

Figure 12.6

Drilonereis longa Webster, 1879:240-241, pl. 7, figs. 84-88.--Pettibone, 1963:272-273, fig. 72.--Hartman, II

1968:801.--Uebelacker, 1984:42-12 -42-15, figs. 42-9, 10a-d.

Drilonereis nr. longa: Lissner et at., 1986:D-12.--Steinhauer & Imamura, 1990:F-1. 1

Drilonereis sp. B Lissner et al., 1986:D-12.

Material Examined. California: Santa Maria Basin, off Purisima Point, Sta. R-5 (1); Western Santa i

Barbara Channel, Sta. 79 (2).

Description. Length to 710 mm, width to 1.5 ram, segments several hundreds. Body very thin,

flexible, cylindrical, with tiny parapodia in anterior setigers (Fig. 12.6A). Color pale, iridescent, maxillae jet 1

black and usually visible through body wall. II

Prostomium spatulate, somewhat longer than wide, with median furrow dorsally. Pharynx armed am

with mandibles and 4 pairs of maxillae. Mandibles small, triangular, free from one another, anterior to i

m

maxillae; maxillae with long carriers, atleast twice as long as MI, and oval, elongate unpaired piece about as

long as MI. First maxillae strongly falcate, heavily sclerotized, with 2 to 3 large basal teeth; MII about half 1

as long, with 3 or 4 teeth; MIII and MIV each with 1very sharply pointed tooth, MIB occasionally with few 1

additional minute denticles; MV absent (Fig. 12.6B).

u

Setigers shortest in anterior part of body, equal to peristomial rings, bearing inconspicuous parapodia •

with few limbate capillaries and single acicular spine (Fig. 12.6C); middle and posterior setigers becoming l

longer than wide, parapodia gradually increasing in length; postsetal lobes turning from short papilla into

conical lobes after setiger 30 to 40; presetal lobes gradually becoming conical after about 50 more setigers •

(Fig. 12.6D); in posterior region, both lobes conspicuous and divergent. Setae including limbate capillaries 1

with very narrow, smooth wings anteriorly and somewhat wider, smooth to very finely serrated wings in

middle and posterior parapodia (Fig. 12.6E); and single, thick acicular spines about asthick as postsetal lobes 1

and longer than the latter in middle setigers, about half as wide in posterior setigers and about as long as i

postsetal lobes.

Ill

Pygidium with 4 short anal cirri. I

Remarks. Most of the specimens from the Santa Maria Basin and Santa Barbara Channel were 1

originally identified asD. nr. longa, but no differences could be found between those specimens and any of the

available descriptions, and they are here considered asD. longa. One specimen, identified asDrilonereis sp. •

B, had been dissected for examination of the jaws. Mandibles are absent, but all other characters match the

description ofD. longa so closely that it was concluded by the author that the small mandibles may have been

overlooked during the dissection of the pharyngeal bulb. Itis also possible thatDrilonereis sheds itsjaws in •

the same manner as is known for lumbrinerids (Colbath, 1987) with the mandibles not yet regenerated in the I

present specimen. Establishment of a new species seems unjustified; the specimen was therefore assigned to

D. longa.

Type Locality. V'trginia. •

Habitat. The species appears to be most common in silt and clay, but has been reported from sand, 1

fine gravel, and shell rubble as well. The specimens from the Santa Barbara Channel were found insediments 1

with more than 70% sand. II

Distribution. Massachusetts to Georgia; Gulf of Mexico, West Indies; Washington to southern

California, intertidal to 2450 m. •

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I Figure 12.6. Drifonereis longa: A, anterior end, dorsal view; B, maxillae and mandibles; C-D, anterior and

posterior parapodium, anterior view; E, limbate seta. (A-D after Uebelacker, 1984; E after Pettibone,

i 1963).

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Drilonereis mexicana Fauchald, 1970 1

Figure 12.7

DriIonereis mexicana Fauchald, 1970:138-140, pl. 23, figs.a-c. I

Description. Length to 240 mm, width to 1.5 mm including setae, segments to 800. Color evenly •

reddish brown. II

Prostomium short, conical, as long as both peristomial rings together. Jaw apparatus consisting of 4

pairs of maxillae; mandibles absent. Paired maxillary carriers very long and slender, almost three times as •

long as first maxillae, fused anteriorly; unpaired carrier roughly rectangular, about as long as first maxillae.

MI falcate, with 4 or 5 basal teeth; MII with 4 teeth, MIII with 2 teeth, and MIV with 1 pointed tooth (Fig.

12.7B). Anterior I

setigers about two to three times as wide as long, gradually lengthening, twice as long as

wide in far posterior setigers. Parapodia minute and inconspicuous in anterior setigers, slightly increasing

after about 30 setigers, with short, papilliform postsetaI lobes throughout (Fig. 12.7A). Setae geniculate, II

f'mely serrated, with moderately wide wings and long, slender tips (Fig. 12.7C). Acicular spines very large, It

projecting much further than postsetal lobes. Aciculae with long filamentous tips.

Remarks. Although no specimens of D. mexicana were reported during the Phase I and Phase II 1

programs, this species may co-occur with D. nuda and may have been misidentified as the latter because of II

the easily observed absence of mandibles shared by both species. Hartman (1944) described a mixture of

these two species as D. nuda, and she used the same erroneous species concept in the Atlas (1968). The 1

differences in the proportions of the parapodia--minute anteriorly and increasing toward the middle of the II

body, postsetal lobes papilliform forD. mexicana, of almost equal size throughout and with digitiform postsetal

lobes for D. nuda--and the presence or absence of serrations on the setae may be sufficient characters to 1

discriminate between those two species without having to dissect the maxillae.

Type Locality and Type Specimens. Mexico, San Cristobal Bay (LACM-AHF). i

Habitat. D. mexicana was found in medium to coarse sand, mixed with mud or shells. 1

Distribution. Southern California to Baja California, 30 - 130 m.

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| A B C

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Figure 12.7. Drilonereis mexicana: A, parapodium, anterior view; B, maxillae; C, seta (all from Fauchald,

I 1970).

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Drilonereis nuda Moore, 1909 I1

Figure 12.8 I

Drilonereis nuda Moore, 1909:254-256, pl. 8, figs. 21-23.--Hartman, 1944:I78-179, pl. 13, figs. 297-302

(in part); 1968:803 (in part).--Fauchald, 1970:140-141, pl. 22, fig. g.--Lissner et al., 1986:D-12. I

Material Examined. California: Santa Maria Basin, off Point Estero, Sta. 6 (rock) (1).

Description. Length to 630 ram, width to 1.3 ram, segments to 935. Body long, cylindrical, of nearly •

uniform diameter, with conspicuous parapodia throughout. Color in alcohol brown to light golden, with dark 1

spots at parapodial bases, sometimes with 2 dark rings across each setiger, with iridescent cuticle.

Prostomium about as long as wide, spatulate, rounded anteriorly, with median dorsal furrow. Nuchal •

organs not discernible; examined specimen with 4 pigment spots on brain, possibly subdermal eyes (Fig. 1

12.8A). Peristomium with 2 rings, the anterior one indistinctly separated from prostomium, about as wide as

prostomial base and of equal length to second ring and subsequent setigers. Pharynx armed with 4 to 5 pairs •

of maxillae, mandibles absent. Maxillae with slender carriers, about twice as long as MI, anteriorly fused I

over short distance; ventral unpaired piece short, oval. MI strongly falcate, very hard, basally smooth; MII

more delicate, but cutting edge with 5 to 7 massive teeth. Mill and MIV each with 1 sharply pointed tooth, •

MH/sometimes with additional tooth or boss at base of main tooth. MV usually absent, when present much 1

smaller than MHI and MIV, with 2 minute teeth (Fig. 12.8B).

Parapodia conspicuous and of equal shape throughout length of body, smallestin few anterior setigers; I

presetal lobes truncate, postsetaI lobes conical, pointing dorsad; notopodial papilla minute (Fig. 12.8C).

Setae narrowly limbate with smooth wings, strongly geniculate in anterior and middle setigers, more straight

and slender in far posterior setigers. Acicular spine first present around setiger 25 to 30, golden, projecting •

about as far as postsetal lobes, usually single, sometimes two per parapodium (Fig. 12.8D-F). Neuroaciculae

fine, with very long filiform tips breaking through integument.

Pygidium without anal cirri, with 2 padlike lobes surrounding slitlike anus. 1

I1

Remarks. The species is defined here as originally described by Moore (1909), with basally smooth

first maxillae; Fauchald's (1970) notion that Hartman (1944) included two species in her definition, one with I

smooth MI and the other with dentate MI, is accepted. The latter was referred to Drilonereis mexicana by 1

Fauchald (1970). The two species may co-occur, but can be distinguished from one another without dissection II

of the jaws by the following differences: the setae are smooth in D. nuda, but finely serrated in D. mexicana;

and the acicular spines project about as far as the postsetal lobes in D. nuda, but far beyond in D. mexicana. I

m

Type Locality and Type Specimens. California, Monterey Bay: holotype (CASIZ 56, ANSP slide

2.24), 2 paratypes (CASIZ 77), 2 paratypes (ANSP 2578). 1

Habitat. Drilonereis nuda typically occurs on rocky bottom; the type locality was a large tide pool. 1

Distribution. Central California to Mexico (Baja California), intertidal to about 100 m.

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i Figure 12,8. Drilonereis nuda: A, anterior end, dorsal view; B, maxillae; C, posterior parapodium, anterior

view; D, supraacicular seta; E, spine; F, subacicular seta.

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Drilonereis spectabilis Hilbig, new species

Figure 12.9 I

Drilonereis sp. A Lissner et al., 1986:D-12. I

Material Examined. California: Santa Barbara Channel, offPoint Conception, Sta. 74, 34°26.84'N, 1

120°38.61 'W, 201 m: holotype (USNM 170424). IID

Description. Length of anterior fragment 100 mm, width 5 mm excluding parapodia; length of

posterior fragment 40 mm, width 3 mm. Body very robust, large for the genus, tapering slightly toward I1

prostomium and more distinctly toward pygidium, almost cylindrical, with several hundreds of setigers. Anterior 1

segments crowded, 8-10 times as wide as long, only as long as parapodial bases; posterior segments only 3-

1

4 times as wide as long, longer than parapodial bases (Fig. I2.9A, B). Color in alcohol uniformly tan. 1

Prostomium spatulate, as long as wide, distinctly narrower than peristomium. Pharynx with well- 1

developed maxillary plates and relatively large, rodlike, slightly curved mandibles resembling a black horseshoe

(Fig. 12.9C). Both mandibles and maxillary carriers broken during previous dissection performed from the •

ventral side, details not readily observed. Maxillary carriers with slender paired pieces, about twice as long 1

as MI, and slender, conical unpaired piece about as long as MI. MI very strongly f',dcate, with 4 large and

about 2-3 small basal teeth; MII stigthly asymmetrical, left one with 8 large and 3 small basal teeth, right one •

with 6 large and about 3 small teeth; M]II with main fang and group of 4 small teeth; MIV and MV with single 1

tooth, the latter with very small base (Fig. 12.9D). Peristomium with 2 rings resembling subsequent setigers.

Parapodia small in first few setigers, rapidly increasing in size after 10 to 15 setigers; presetal lobes •

in anteriormost parapodia obliquely truncate, in middle and posterior parapodia developed into short conical 1

lobes; postsetal lobes conical, very conspicuous throughout, about as long as parapodial stem to about setiger

70; after setiger 70, postsetal lobes turning digitiform to almost foliaceous, at least twice as long as parapodial •

stem, inserting ventrally and pointing dorsad (Fig. 12.9E). Limbate setae weakly geniculate, with f'mely 1

serrated wings; acicular spines present on all setigers, longest in posterior fragment; all setae amber colored

(Fig. 12.9F-H). Notoaciculae delicate, clear, in fascicles of about 3; neuroaciculae amber, about half as thick •

as acicular £pines, in fascicles of about 5 (Fig. 12.9E). 1

Remarks. The presence of long, digitiform to foliaceous postsetal lobes is unique in the genus, so

that the establishment of a new species based on only a single incomplete specimen isjustifiable. The habitus •

of D. spectabilis is similar to Arabella (A.)semimaculata.

Etymology. The species name was chosen to indicate the very unusual outer appearance of this

species. I

Habitat. The type specimen was found in silt mixed with sand (27%) andclay (16%).

Distribution. Known only from type locality, central California, 220 m. ==

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336 I
i Figure 12.9. Drilonereis spectabUis Hilbig, new species: A, anterior end, dorsal view; B, posterior segments, dorsal view; C, mandibles; D, maxillae; E, posterior parapodium, anterior view; F-G, seta, lateral

and frontal view; H, spine. (all from holotype).

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Literature Cited

!

Blake, J.A. 1975. Phylum Anne[ida: Class Polychaeta. Pp. 151-243 In: R.I. Smith and T.J. Carlton (eds.),

Light's Manual: Intertidal Invertebrates of the Central California Coast. University of California •

Press, Berkeley.

Colbath, G.K. 1986. Jaw mineralogy ineunicean polychaetes (Anne[ida). Micropaleontology 32:186-189.

I

Colbath, G.K. 1987. Evidence for shedding of maxillary jaws in eunicoid polychaetes. Journal of Natural II

History 21: 443-447.

Colbath, G.K. 1989. Revision of the family Lysaretidae, and recognition of the family Oenonidae Kinberg, 1

1865 (Eunicida: Polychaeta). Proceedings of the Biological Society of Washington 102 (1): 116-123.

Crossland, C. 1924. Polychaeta of tropical East Africa, the Red Sea, and Cape Verde Islands collected by I

Cyril Crossland, and of the Maldive Archipelago collected by Professor Stanley Gardiner, M.A., I

F.R.S. Proceedings of the Zoological Society of London 1:1-106.

Dean, H. 1992. A new arabellid polychaete living in the mantle cavity of deep-sea wood boring bivalves I

(family Pholadidae). Proceedings of the Biological Society of Washington 105(2):224-232.

m

Emerson, R.R. 1974. A new species of p[olychaetous annelid (Arabellidae) parasitic in Diopatra ornata 1

(Onuphidae) from Southern California. Bulletin of the Southern California Academy of Sciences

73:1-5.

Fauchald, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae, 1

Lysaretidae and Dorvilleidae from western Mexico. Allan Hancock Monographs in Marine Biology

5:1-335. l

Fauchald, K. 1977. The Polychaete Worms. Def'mitions and Keys to the Orders, Families and Genera. 1

Natural History Museum of Los Angeles County, Science Series 28:1-190.

Fauchald, K. and P.A. Jumars. 1979. The diet of worms: a study ofpolychaete feeding guilds. Oceanography 1

and Marine Biology Annual Review 17:193-284.

m

Hartman, O. 1944. Polychaetous Annelids Part V.Eunicea. Allan Hancock Pacific Expeditions 10:1-237. 1

Hartman, O. 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foundation,

University of Southern California, Los Angeles. 828 pp. 1

Kinberg, J.G.H. 1865. Annulata nova. t)fversigt af Kongliga Vetenskaps-Akademiens Frrhandlingar !

Stockholm 21:559-574.

Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bernstein, R. Cimberg, T. Kauwling, and W. Anikouchine. 1

1986. Assessment of long-term changes in biological communities of the Santa Maria Basin and m

Western Santa Barbara Channel - Phase I. Report prepared for the U.S. Department of the Interior, I

Minerals Management Service, Pacific OCS Region, under contract No. 14-12-0001-30032. I

1

Mclntosh, W.C. 1910. A monograph of the British annelids. Vol. 2, pt. 2. Polychaeta. Syllidae to Ariciidae.

London, Ray Society, pp. 233-524, pls. 51-61, 71-87. I_

Montagu, G. 1804. Descriptions of several marine animals found on the south coast of Devonshire.

Transactions of the Linnaean Society of London 7:80-84, pl. 7.

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i

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Moore, J.E 1909. Polychaetous annelids from Monterey Bay and San Diego, California. Proceedings of the Philadelphia Academy of Natural Sciences 61:235-295, pls. 7-9.

i Moore, J.California P. 1911. inThe1904. polyEuphrosynidae chaetous annelids to Goniadid dredgedab

e.y theProceedings U.S.S. Albaoftrthe oss Philadelphia off the coastAcademy of southemof

Natural Sciences 63:234-318, pls. 15-21.

I Orensanz, J.M. 1974. Los anelidos poliquetos de la provincia biogeografica Argentina. VI. Arabellidae. Physis Seccion A 33(87):381-408.

i Orensanz, J.M. 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas. Biology of the Antarctic Seas XXI. Antarctic Research Series 52: 1-183.

Perkins, T.H. 1979. Lumbrineridae, Arabellidae, and Dorvilleidae (Polychaeta), principally from Florida,

I with 465. descriptions of six new species. Proceedings of the Biological Society Washington 92(3) :415-

i Pettibone, M.H. 1957. Endoparasitic polychaetous annelids of the family Arabellidae with descriptions of new species. Biological Bulletin 113(1): 170-187.

Pettibone, M.H. 1963. Marine polychaete worms of the New England region. I. Aphroditidae through

I Trochochaetidae. United States National Museum Bulletin 227:1-356.

Steinhauer, M. and E. Imamura (eds.). 1990. California OCS Phase ITMonitoring Program. Year-Three

Annual Report. Prepared for the U.S. Department of the Interior, Minerals Management Service,

I Pacific OCS Region, under contract No. 14-12-0001-30262.

Uebelacker, J.M. 1978. A new parasitic polychaetous annelid (Arabellidae) from the Bahamas. Journal of

I Parasitology 64:151-154.

Uebelacker, J.M. 1984. Family Arabellidae Hartman, 1944b. Chapter 42 In: Uebelacker, J.M. and P.G.

Johnson (eds.). 1984. Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Barry

I A. Vittor & Associates, Inc., Mobile, Alabama. 7 vols.

Verrill, A.E. 1881. New England Annelida. Pt. 1. Historical sketch, with annotated list of the species hitherto

I recorded. Transactions of the Connecticut Academy of Arts and Science 4:285-324, 12 pls.

Verrill, A.E. and S.E. Smith. 1874. Report upon the invertebrate animals of Vineyard Sound and adjacent

Ill waters, with an account of the physical features of the region. Washington, pp. 1-478, 38 pls. (reprinting

[ I of Verrill, 1873a).

Webster,H.E. 1879. Annelida Chaetopoda of the Vh'giniacoast. Transactions of the Albany Institute 9:202-

I 272, 11 pls.

Webster, H.E. 1884. Annelida from Bermuda collected by G. Brown Good. Pt. 7. Bulletin of the U.S.

National Museum 25:305-327, 6 pls.

I Webster, H.E. 1886. The Annelida Chaetopoda of New Jersey. Reports of the New York State Museum

39:128-159, pls. 4-10 (reprint of No. 32, 1879, with plates included).

I Webster. H.E. and J.E. Benedict. 1884. The Annelida from Provincetown and Wellfleet, Mass. Chaetopoda

Reports of the U.S. Commision of Fisheries, Washington, volume for 1881:699-747, 8 pls.

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13. FAMILY DORVILLEIDAE CHAMBERLIN, 1919

I

by

I Brigitte Hilbig 1

I

i Introduction

The dorviUeidsaremostly small, only afew millimetersinlength, andmost species have been discovered

I and Somdescribed e species ofonlyOphryotrocha inthe last 30hayveearsbeenwheinnvessampl tigatedes weextensively re more routinely because ofsietheir ved trole hrougashpol finelutionmesindicators, h screens.

and quantitative sampling of the deep-sea floor hasresulted in a wealth of new genera and species of unexpected

I mbeocrph auseologtheyicalarediver thesity. onlyAmextant ong the familyEunicwithida,athectenognath dorvilleidsjawareapparatus, consideredi.e.phylogenetically the maxillae consist the oldest of several group

rows of small denticles rather than 4 to 6 pairs of larger pieces. They also differ from all other Eunicida in

I having an even number of prostomial appendages.

I Morphology

a pair of The antennae, prostomium and a pair mayofbepalps. pear-shaped, The antennae conical, mayorberounded, with or without one or two pairs of eyes, simple, papilliform to digitiform, or they may be

biarticulate or moniliform; the palps are simple or biarticulate. The presence or absence, relative lengths, and

I shoratwo pes ociliary f the prostomial bands and appen sometimes dages aren apical of greaciliary t taxonomi tuft.c The importan phrynx ce. Us is eversible ually the prostomium and forms a bears muscular one

proboscis armed with numerous maxillary plates and a pair of mandibles. The jaws differ a great deal in

I compo may besitipresent on amoinntwo, g thefour, dorvilleids or up toand14 rows, axe therefore consistingtheofmost several impoplates rtant taxthatonmay omicbecharacter. heavily sclerotized The maxiland lae

bear large fangs and smaller teeth along the cutting edges, or be more delicate, soft, and bear f'me serrations

I aMeiodorvillea long the cutti, nand g edge. ProtodorviIlea Maxillary (Fig. carriers, 13.2A, located C). dPosterior orsal to the denticles denticlemayrowbes,fused are prtoesent forminbasal Dorvill plates ea,

(Fig. 13.2A); in Ophryotrocha, Exallopus, and Parophryotrocha, the posterior maxillae may be icetongi shaped and much larger than the anterior free denticles (Fig. 13.2B). The maxillae are reduced in anumber of

interstitial forms such Coralliotrocha, Microdorvillea, EIiberidens, and Pseudophryotrocha (Fig. 13.2D),

and generally in deep-sea species. The genus Pettiboneia is characterized by the presence of numerous very

i thin mandibles denticles covered with surficial spines that give the anterior end of the pharynx a rasp-like surface. The are either triangular with several coarse teeth along the cutting edge (Dorvillea and related genera,

Fig. 13.2F), elongate bifid with finely serrated cutting edges (Ophryotrocha, Fig. 13.2G; alsoAnchidorvilIea,

i Fig. 13.2H), or L-shaped with variable shapes and degrees of dentition (Fig. 13.2I).

I 1ENSRConsultingandEngineering,89 WaterStreet,WoodsHole,Massachusetts02543

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The peristomium consists of two achaetous tings, not true segments, that are usually encircled by a I

ciliary band. The setigerous segments are usually ciliated as well, either bearing segmental bands or ciliary

tufts at the parapodial bases. Parapodia are uniramous or subbimmous, with the notopodia consisting of the []

base of the dorsal cirri and a fine acicula extending into the dorsal cirrus (Fig. 13.1L). Pettiboneia possesses il

subbiramous parapodia on a species-specific number of anterior parapodia. The neuropodium may bear a

conical, retractable ventral lobe in addition to the acicular lobe (Fig. 13.3C). Dorsal and ventral cirri are m

generally simple and often very small, and one or both may be absent in small forms. Branchiae are present il

in a few species ofOphryotrocha, Pettiboneia, andDiapharosoma assimple digitiform or cushion-like structures

on the dorsal or ventral side of the parapodium. Notopodia often contain large vessel loops and thus are •

functional gills. II

All setae are neuropodial; they are generally simple above the neuroacicula and compound below,

with the exception of Parophryotrocha that has simple setae only and Ikosipodus that has compound setae []

only. Supraacicular setae include smooth or serrated capillaries and spines and, in some genera, furcate and

geniculate setae (Fig. 13.1A-D). Subacicular setae are homogomph or heterogomph, smooth or serrated, and

the blades may be falcigerous or spinigerous (Fig. 13.1E-G). Sometimes the inferiormost seta in a fascicle is []

a simple capillary or cultriform seta (Fig. 13.1K). In the genus ExaUopus, all or some setae in anterior II

setigers may be greatly modified into recurved hooks (Fig. 13.1H-J).

The pygidium bears a central or slightly dorsal anus and often one or two pairs of anal cirri that may i

be simple or articulated; sometimes a conical dorsal stylus is also present. []

Taxonomic History I

Due to the small size of the dorvilleids, the history of their classification is relatively short, and its I

beginning is marked by confusions and errors. In 1865, Kinberg established the Staurocephalidea, and a

number of genera had been established during the first half of the 19th century for Dorvillea-like forms, most •

of which are now synonymized. Staurocephalus, erected by Grube in 1855, was found to be preoccupied by !1

Verrill (1900), and he proposed Stauronereis (andthe family name Stauronereidae) as a replacement. However,

since he also changed the type species from S. rubrovittatus toS. rudolphi (which had actually been described •

as Nereis rudolphi by delle Chiaje in 1828), he violated an ICZN rule, and the genus Stauronereis became n

invalid. Mclntosh (1910) movedS, rubrovittatus to Dorvillea, a genus established by Parfitt in 1866 with the

type-species Dorvillea lobata, which McIntosh synonymized with Stauronereis rubrovittatus. Dorvillea and •

Ophryotrocha Clapar_de and Mecznikov, 1869, were the only valid genera used for the dorvilleids for nearly U

100 years, with the former containing a mixture of many different forms.

Pettibone's (1961) revision of the family was the beginning of a substantial change in dorvilleid Ill

systematics, even though two of her four genera have since become invalid. In her genetic definitions, she l

relied on setal types and the presence of notopodia, and she removed forms with furcate setae, but without

notopodia from Dorvillea and placed them in a new genus Protodorvillea. In the 1970's, the importance of []

structural details of the jaw apparatus was recognized, mainly by two authors. Orensanz (1973) described a U

new genus with a very unique maxillary structure fully described by Blake (1979), Pettiboneia. Jumars

(1974) was the first to describe deep-sea dorvilleids, a group that since has proven to include a host of genera •

and many unusual species. Two of his new genera, Meiodorvillea and ExaUopus, are still valid, and U

Schistomeringos now exists as a subgenus of DorviUea. Hartmann-Schr6der (1971) established the genus

Parophryotrocha based on the absence of compound setae. At the same time, Akesson had worked extensively •

with shallow-water species of Ophryotrocha, and he found many new species that were quite uniform |

morphologically, but differed in their reproductive behavior and the number of chromosomes (Akesson, 1967,

1973a-b, 1975a, 1976). I

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I During the following decade, important contributions were made by Wolf (1986a-b) and Westheide

and his coauthors (Westheide, 1982; Westheide and Riser, 1983; Westheide and yon Nordheim, 1986; von

i Nordheim, 1987). Wolf divided theDorviUea-like forms into Ougia and Parougia, based on the anatomy of the maxillae, and described some very unusual genera from the Gulf of Mexico, including Westheideia,

Eliberidens, and Diapharosoma. Westheide and his colleagues discovered a number of minute interstitial

i dorvilleids, including Petrocha, Arenotrocha, Pusillotrocha, and Apodotrocha. In 1990 and 1991, the description of deep-sea dorvilleids initiated by Jumars was continued by Hilbig and coauthors (Hilbig and

Ruff, 1990; Hilbig, 1991; Hilbig and Blake, 1991), including the description of two new genera, Anchidorvillea

I and Pseudophryotrocha The publication. of Orensanz' (1990) revision of the eunicemorph polychaetes, including fossil species,

proposed further change in the systematics of the dorvilleids. He thought the split within the family into

I

types Ophryotrocha of ctenognath -like andmaxillae--many Dorvillea-like free specdenticles ies to be insignificant several rows enoughin theto divide Drovilleathe group family.andThelarge, two fused main

proximal pieces and free distal denticles in the Ophryotrocha group--have been separate from each other

i since generathePaIpiphitime Jurassic. Orensanz , Pinniphitime therefore , andproposed Mammiphitime tojoin, Ophryotrocha together with and the dinophilids, related genera, withincluding the Iphitimidae. his new

This proposition is rejected here, mainly because of f'mdings in Hilbig and Blake (1991) of which Orensanz

i was not aware while preparing his revision. In particular, the genus Anchidorvillea would fall between the two families, because it is linked toDorvillea-like forms by itslong antennae and palps, but to the Ophryotrochalike genera by the uniramous parapodia and the arrangement of its maxillae that is reminiscent of P-type

i forceps and a (reduced) number of free denticles. Furthermore, Orensanz (1990) overstated Westheide's (1982) position on the relationship between dinophilids and dorvilleids. Although Westheide's and Westheide

and Riser's (1983) cladograms suggest that the dinophilid genera should be moved to the dorvilleids, these

i authors never actually merged these two families; in fact, in Westheide and Riser (1983), they specifically state the possibility of different evolutionary pathways leading to neotenic species like the dinophilids. It may

be difficult to resolve this issue because neotenic interstitial forms have so few characters that could be used

i to identify them as sister groups to any morphologically complex taxa.

i Distribution and Biological Notes

Dorvilleids are mostly shallow-water forms, although a great number of deep-sea species have been

I found during the last 20 years. Many species are opportunists and flourish in organically enriched environments, such as sewer and pulp mill outfaUs and, in the deep sea, locations of temporal enrichment such as whale

carcasses (Hitbig, personal observations). These opportunistic species are likely to be exposed to pollutants,

i and/_esson (1975b) developed a bioassay method using Ophryotrocha Iabronica as test organism.

The heavily sclerotized, elaborate maxillae of many forms belonging or related to Dorvillea suggests

that they are predatory, and the fact that they are usually captured with empty guts supports that assumption.

I However, occasionally specimens are seen with sediment-filled intestines, and Ophryotrocha can be cultured and raised both with plant matter such as algae or spinach and with small prey such as crustacean larvae

(,2_l_esson,1975a). The rasp-like surface of most denticles of Pettiboneia may be used to graze, much as

I gastropods use the radula.

One of the most interesting features in dorvilleid biology, and probably one of the reasons for their

success, is the plasticity of their reproductive behavior. It has been extensively investigated in the genus

I Ophryotrocha. (1975, 1976) described a number of species that were morphologically similar, but

t_esson

differed in their reproductive cycles; he also divided the genus into several species groups characterized by

I their reproductive mode (gonochoric, hermaphroditic, and viviparous), chromosome numbers, and morphology

i 343
!

of egg cases. Sex determination by environmental factors, a phenomenon known as Paarkultureffekt (pair I

culture effect), was f'n-stobserved for O.puerilis by Hartmann and Huth (1936), and further investigated

mainly in Italy by Bacci and coworkers (see, for example, Bacci & Bortesi (1967)) and in Germany by II

Pfannenstiel and coworkers (see Kegel & Pfannenstiel (1983a, b) for synopsis). Richards (1967) investigated i

the reproduction and development of a species of Dorvitlea, probably D. longicornis, in the laboratory. He

was able to raise two generations to maturity within approximately two years. Eggs and sperm are released I1

directly into the water, and larvae settle and start to feed after 8 days. Recently, Akesson & Rice (1992) I

described two new species of Dorvillea with obligate asexual reproduction. Both species autotomize into

small pieces of about 5-10 setigers which develop new heads and tails within a matter of days. I

Key to the Dorvilleidae 1

The following key can be used not only to identify the species described in this chapter, but to identify l

all known genera. The presentation of such an extensive key seems to be appropriate because of the large 1

number of recently described new genera and species and the likelihood that many previously unreported

genera will eventually be found in California. Ifjaw structures have to be examined andthe specimens are too /

small for dissection, the whole animal should be cleared in 10% KOH for about 30 to 60 minutes or until the I

muscular tissue is macerated away and the body is transparent. The clearing process should be checked every

10 to 20 minutes to avoid loss of the specimen. Details of the setae must be examined under oil immersion, i

1A. Notopodia (= "dorsal cirri" with embedded aeicula) present in at least some setigers (Fig. 13.1L) ..

........................... .............................................................................................................................. 2 Hi

I

i

lB. Notopodia absent; dorsal cirri if present short, never with acicula (Fig. 13.1 M) .......................... 10

2A. Notopodia present throughout body (may be absent on setiger 1); antennae and palps well developed, I

antennae moniliform, palps biarticulate; maxillae in 4 rows, with or without maxillary carders, with

at least 1 pair of basal plates (Fig. 13.2A) ....................................................................................... 3 I

2B. Notopodia with acicutae present on limited number of anterior setigers; antennae and palps well

developed or reduced; maxillae in 2, 4, or numerous rows, consisting of free denticles only .......... 8

l

3A. Maxillae with maxillary carriers and both superior and irfferior basal plates; furcate setae if present

with short tines (Fig. 13.1D) ....................................................................... genusDorvillea .......... 6 i

3B. Maxillae without inferior basal plates; furcate setae if present with long, slender tines (Fig. 13.1C) I

.......................................................................... ............................ .................................. ................. 4

II

4A. Maxillary carriers present ............................................................................................ genus Ougia 2 I

4B. Maxillary carders absent ............................................................................ genus Parougia ......... 5

!

2 Genus not treated in this Atlas. For recent diagnoses or revisions, see Hilbig and Blake 0991) for Anclu'dorvitlea, l_

Exatlopus, Ophryotrocha, and Pseudophryotrocha, Jumars (1974) for Meiodorvillea, von Nordheim 0987) I

for Petrocha, Orensartz (1990) for Pinniptu'time, Petlibone (1961) for Protodorvittea, Wainwright and Perkins

(1982) for Gymnodorviflea, Westheide 0982) for Ikosipodus, Westheide and yon Nordheim (1985) for

Arenotrocha, Coraftiotrocha, Microdorviltea, and Pusitlotrocha, and Wolf (1986a) for Diapharosoma, I

Eliberidens, and Ougia.

|
I Figure 13.1. Dorvilleidae: A, capillary supraacicular setae; B, cultriform supraacicular seta (Ophryotrocha); C,

long-tined furcate seta; D, geniculate and short-tined furcate setae; E, bidentate subacicular setae;

I F, modified spinigerous subaeicular seta (Exallopus); G, falcigerous subacicular seta (Ophryotrocha); H, supraacicular setae (Exallopus); I-J, modified subaeicular setae, heterogomph and

homogomph (Exallopus); K, cultriform ventralmost seta (Ophryotrocha); L, subbiramous

parapodium; L, uniramous parapodium; N, habitus of interstitial species (Arenotrocha). (from various

I sources).

i 5A. Body large (more than 10 mm long), rigid; furcate setae usually present; all setae with serrations at

least distally; maxillae heavily sclerotized, visible through body wall as V-shaped structure; mandibles

triangular, dark ....................................................................................................... Parougia caeca 3

I 5B. Body maxillaesmall reduced, (about transparent, 5 mm long),notfragile; visible furcate through setae body awall; bsent;mandibles all setae L-shaped smooth andwithverytransparent slender;

center (Fig. 13.6C) ..................................................................................................... Parougia batia

I 3 in either Phase 1 Phase 2 material, the is to in the

Although no specimens Wel_ present O1" SpeCieS likely occur area

i 345
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6A. Furcate setae absent (check several parapedia) ................................................ subgenus Dorvillea 2 U

6B. Furcate setae present ...................................................................... subgenus Schistomeringos ..... 7

i

7A. Dorsal cirri tapering, with cirrophores as long as cirrostyles; ventral cirri inserting subdistally; furcate

setae with short fines half as long as long tines; anterior dentictes with straight, finely serrated cutting III

edge ........................................................................................ Dorvillea (Schistomeringos) annulata m

7B. Dorsal cirri cylindrical, distally inflated, with cirrophores much longer than cirrostyles; ventral cirri

inserting distally (may look like subdistal insertion when ventral setal lobe is extended); furcate setae

with short tines one-third as long as long tines (setiger I0); most anterior denticles with crescentic,

wing-like serrated cutting edge and some larger distal teeth ..............................................................

DorviIlea (Schistomeringos) longicornis I

8A. Maxillae in 8 rows, most denticles covered with surficial spines; antennae simple, palps biarticulate,

shorter than antennae, palpophores very short; notopodia slightly longer than neuropodia, present in l

setigers 2 to 12 ............................................................................................... Pettiboneia brevipalpa

8B. Maxillae in 2 or 4 rows, none covered with surflcial spines ............................................................ 9 i

9A. Maxillae in 4 rows, maxillary carriers absent; antennae moniliform, palps biarticulate, palpophores !11

much longer than palpostyles; anterior notopodia with aciculae, posterior ones without aciculae ....

....................................................................................................................... genus Diapharosoma _

U

9B. Maxillae in 2 rows, maxillary carders present; antennae indistinctly articulate, palps biarticulate; II

palpophores about as long as palpostyles; notopodia present in limited number of anterior setigers II

............................................................................................................................. genus Westheideia 2

10A. Antennae and palps well developed, antennae moniliform; maxillae in 2 rows (Fig. 13.2E); ventral I

cirri much longer than dorsal cirri .................................................................. genus Anchidor_iIlea _

10B. Antennae and palps well developed or reduced, antennae never monilfform; ventral cirri always shorter !

than dorsal cirri .............................................................................................................................. 11 u

11A. Maxillae with superior and inferior free denticles and forceps or icetongs formed by fused carriers U

and basal plates (Fig. 13.2B); prostomial appendages and parapodial cirri present or absent, well

developed or reduced ...................................................................................................................... 13 I

liB. Maxillae in 2 or 4 rows, without forceps or icetongs ..................................................................... 12

12A. Maxillae in 2 rows ......................................................................................................................... 20 I

12B. Maxillae in 4 rows, with superior and inferior free denticles, superior and inferior basal plates, and

maxillary carriers (some elements may be reduced); antennae and palps well developed or reduced i_

...................................................... . ............................................................................................. ,°. 15

!

|
I

i Figure 13.2. Dorvilteidae: A-E, maxillae; A, with maxillary carriers, two pairs of basal plates, and and four rows of free denticles; B, with forceps and seven pairs of free denticles in four rows (Ophryotrocha);

C, with two rows of denticles; D, reduced to smooth, soft plates (Pseudophryotrocha); E, modified,

woth two rows of denticles (Anchidorvillea); F-J, mandibles; F, DorviUea; G, Ophryotrocha; H,

I Anchidorvillea SOUrCeS). ; I, modified, L-shaped (Ophryotrocha); J, rodlike (Pseudophryotrocha). (from various

I

i 347
!

!

13A. All setae simple; prostomium wider than long, with well-developed clavate antennae and palps; median i

and posterior setigers with dorsolateral and ventrolateral segmental lobes; setae including smooth

spines and free capillaries ................................................................... Parophryotrocha brevicapitis

13B. Supraacicular setae simple, subacicular setae compound (ventralmost seta may be simple) ........ 14 i

14A. Some or all setae in anterior setiger(s) greatly modified into recurved hooks (Fig. 13.1H-J) ............ /

................................................................................................................................ genus Exallopus 2 l

14B. Anterior setae if modified only slightly different from regular setae, never recurved; prostomial

appendages and parapodial cirri usually short and simple ................................ genus Ophryotrocha 2 i

15A. Maxillae consisting of basal plates only; antennae and palps short, digitiform ................................. •

.............................................................................................................................. genus Eliberidens 2 II

1513. Maxillae including free denticles .................................................................................................. 16

i

16A. Minute interstitial forms, about 1 mm long, with maximally 15 setigers (Fig. 13.1N) ................. 17

16B. Animals not interstitial, adults several millimeters long; antennae papilliform, palps multiarticulate, i

much longer than antennae; maxillary apparatus well-developed .................. genus Protodorvillea 2 II

17A. Maxillae with superior and inferior basal plates and superior and inferior free denticles .............. 18 I

17B. Maxillae with superior basal plates and superior and inferior free denticles; antennae simple, palps

biarticulate, palpophores as long as palpostyles; all supraacicular setae simple spines .................... •

....................................................................................................................... genus Microdorvillea 2 If

• !

18A. Antennae moniliform, palps biarticulate, with long palpophore; supraacicular setae including furcate

setae with long, slender tines (Fig. 13.1C) ...................................................... genus Coralliotrocha 2

18B. Antennae simple or absent; furcate setae absent ........................................................................... 19 i

i

19A. Setae including serrated capillaries and compound falcigers with serrated shaft and blade; prostomium

with simple palps, antennae absent; parapodia without cirri; mandibles ornate; maxillae with at least

2 pairs of free denticles ............................................................................................. genus Petrocha 2

19B. Both simple setae and blades of compound falcigers unidentate; capillaries serrated, compounds

smooth; prostomium with simple palps, antennae absent; maximally 18 setigers, parapodia without i

cirri ................................................................................................................... genus Pusillotrocha 2

i

20A. Maxillae consisting of 3 pairs of plates, the posteriormost pair basally fused or linked ............... 21 i

20B. Maxillae consisting of serrated, rounded free denticles ................................................................. 22

!

!

348 I
!

!

I 21A. Maxillary plates smooth, elongate, weakly sclemtized (Fig. 13.2D); mandibles rodlike or L-shaped (Fig. 13.2J); furcate or geniculate setae absent ........................................ genus Pseudophryotrocha 2

2lB. Maxillary plates dentate, oval, heavily sclemtized; mandibles subterrninally fused along symphysis

I (Fig..2K) .................................................................................................... genus Pinniphitime 2'

13

4

I 22A. Small, interstitial forms with reduced prostomial and parapodial appendages .............................. 24 22B. Adults several millimeters long, not interstitial .............................................................................. 23

I 23A. Maxillary carriers absent; supraacicular setae including serrated capillaries, fumate setae with short

tines (anterior setigers), and geniculate setae (median and posterior setigers); inferiormost subacicular

i 23B. Msetaaxillary cultriform; cardersantennae presentand(Fig. palps13.2C); absentsupraacicular ................................................... setae including capillagenus ries Gymnodorvillea and furcate setae2

with short tines, occasionally replaced by geniculate seta in one or few anterior parapodia (Fig.

I 13.1D); antennae and palps present (palps may be absent) ............................ genus Meiodorvillea 2

24A. All setae compound; maximally 10 setigers, parapodia lacking cirri; prostomium with palps, antennae

i eyes ..................................................................................................... genus

and absent Ikosipodus 2

24B. Supraacicular setae simple, serrated, bidentate; compound falcigers with smooth, distally bidentate

blades; up to 10 setigers, parapodia without cirri; prostomium with digitiform antennae and thicker

I paIps equal length, eyes absent; organs pads .............. genus

of nuchal with 4 ciliated Arenotrocha 2

!

. Description of Species

As a result of the many recent publications, the Dorvilleidae have become a large family with well

I over 100 species in 22 genera. The dorvilleid fauna in the Santa Mafia Basin and Santa Barbara Channel is depauperate with only five species, none of which belong to Ophryotrocha. The following species are treated:

I Dorvillea (Schistomeringos)annulata (Moore, 1906)

DorviIlea (Schistomeringos ) longicornis (Ehlers, 1901)

I Parophr Parougiayotbatia rocha(Jumars, brevica1974), pitis Hilbig, new comb newination species

Pettiboneia brevipalpa Hilbig and Ruff, 1990

!

!

4TheothertwogeneradescribedbyOrensanz(1990),PalpiphitimeandMammiphitimea,reconsidereddoubtfulandare

I thereforenotincludedin thiskey.

i 349
I

!

Genus DorviIlea Parfitt, 1866 I

Type Species. Staurocephalus rubrovittatus Grebe, 1855 I

Diagnosis. Parapodia subbiramous throughout (except for setiger 1), with notopodium consisting of

base of dorsal cirrus and acicula projecting into the cirrostyle; setae including simple capillaries and heterogomph •

falcigers (subgenus Dorvillea) or capillaries, furcate setae with short tines, and heterogopmph falcigers I

(subgenus Schistomeringos). Antennae and palps well-developed, antennae usually moniliform, palps

biarticulate. Maxillae heavily sclerotized, visible through body wall, consisting of maxillary carriers and 4 i

rows of denticles, the posterior ones fused to form superior and inferior basal plates; mandibles dark, triangular, III

with few large teeth along cutting edge and often several lateral free conical teeth.

Remarks. Thanks to the revisions by Pettibone (1961) and Wolf (1986a-b), the formerly ill-defined !

genus now has a clear and concise definition. It appears that true Dorvillea are limited to shelf depths,

u

whereas species from deep water are assigned to Ougia and Parougia. !

DorviUea (Sehistomeringos) annulata (Moore, 1906)

Figure 13.3 I

Stauronereis annulatus Moore, 1906:225-227, pl. 10, figs. 12-13, pl. 11, figs. 18-22. im

Stauronereis rudolphi: Pettibone, 1963:231-233, fig. 60a-f (in part). |

Dorvillea annulata: Fauchald, 1970:152-154.

Schistomeringos nr. annulata: Lissner et al., 1986:D-12. m

Schistomeringos longicornis: Lissner et al., 1986:D-12 (in part). 1

u

Material Examined. California: Santa Maria Basin, offPoint Buchon, Sta. 6 (1), Sta. 13 (2). i

!

Description. Length to 13 mm, width to 1 mm, segments to 72. Body slender, dorsoventrally

compressed, with conspicuous parapodia, widest in pharyngeal region, gently tapering toward pygidium within

last 10 setigers or so. Color in alcohol light tan.

Prostomium rounded, somewhat longer than wide, with antennae and palps of equal length and about

twice as long as prostomium. Antennae with 7 to 8 moniliform articles, palps with long patpophore and short, m

conical palpostyle. Two pairs of eyes near antenna] base, side by side or arranged in a square; anterior (inner)

pair large, posterior (outer) pair minute spots (Fig. 13.3A). Proboscis with well-developed mandibles and

n

maxillae. Maxillae with smooth to finely serrated carriers, posteriorly fused or free from one another; superior

basal plates large, with 6 to 7 large teeth alternating with smaller ones; superior free denticles about 30; E

proximal denticles with narrow base and wide cutting edge beating large main fang and several smaller lateral

I

teeth; remaining free denticles gradually becoming elongate, more delicate plates, main fang diminishing in

size and eventually completely absent, cutting edge with several f'me, long teeth. Inferior basal plates smaller

and more delicate than superior ones, with about 5 large fangs alternating with several smaller teeth. Inferior I

free denticles also numbering about 30, resembling corresponding superior ones, but generally less sclerotized m

(Fig. 13.3B). Mandibles large, anteriorly flared, with about 10 teeth on cutting edge and 4 to 8 free conical •

teeth (Fig. 13.3C). m

Parapodia uniramous in setiger 1, subbiramous from setiger 2, with notopodium reduced to base of n

dorsal cirrus and f'me acicula projecting into cirrostyle; with rounded acicular lobe, conical postsetal lobe, and •m

retractable ventral lobe. Ventral cirri inserting subdistally, projecting about as far as postsetal lobe; dorsal

cirri slender, with conical cirrostyle about as long as cirrophore (Fig. 13.3D). Supraacicular setae including ,,

l

350 |
I

!

i ao

i

, i

i

I infBp P

!

mC

!

!

II

!

!

I Figure 13.3. Dorvillea (Schistomeringos) annulata: A, anterior end, dorsal view; B, maxillae; C, mandibles; D,

middle parapodium, ventral setal lobe extended; E, subacicular setae, shortest and longest blade of

I fascicle; F, furcate supraacicular seta; G, capillary supraacicular seta (after Fauchald, 1970).

i 351
I

2 to 3 finely serrated capillaries and usually one or two furcate setae with long tines about twice as long as i

short fines; shaft serrated subdistally below shorter tine (Fig. 13.3F,G). Subacicular setae numerous compound

setae with bidentate blades of varying length; longest blades on dorsalmost setae, about 3 times as long as i

shortest blades (Fig. 13.3E). l

Remarks. The largest Phase I specimen examined was originally identified as Schistomeringos nr.

annulata, probably because of the differences in the jaw apparatus to Fauchald's (1970) redescription. However,

since characters such as the number of free denticles and the number of teeth on a denticle or basal plate II

appear to be growth-dependent (Hilbig, personal observations), the only remaining discrepancy is the shape of

the maxillary carriers, which were found to be fused and serrated in the specimen at hand, but were described i

as being smooth and posteriorly free from one another by Fauchald (1970). All other characters, such as the |

proportions of the pampodia and setae, agree very well with Moore's (1906) original description, and it seems

unlikely that two species would differ only in details of the maxillary carriers. The specimen is therefore i

considered to be D. annulata.

Type Locality and Type Specimens. Quarantine Rock, Port Townsend, Washington: holotype (USNM

5541) and paratype (ANSP 1980). I

1

Habitat. Silt, silty and shelly sand.

Distribution. Washington to western Mexico, 80 - 220 m. i

If

DorviUea (Schistomeringos) longicornis (Ehlers, 1901)

Figure 13.4 i

Stauronereis longicornis Ehlers, 1901:150-151, pl. 19, figs. 18-21, pl. 20, figs. 4-6. i

Stauroneries articulatus Hartman, 1938:101-102, figs. 39-44. i

Dorvillea articulata: Hartman, 1968:817, figs. 1-5.

im

Stauronereis rudolphi: Pettibone, 1963:231-233, fig. 60a-f (in part).

Dorvillea rudolphi: Fauchald, 1970:156-159, pl. 27, figs. a-j (in part), i

Dorvillea atlantica: Hartman, 1968:819, figs. 1-5.

m

Schistomeringos longicornis: Jumars, 1974:107-109, fig. 2.--Lissner et al., 1986:D- 12. i

Schistomeringos annulata: Lissner et al., 1986:D-12 (in part), i

Material Examined. California: Santa Mafia Basin, off Point Sal, Sta. 104 (1), Sta. 103 (1); off

Purisima Point, Sta. R4 (1). B

u

Description. Length to 70 mm, width to 1 mm, segments to 80. Body stout, with rounded dorsum

and slightly flattened ventrum, widest in middle, tapering toward pygidium and slightly toward prostomium. Hi

Color in alcohol uniformly light tan. i

Prostomium bluntly rounded, longer than wide, in alcohol sometimes divided into 2 or 3 tings, bearing

2 pairs of eyes, the anterior pair largest; antennae somewhat longer than prostomium, moniliform, with 5 to 12 in

articles; palps about as long as antennae, with long palpophore and short palpostyle (Fig. 13.4A). Maxillae |

with fused, serrated carriers; superior basal plates fused posteriorly, with about 14 large fangs alternating

with smaller teeth (Fig. 13.4B); superior free denticles about 30 to 40 elongate plates with cutting edges i

bearing different kinds of dentitions from proximal to distal position within row; denticles close to basal plate |

with comb-shaped cutting edge bearing several equal teeth (Fig. 13.4C); more distal denticles with main fang

and small lateral teeth (Fig. 13.4D, E); main fangs gradually diminishing in size, cutting edges becoming i

crescentic with progressingly freer dentitions, distalmost denticle soft, with delicate, rounded teeth (Fig. 13.4F, i

G). Inferior basal plates free, smaller than superior ones, also bearing about 14 large fangs and several

smaller teeth; about 40 to 50 inferior free denticles present, similar in shape to corresponding superior denticles, I

352 i
I Figure 13.4. Dorv base illplate, ea (Sdetail chistomering of proximal os) long teeth; icornC-G, is: superior A, anteriofree r enddenticles , dorsal fromviewproximal (eyes fadetoddistal: ); B, sC, uperi veryor

close to base plate, D-E, middle of superior row, F-G, first and fifth denticle; H-J, distal inferior

I free denficles; K, parapodium, anterior view, ventral setal lobe extended; L, same, ventral setal lobe retracted; M, furcate supraacicular seta; N, subacicular seta. (D, E, G after Fauchald, 1970).

i 353
!

!

generally somewhat more delicate; distal inferior denticles soft, oblong plates with numerous very fine teeth m

(Fig. 13.4H-J). Mandibles heavily sclerotized, flared anteriorly, with about 10 teeth on cutting edge and i

several free conical teeth. INN

Parapodia uniramous in setiger 1, subbiramous from setiger 2 (Fig. 13.4K, L); with low presetal t

lobe, conical postsetal lobe, and retractable ventral lobe. Ventral cirri inserted distally, insertion appearing |

subdistal when ventral parapodial lobe fully extended (Fig. 13.4K); dorsal cirri with long cylindrical cirrophore

and short, conical, slightly inflated cirrostyle, extending about as far as postsetal lobe. Supraacicular setae am

including serrated capillaries and furcate setae with subdistally serrated shafts and unequal tines; around i

setiger 10, tong tine about 3 times as long as short tine (Fig. 13.4M). In anterior setigers occasionally one of

the furcates replaced by bidentate geniculate seta. Subacicular setae compound falcigers with bidentate, m

serrated blades of varying length, dorsalmost setae with longest blades, up to 4 times as long as shortest

blades in same fascicle (Fig. 13.4N).

m

Pygidium with 2 short ventral and 2 longer ventrolateral anal cirri. Im

Remarks. Among thePhase I voucher material, some specimens were misidentified as Schistomeringos II

annulata, most likely because the overall shape of the parapodia resembles that species when the ventral lobe

is extended; the ventral cirrus then seems to arise from the middle of the ventral parapodial edge rather than R

distally. However, the shape of the dorsal cirrus is a good character to discriminate between those two II

species; the cirrostyle is slender and as long as the cirrophore in D. annulata, but much shorter than the

cirrophore and somewhat inflated in D. longicornis. The cresecentic, wing-like cutting edges of the anterior Ill

free denticles are also characteristic for D. longicornis and can be easily observed through a short dorsal II

incision just below the prostomium. The length ratio between the shorter and longer tines of a furcate seta

from setiger t0, recommended for species discrimination between D. longicornis and D. rudolphi by Jumars Im

(1974), can also be used to distinguish D. longicornis from D. annuIata; the ratio is about 2 forD. annulata |

and 3 for D. longicornis. However, several setae should be examined because this character is somewhat

variable. n

Type Locality and Type Specimens. According to Jumars (1974), Ehlers' type specimens have to J

be considered lost. Type locality is Chile.

Habitat. The species is found in mixed sediments, mud with high silt content, and ooze. I

IN Distribution. Alaska to Chile, intertidal to 575 m.

Genus Parophryotrocha Hartmann-Schrrder, 1971 I

Type Species. Ophryotrocha isochaeta Eliason, 1962 I

Diagnosis. Parapodia uniramous, with simple setae of one or two kinds in both supra- and subacicular IN

fascicles; compound setae absent. Antennae and palps if present short and simple. Maxillae with icetong- II

shaped posterior pieces, formed through fusion of maxillary carriers and superior and inferior basal plates,

and usually 7 pairs of free denticles in 4 rows. II

Remarks. This genus is very similar to Ophryotrocha except for the absence of compound falcigers II

in the subacicular fascicle.

!

!

354 I
!

I Parophryotrocha brevicapitis Hilbig, new species

Figure 13.5

I Dorvilleidae sp. A Hyland et al., 1990:F-1.

I 120°49.16'W, Materi9Ial m:Examholotype ined. (USNM California: 170427). Santa Maria Basin, off Point San Luis, Sta. R-l, 35°05.8YN,

Description. Length 2 mm, width 0.2 mm, segments 19. Body long, slender, widest in pharyngeal

I region, gently tapering toward pygidium. Median and posterior setigers drawn out to rounded dorsotateral and ventrolateral lobes extending over the parapodial bases, ventral lobes largest (Fig. 13.5A, B). Ciliation

including ring on prostomium between antennae and palps and tufts at dorsal and ventral parapodial bases

I andalong edge of lateral segmental lobes. Color in alcohol pale.

Prostomium bluntly triangular, wider than long, with two digitiform antennae arising from ridge

running across middle of prostominm and large biarticulate palps inserted at postectal comers of prostomium;

I cirrophores short, cylindrical, bases as wide as lateral edges of prostomium; cirrostyles slender, digitiform

(Fig. 13.5A). Jaws observed through body wall only; maxillae with P-type forceps, weakly sclerotized, and

probably 7 pairs of free denticles arranged as typical for the genus; mandibles L-shaped, anteriorly flared,

I with smooth cutting edge, also weakly sclerotized except for median rod-shaped handles. Peristomium with 2

equal rings, similar to subsequent setigers.

i Parapodia simple, with long, slender base and conical acicular lobe; dorsal and ventral cirri absent, but ciliated pad present just above insertion of dorsalmost seta; no retractable parapodial lobes observed (Fig.

13.5C). Setae all simple, in fascicles of 4 or 5, of 2 kinds: (1) 3-4 slender, smooth, f'me-tipped spines, (2) in

ventralmost position single, very thin capillary (Fig. 13.5D, E).

I Pygidium unknown.

Etymology. The species is named for its characteristically short head (brevis= Latin for short,

I caput=-Latin for head). Remarks. Although only one incomplete specimen was available for examination, the establishment

of a new species is justifiable because of the very unusual external appearance of this animal. The other two

I known species ofParophryotrocha, P. isochaeta and P. gesae, resemble typical shallow-water Ophryotrocha species, whereas P.brevicapitis is most similar to a group of somewhat aberrant species within Ophryotrocha

including four provisional SCAMIT species from polluted environments in California and the hydrothermal

I vpalps ent species and the Oheavily . platykephale ciliated. segmental The mostlobes charaabove cteristicandfeatures below theof median P. brevicap anditisposterior are theparapodia. large biarticulate

Distribution. Only known from the type locality.

I

Genus Parougia Wolf, 1986

I Type Species. Schistomeringos nigridentata Oug, 1978

I Diagnosis. Parapodia subbiramous throughout except for setiger 1, with notoaciculae projecting into dorsal cirrostyles; supraacicular setae simple capillaries, in some species also 1 or 2 furcate setae with

long fines; subacicular setae compound falcigers or spinigers. Antennae and palps well-developed; antennae

I simple or moniliform, palps biarticulate. Maxillae with superior basal plates and superior andinferior free denticles; maxillary carders and inferior basal plates absent. Mandibles dark, triangular, with conical teeth

along cutting edge, or more delicate, wing-shaped, and smooth anteriorly.

!

i 355
Figure 13.5. Parophryotrocha brevicapitis Hilbig, new species: A, anterior end and last setigers of fragment, I

dorsal view; B, same, ventral view; C, parapodium; D, spines; E, ventralmost capillary seta. (all

from holotype). I

I

356 |
!

I Remarks. Parougia contains some species formerly assigned to Schistomeringos as defined by

Jumars (1974). The presence of furcate setae seems to be a fairly inconsistent character; for example, Parougia

I certain caeca, one portionof theof most the body common (sometimes species only in thison genus, one side), is found or withwithnone furcate at all. setaeThein inconsistency all parapdia, along of thisa

character was one of the initial reasons to restrictSchistomeringos (as a subgenus of Dorvillea, see above) to

i those forms that possess short-tined furcates which occur regularly in all parapodia.

Parougia batia (Jumars, 1974), new combination

I Figure 13.6

DorviIlea batia Jumars, 1974:115-117, fig.6.--Hyland et al., 1990:F-1.

Dorvillea sp. 2: Blake et aI., 1987:C-2.

Material Examined. California: Santa Mafia Basin, off Point Sal, Sta. R-7 (2). North Carolina:

I Hatteras, MMS ACSAR Sta. $9, 35°28.3'N, 74°47.6"vV,604 Lookout, MMS ACSAR Sta. Cape m (6); Cape

S1, 34°16.2'N, 75°45.8'W, 583 m (1).

Description. Length to 3.3 mm, width to 0.3 ram, segments to 44. Body slender, dorsoventrally

I flattened; color in alcohol pale.

Prostomium bluntly rounded, wider than long, with antennae arising close to posterior margin at

I outer postectal edgesmargin of central of prostomium transverse ridge; and tapering, palps slightly cirriformshorter palpostyles than antennae, (Fig. 13.6A); with sheyes ort palpophores absent. Peristomium located at

with 2 rings, the first one much shorter and narrower than the second one, bearing large, bilobed, heavily

I ciliated altematingnuchal withorgans. several Maxillae small teeth,weakly fusionsclerotized, lines of original reduced; denticles superior visible basalas plates typicalfused, for thewith genus; 5 large 5 superior fangs

free denticles present, all but distal one with main fang and long serrated cutting edges below main fang; distal

I superior oblong plates denficlewithredrounded uced to very cuttingthin,edge finelybearing serratednumerous plate. Inferior very fine rowteeth consisting (Fig. of 13.6B). 5 free denticles, Mandiblesallwing- soft,

shaped, with short posterior handle and medially serrated cutting edge; center of mandible soft, transparent,

i margins and cutting edge relatively heavily sclerotized and visible through body wall even in dorsal view (Fig.

13.6C).

Parapodia uniramous in setiger 1, subbiramous from setiger 2; neuropodia conical, with retractable

I ventral cirri about lobe,asbearing long aslarge neuropodia, setal fascicles indistinctly arrangeddivided in broadintofan; cirrophore ventral cirri and short, inslightly serted subdistally; inflated cirrostyle dorsal

(Fig. 13.6D). Supraacicular setae numerous smooth capillaries, subacicular setae compound spinigers with

I blades 13.6E-G). of varying lengths, shortest blades almost falcigerous; shafts with very few serrations distally (Fig.

Pygidium with two articulated anal cirri and middorsal style.

I Remarks. Unlike typical members of this genus,P, batia does not preserve well, and proportions of the soft body parts, such as prostomial appendages and parapodial cirri, vary somewhat depending on the

condition of the specimens. The soft,small body,smoothsetae, indistinctlyarticleddorsal cirri,andcharacteristic

I outline of the mandibles readily distinguish this species from its congeners and from co-occurring species of Dorvillea. An examination of the maxillae in situ after clearing in 10%KOH revealed that maxillary carriers

and inferior basal plates areabsent, and consequently the species is transferred to Parougia. Jumars, in his

I original description, had interpreted the superior basal plates as being afusion product of the carriers and both pairs ofbasal plates. P. bat/a belongs to agroup of deep-water species within thegenus that areall characterized

by reduced, weakly sclerotized maxillae and mandibles of aberrantshapes. At least one Atlantic and one

I Pacific species are still awaiting description.

i 357
I

I

..... l I

/ I

" . - . ..

•._:.i !

:-.:..i_.. . . . . ....i:,._. _

'' .... I

• ;' ..,.,:_

" I -- |

,

supBp

_eAc "_-

D \ \ I setLo

I

I

Eigure 13.6. Parougia batia: A, anterior end, dorsal view; B, maxillae; C, right mandible (after Jumars, 1974);

seta;D' parapodium,G, supraacicularanteri°rseta.View; E-F, two upper, short-bladed and one lower, long-bladed subacicular I

I

358 i
!

I Type Locality and Type Specimens. San Diego Trough, Califomia: holotype (LACM-AHF Poly

1074) and 4 paratypes (LACM-AHF Poly 1075, 1076).

I epifaunaBiofoloastrorhizid gy. Parougia foraminiferans. batia was foundIt isinpossible silty mud;thatJumars P. batia(1974) feedssuggested in a similar that thewayspecies on epifauna grazes oofn

another large foraminiferan characteristic for the slope off Cape Hatteras, Bathysiphonfiliformis.

I 560 Distribution. Central and southern California; North Carolina, Cape Hatteras to Cape Lookout, -1229 m.

I Genus Pem'boneia Orensanz, I973

i Type Species. Pettiboneia sanmatiensis Orensanz, 1973

Diagnosis. Parapodia subbiramous in a limited number of anterior setigers (usually about 8 to 12),

with notoaciculae projecting into dorsal cirrostyles; median and posterior parapodia uniramous. Setae including

I simple capillaries geniculate or supraacicular compound falcigers

and furcate setae in fascicles and in

subacicular fascicles. Antennae simple, palps biarticulate. Maxillae in 8 to 22 rows, with flee denticles only;

most denticles weakly sclerotized, rounded, rasplike plates densely covered with surficial spines. Mandibles

I delicate, wing- or L-shaped, with scalloped cutting edge.

Remarks. The often spectacular maxillae, although usually visible only after clearing, and the presence

i of notopodia in a defined number of anterior setigers makes this genus easy to recognize. To date, three species of Pettiboneia have been reported from California: the type species P.sanmatiensis that was found in

Tomales Bay, P. dibranchiata with the type locality in the Santa Catalina Basin, and P brevipalpa which was

found during Phase I and is described below. Descriptions of the other two species can be found in Blake

I (1979) and Blake and H_ilbig(1990).

I Pettiboneia brevipalpa Hilbig and Ruff, 1990

Figure 13.7

I Pettiboneia brevipalpa Hilbig and Ruff, 1990:115-118, fig. 1.

Pettiboneia sanmatiensis Lissner et al., 1986:D-12. Not Orensanz, 1973.

I Material Examined. California: Santa Mafia Basin, off Point San Luis, Sta. 28 (1); Pioneer Canyon, Sta. C-3, 37°20.21'N, 123°08.84'W, 1400 m (1); north of Pioneer Canyon, Sta. 3-4, 37°27.83'N, 123° 11.60%V,

1040 m(1), Sta. 3-5, 37°27.39'N, 123°14.42%V, 1195 m(2), Sta. 3-10, 37°26.00'N, 123°10.83"_V,985 m(1),

I Sta. 3-11, 37°25.50'N, 123°15.02'W, 1225 m (3), Sta. 3-15, 37°24.71'N, 123°12.16'W, 1010 m (3), Sta. 3- 16, 37°23.20'N, 123°14.44_N, 1220 m (2),Sta. 3-17, 37°22.18'N, 123°16.76'W, 1457 m (1); south of Pioneer

Canyon, Sta. 4-2, 37°15.29'N, 123°07.13'W, 995 m (1).

I Description. to 4.5 width to 0.1 to 66. Body slender,fragile, unpigmented Length mln, ram, segments

in alcohol.

i Prostomium pear-shaped, about as long as wide (Fig. 13.7A); antennae clavate, about as long as greatest prostomial width; biarticutate palps half antennal length, inserted ventrolaterally behind antennae,

with short, inconspicuous palpophores; eyes absent; narrow band of cilia encircling prostomium between

antennae and palps; large yellowish-brown nuchal organs on posterior margins of prostomium. Maxillae with

I two main rows of free denticles and about six additional rows on each side; maxillary carriers and basal plates

absent (Fig. 13.7F). Basal teeth of main rows smooth, delicate, rounded plates with serrated cutting edge;

Ii

I

i 359
I

Figure 13.7. Pettiboneia brevipafpa: A, anterior end, dorsal view; B, anterior parapodium; C-D, capillary and •

fln'catesuprancicu1_setaeE;, subacicularsereF,; maxillae;G, mandibles,(fromHilbig& Ruff,

I

1990).

I

36o I
I

i middle and upper teeth rounded plates covered with surficial spines. Teeth of additional rows including

smooth, anteriorly serrated plates proxirnally and spinose plates distally. Mandibles elongate, slightly curved,

I anteriorly flared, smooth and weakly incised (Fig 13.7G). Two subequal asetigerous peristomial rings, shorter than setigerous segments; complete ciliary bands on both rings and on anterior setigers.

Cirrifonn notopodia with embedded acicula present from setiger 2 through sefiger 8-12, as long as or

I slightly longer than neuropodial acicular lobe. Neuropodia with conical acicular lobe and inferior retractable

setal lobe supported by inferiormost seta (Fig. 13.7B); ventral cirri short, cirriform. Supraacicular setae

including 1 to 3 serrated capillary setae (Fig. 13.7C) and 1 (occasionally 2) furcate seta with unequal tines

I and 1-2 of fine serrations below shorter fine with delicate inner side 13

roWS tine; longer wing on (Fig. o7D),

Subacicular fascicle with 2-4 compound setae having unidentate, finely serrated blades of varying lengths;

shafts bifid with coarse subdistal serrations (Fig. 13.7E). Far posterior setigers occasionally with simple

I pointed inferior setae.

Pygidium rounded, longer than preceding setigers, with 2 pairs of clavate subterminal cirri.

I P. sanmatiensis Remarks.Orensanz, To date,1973, three species P. dibranchiata of Pettiboneia (Armstrong have been and reported Jumars, from 1978),California: and P. brevipalpa the type species Hilbig

and Ruff, 1990. Pettiboneia brevipalpa is similar toP. sanmatiensis in the number of notopodia, the shape of

I the easilpros y ditomium, the number of maxillary rows, and the size and shape of the pygidial cirri. However, it is stinguished from the latterspecies by the palps that aremuch shorterthanthe antennae, withpalpophores

shorter than the palpostyles, tn addition, P brevipalpa lacks eyes, andthenotopodia are longer in relation to

I the neuroType podiaLothan calityin Pand.sanmatien Type Speeir sis. nens. Boca de Quadra, Alaska: holotype (USNM 130084) and 15

paratypes (USNM 130086 - 130091), 3 paratypes (BMNH ZB 1990.29-31), 1 paratype (ZMH P-20325);

I Smeaton Bay, Alaska: 1 paratype (BMNH ZB 1990.32). Biology. Hilbig and Ruff (1990) reported the occurrence of mature males and females in Alaska

between April and July. There are up to 8 round or elongated eggs per setiger averaging 40 im in length and

I 27 im in width. The eggs are first present between setigers 14 and 21, and the sperm appear between setigers 15 and 19. The gametes are associated with the bases of the parapodia and continue for most of the length of

the worm. The species is found in f'me silts and silt mixed with sand.

I Distribution. Alaska to central California, 140 - 1457 m.

I Literature Cited

I _&esson, B. 1967. On the biology and larval morphology ofOphryotrochapuerilis Clapar&te & Metschnikov

(Polychaeta). Ophelia 4:111-119.

I Akesson, B. 1973a. Morphology andlife history ofOphryotrocha maculata sp.n. (Polychaeta, Dorvilleidae).

Zoologica Scripta 2: 141-144.

I _kesson, B. 1973b. Reproduction and larval morphology of five Ophryotrocha species (Polychaeta, DorviUeidae). Zoologica Scripta 2: 145-155.

I /_dcesson, della B. Stazione 1975a. Reproduction Zoologica Napoli in the39genus Suppl.:Ophryo 377-398. trocha (Polychaeta, Dorvilleidae). Pubblicazioni

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._&esson, B. 1975b. Bioassay studies with polychaetes of the genus Ophryotrocha as test animals. In: •

Koeman and Strik (eds.), Sublethal effects of toxic chemicals on auqatic animals, Elsevier 1975, pp.

121-135. I

]kkesson, B. 1976. Morphology and life cycle of Ophryotrocha diadema, a new polychaete species from

California. Ophelia 15(1): 23-35. i

Bacci, G. and O. Bortesi. 1967. The reestablishment of sex balance in Ophryotrocha puerilis through i

interactions between individuals from arrhenogenous and thelygenous lines. Nature 209:448-449.

Blake, J.A., B.Hecker, J.E Grassle, B.Brown, M. Wade, P.D. Boehm, E. Baptiste, B. Hilbig, N. Maciolek, R. i

Petrecca, R.E. Ruff, V. Strarczak, and L. Watling. 1987. Study of biological processes on the U.S. i

South Atlantic slope and rise. Phase 2. Final Report. Prepared for the U.S. Department of the Interior,

Minerals Management Service, Washington, D.C., under Contract No. 14-12-0001-30064. 415 pp + i

Appendices A-M. NTIS No. PB87-214-359. i

Chiaje, S. delle. 1828. Memorie sulla storia e anatomia degli animal] senza vertebre del regno di Napoli 3: 1- •

232. Naples. Ii

Ehlers, E. 1901. Die Polychaeten des magellanischen und chilenischen Strandes. Ein faunistischer Versuch.

In: Festschrift zur Feier des hundertffinfzigjahrigen Bestehens der ktniglichen Gesellschaft der B

Wissenschaften zu Gtttingen (Abh. Math.-Phys). Wiedemannsche Buchhandlung, Berlin, 232 pp. i

Fauchald, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae, •

Lysaretidae and Dorvilleidae from western Mexico. Allan Hancock Monographs in Marine Biology I!

5:1-335.

Grube, A.-E. 1855. Beschreibungen neuer oder wenig bekannter Anneliden. Archly fiir Naturgeschichte R

21(1): 81-128.

Hartman, O. 1938. Descriptions of new species and new generic records of polychaetous annelids from i

California of the families Glyceridae, Eunicidae, Stauronereidae and Opheliidae. University of i

California Publications in Zoology 43: 93-112.

Hartman, O. 1968. Atlas of errantiate polychaetous annelids from California. Allan Hancock Foundation, i

Los Angeles, University of Southern California, 828 pp.

Hartmann-Schrtder, G. 1971. Annelida, Borstenwiirmer, Polychaeta. Tierwelt Deutschlands 58: 1-594. •

II Hilbig, B. and J.A. Blake. 1991. Dorvilleidae (Annelida: Polychaeta) from the U.S. Atlantic slope and rise.

Description of two new genera and 14 new species, with a generic revision ofOphryotrocha. Zoologica

Scripta 20(2): 147-183. i

i

Hilbig, B. andR.E. Ruff. 1990. Remarks on the genus Pettiboneia (Polychaeta: Dorvilleidae) with descriptions

of two new species. Bulletin of the Southern California Academy of Sciences 89(3): 115-123. •

Hyland, |J., E. Baptiste, J. Kennedy, J. Campbell, R. Kropp, C. Robinson, and S. Williams. 1990. Macroinfaunat

Assemblages in the Santa Maria Basin offthe Coast of Southern California. Chapter 7 In: California

OCS Phase II Monitoring Program. Year-Three Annual Report (M. Steinhauer and E. Imamura, •

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Region, under contract No. 14-12-0001-30262.

Jumars, P. 1974. A generic revision of the Dorvilleidae (Polychaeta), with six new species from the deep H

North Pacific. Zoological Journal of the Linnaean Society 54: 101-135.

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I Kegel, B. and H.-D. Pfannenstiel. 1983a. Evaluation of the pair-culture effect in Ophryotrocha puerilis

(Polychaeta: Dorvilleidae). I. Pair-culture effect and sex ratio. Helgolander Meeresuntersuchungen

i 36: 205-213. Kegel, B. and H.-D. Pfannenstiel. 1983b. Evaluation of the pair-culture effect in Ophryotrocha puerilis

(Polychaeta: Dorvilleidae). II. Conditions for the moult of the upper jaw. Helgol_inder

I Meeresuntersuchungen 36:215-222.

Kinberg, J. 1865. Annulata nova. Oefversigt F6rhandlinger Svenska Vetenskaps Akademien 21: 559-574.

I Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bemstein, R. Cimberg, T. Kauwling, and W. Anikouchine. 1986. Assessment of long-term changes in biological communities of the Santa Maria Basin and

Western Santa Barbara Channel - Phase I. Report prepared for the U.S. Department of the Interior,

I Minerals Management Service, Pacific OCS Region, under contract No. 14-12-0001-30032.

Mclntosh, W. 1910. A monograph of the British annelids. II. Polychaeta. 2. SyUidae to Ariciidae. London,

i Moore, Ray Society, pp. 233-524. J.P. 1906. Additional new species of Polychaeta from the North Pacific. Proceedings of the Academy

of Natural Sciences Philadelphia 58: 217-260.

I Nordheim, H. von. 1987. Petrocha notogaea gen. et sp.n., a neotenic interstitial polychaete (Dorvilleidae)

from the Hauraki Gulf, New Zealand. Zoologica Scripta 16(1): 33-38.

i Orensanz, J.M. 1973. Los anelidos poliquetos de la provincia biogeographica Argentina. III. Dorvilleidae. Physis Seccion A 32(85): 325-342.

i Orensanz, J.M. 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas. With addenda to the Eunicemorpha of Argentina, Chile, New Zealand, Australia, and the southern Indian

Ocean. Biology of the Antarctic Seas XXI. Antarctic Res. Ser. 52:1-183.

I Parfitt, E. 1866. Description ofaNereisnew to science. Zoologist (Ser. 2) 1:113-114.

Pettibone, M.H. 1961. New species of polychaete worms from the Atlantic Ocean, with a revision of the

I Dorvilleidae. Proceedings of the Biological Society of Washington 74:167-186. Pettibone, M.H. 1963. Polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae.

Bulletin of the United States National Museum 227(1): 1-356.

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Richards, T.L. 1967. Reproduction and development of the polychaete Stauronereis r_Molphi, including a

I summary of development in the superfamily Eunicea. Marine Biology I: 124-133.

Verrill, A. 1900. Additions to the Turbellaria, Nemertina and Annelida of the Bermudas, with revisions of

some New England genera and species. Transactions of the Connecticut Academy of Arts and Sciences

I 10: 595-671.

Westheide, W. 1982. lkosipodus carolinensis gen. et sp.n., an interstitial neotenic polychaete from North

I Carolina, 117-126. U.S.A., and its phylogenetic relationships within Dorvilleidae. Zoologica Scripta 11(2):

i Westheide, W. and H. von Nordheim. 1985. Interstitial Dorvilleidae (Annelida, Polychaeta) from Europe, Australia and New Zealand. Zoologica Scripta 14(3): 183-199.

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Westheide, W., and N.W. Riser. 1983. Morphology and phylogenetic relationships of the neotenic interstitial I

polychaete Apodotrocha progenerans n.gen., n.sp. (Annelida). Zoomorphology 103: 67-87.

Wolf, ES. 1986a. Four new genera of Dorvilleidae (Annelida: Polychaeta) from the Gulf of Mexico. I

Proccedings of the Biological Society of Washington 99(4): 616-626.

Wolf, P.S. and1986b. a new genus Threefornewdorvilleids species offromDorville Scandinavia idae (Anneli anddaNorth : Polychae America. ta) from Proceedings Puerto Rico of theanBiological dFlorida I

Society of Washington 99(4): 627-638.

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364 I
I

I Appendix A

i List of Abbreviations on Figures

The explanation of letter symbols on figures follows. Roman numerals indicate body segments (in

I Phyllodocidae), areas of the proboscis (in Nereididae and Goniadidae), and maxillary jaw pieces (in Ennicea).

I aC, aK, anal achaetous cirrusknob ey, fAn,eyefrontal antenna

aLa, anterior lamella fPa, frontal palp (in onuphids)

I aP, anal or pygidial plate fTu,

facial tubercle

ac, acicula g, gap

I acB, accessory branchia gl, gland acL, or acLa, acicular lobe hG, hyaline gland

acS, acicular seta iLan, inner lateral antennae

I ai, aileron or wing on jaw InfLatAn, inferior lateral antenna

an, antenna intC, interramal cirrus or intercirrus

I ap, accessory papilla intSo, interramal sensory organ

br, branchia 1An,lateral antenna

I buS, bv, blood buccalvessel segment 1C, li, ligule lateral cirri

ca, capsule lig, ligament

I car, caruncle lo,

lobe

cer, ceratophore IPa, labial palps (in onuphids)

I cG, chromophile gland 1Sup,lateral support piece (lumbrinerid jaw) ch, chevron m, mouth

co, cortex mAn, median antenna

I cP, cephalic peak of prostomium mC, maxillary carriers

cph, cirrophore or base of cirrus mPa, median papilla

I cr, crotchet mR, maxillary ring

ct, ctenidium or ciliated cushion rot, macrotubercle

i cutP1, D, dorsal cutting plate macG,n, mandible macrognath

dC, dorsal cirrus or notocirrus max, maxilla

I dLa, dorsal lamella medulla

me,

dT, or dTu, dorsal tubercle microPap, micropapilla

I DtC, dorsal tentacular cirrus mid Pc, middle piece el, elytron or scale micG, micrognath

elph, elytrophore ne, neuropodium or ventral ramus of parapodium I

i 365
!

neAc, neuroacicula poL, podial lobe i

neLi, neuropodial ligule postL, postsetal (postacicular) lobe or lamella m

neL, neuropodial lobe or lamella postPap, postsetal papillae or podial fringe m

no, notopodium or dorsal ramus of parapodium pr, prostomium

noAc, notoacicula preL, presetal (preacicular) lobe or lamella i

noL, notopodial lobe or tamella prob, proboscis

i

noLi, notopodial ligule pmbO, proboscideal organs i

noRu, notopodial rudiment ro, rosette organ l

noS, notoseta sO, sense organ

huE, nuchal epaulet sS, swimming seta m

nuF, nuchal fold semR, seminal or sperm receptacle or pouch

m

null, nuchal hood set, setigerous segment or setiger i

nuO, nuchal organ setLo, setigerous lobe |

nuP, nuchal papilla st, style m

nuT, nuchal tubercle sty, stylodes of parapodia or parapodial fringe m

ocAn, occipital antenna subF, subpodial fringe or ventral pad

ocP, ocular peduncle SupLatAn, superior lateral antenna m

1Lan, outeral lateral antennae supNeL, superior neuropodial lobe

U

oR, oral ring subPap, subpodial lateral papilla or ventral cirrus

ov, ovary tC, tentacular cirrus m

pan, palpal antenna ts, tentacular segment

pLa, posterior lamelta tPa, tentacular palp i

pa, palp to, tooth

am

paPh, palpophore tOt, terminal organ •

paSty, palpostyle uNeL, upper neuropodial lobe

pap, papilla V, ventral m

parag, paragnath vC, ventral cirrus or neurocirrus i

PC, peristomial cirrus vPap, ventral or stomach papillae

per, peristomium VtC, ventral tentacular cirrus m

pi, pinnule

i

!

366 i
Appendix B

Lists and Maps of Stations

Table A.1. Position of soft-substrate stations taken during the Phase I Reconnaissance.

Station Latitude Longitude Depth

(m)

1 35°27.86"N 121°05.33'W 98

2 35°27.70'N 121°06.52"W 200

3 35°27.07'N 121°I0.20"W 291

4 35°26.56'N 121°14.93'W 393

5 35°25.77'N 121°21.69"W 585

6 35°20.88"N 120°59.62"W 109

7 35°20.65"N 121°02.57'W 197

8 35°20.00eN 121°06.58"W 308

9 35°19.48'N 121°10.06'W 398

10 35°18.28'N 121°18.65"W 591

11 35°17.80'N 121°22.13'W 690

12 35°15.03'N 120°57.3l'W 98

13 35o14.54'N 120°59.77"W 197

14 35°14.15"N 121°02.04'W 299

15 35°13.98'N 121°04.54'W 393

16 35°12.23"N 121°16.29'W 591

17 35°11.61"N 121°22.55'W 654

18 35°09.08'N 120°56.55'W 197

19 35°08.93'N 120°59.66'W 296

20 35° 15.72'N 121°04.68"W 396

21 35°06.11"N 120°44.82'W 49

22 35°05.85'N 120°50.23'W 99

23 35°05.60'N 120°55.18"W 195

25 35°05.07'N 121°00.75'W 390

26 35°04.38'N 121°15.99"W 590

27 35°04.30'N 121°19.27'W 611

28 35°04.22'N 121°19.65'W 603

30 34°54.19'N 120°47.07'W 98

31 34°53.76'N 120°52.96'W 200

32 34°53.56'N 120°56.81'W 297

33 34°53.43"N 120°59.66'W 396

34 34°53.15'N 121°04.40'W 492

35 34°52.96'N 121°10.30'W 548

36 34°52.77'N 121o15.37'W 492

38 34°49.8 I'N 120°52.66'W 197

39 34°49.53"N 120°56.85'W 294

40 34°49.24"N 121°00.8 I'W 392

41 34°48.35"N 121°19.14'W 495

42 34°48.04'N 120°47.50'W 100

43 34°46.59'N 120°52.92'W 197

45 34°44.91'N 120°59.59'W 395

46 34°41.22'N 121o13.56'W 597

47 34°41.99'N 121°10.81'W 378

48 34°45.11'N 120°52.85'W 196

49 34°45.03'N 120°56.3 I'W 290

50 34°37.80'N 121°01.66'W 591

52 34°39.56'N 120°47.64"W " 98

53 34°37.69'N 120°50.38'W 196

54 34°36.57'N 120°52.02'W 396

55 34°33.66'N 120°56.31'W 590

56 34°30.32'N 121°01.02'W 900

367
!

Table A.1 (Continued)

Station Latitude Longitude Depth i

l

(m) I

58 34°34.35"N 120°45. l 8'W 99 •

|

59 34°33.65"N 120°47.18"W 216

60 34°33.25"N 120°48.34'W 275

61 34°33.01'N 120°48.89'W 345

62 34°30.46'N 120°52.13"W 582 1

63 34°26.29'N 120°58.08'W 930

64 34°33.15'N 120°40.90"W 59

65 34°31.27'N 120°43.27'W 107

66 34°30.46'N 120°44.55'W 201 •

67 34°30.29'N 120°45.50"W 282

68 34°29.24'N 120°45.99'W 390

69 34°22.88'N 120°54.20'W 927 B

70 34°29.67'N 120°43.70'W 200 •

71 34 II °29.04'N t20°44.01'W 306

72 34°28.4 I'N 120°44.76'W 401

73 34°28.21"N 120°36.80"W 98 •

II

74 34°26.84"N 120%8.61'W 201

75 34°26.08'N 120°39.65'W 293

76 34°25.59'N 120°40.98"W 387

77 34°22.62'N 120°44.02'W 578 •

78 34°18.78'N 120°49.30"W 762

79 34°24.12'N 120°28.32"W 98

80 34°22.86'N 120°28.34'W 196 atom

81 34°21.26'N 120°28.83"W 294 •

82 34°18.7 I'N 120°29.55"W 394

83 34°17.20'N 120°30.20'W 444

84 34°13.54'N 120°31.19'W 394 1

|

85 34°25.88'N 120°16.3 I'W 113

86 34°24.45 'N 120°17.02'W 197

87 34°2 1.60'N 120°17.1 I'W 299

88 34°17.89'N 120°16.86"W 393 •

l

89 34013.79'N 120°16.56"W 471

90 34°09.44"N 120°16.30'W 375

91 34°11.73"N 120°07.43'W 540

92 34°08.70"N t 20°07.50'W 444 •

93 34°07.63'N 120°07.5 I'W 357

96 34°22.91"N 120°05.42'W 296

94 34°24.54"N 120°05.47'W 96

95 34°23.70'N 120°05.47'W 198 •

97 34°22.28'N 120°05.49'W 393

98 34°12.87'N 120°05.59'W 561

99 34°11.22"N 120°05.86'W 540 n

I

100 34°08.67'N 120°05.50'W 443

101 34°07.51'N 120°05.65"W 357

102 34°59.7 I'N 120°48.22'W 99

103 34°59.63'N 120°53.56'W 197 •

1

104 34°59.45'N 120°56.49'W 294

105 34°59.23'N 120°59.60'W 392

106 34°58.95'N 121°04.42'W 492

107 34°58.65'N 121°15.08'W 573 •

108 34°58.21'N 121°17.88"W 492

Note: Sample labels from the Soft-subslrate stations have several identification codes which include a station number, sample

replicate number, and analysis type. These are as follows: 001 to 200 = the range of station numbers; BSS = Benthic •

Sediment Single (i.e., a non-replicated station); BSR = Benthic Sediment Replicate (three replicates taken at this station); BSV =

Benthic Sediment Variance (subsamples); 01-09 = replicate numbers; TX = a taxonomy sample. Sample labels having the

designation BRA, represents a sample from rocks taken as part of the hard bottom survey. I

368 I
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10 •

11 •

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2'7 26 •

28 104 103 : 1(_

108 107 106• • • 6 o •

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I Figure A.1. Map showing location of soft substrate stations from Phase I and Phase II MMS Reconnaissance and Monitoring Programs.

I

i 369
I

Table A.2. Location of soft-subslrate stations taken during the Phase II Monitoring Program. I

Station Latitude Longitude Depth I

(m)

n

R-1 35°05.83'N 120°49.16'W 91 •

R-2 35°05.50'N 120°53.40'W 161

R-3 35°05.30'N 121°00.90'W 409

R-4 34°43.01'N 120°47.39'W 92 1

R-5 34°42.69"N 120°50.83"W 154 •

U R-6 34041.40"N 120°57.90"W 410

R-7 34°52.9tYN 121° 10.30"W 565

R-8 34°55.3tYN 120°45.87'W 90 •

R-9 34°53.68'N 120°59.12'W 4t0 1 PJ-1 34°55.79'N 120°49.91"W 145

PJ-2 34055.32 'N 120°49.59'W 142

PJ-3 34°56.26'N )20049.58'W 138 •

PJ-4 34°56.26'N 120°50.24'W ) 50

PJ-5 34°55.32'N 120°50.24'W 152

PJ-6 34054.7 I'N 120°49.91"W 148 i

PJ-7 34°55.79'N 120°48.60"W 123 •

PJ-8 34°56.87'N 120°49.91'W 142

PJ-9 34°55.79'N 120°51.23"W 169

PJ-10 34°53.63"N 120°49.91"W 147 n

PJ- 11 34°57.95"N 120049.9 I'W 136 | PJ-12 34°55.58'N 120°49.91"W 145

PJ-13 34°56.01'N 120°49.91"W 144

PJ- t4 34°55.79'N 120°49.26'W 134 •

PJ-15 34°55.79'N 120°50.57"W 155 I

PJ-16 34°55.03'N 120°48.99"W 130

PJ-17 34°56.56'N 120°48.98"W 126

PJ-18 34°56.56'N 120°50.84"W 158 •

PJ- 19 34°55.03'N 120°50.84"W 167

PJ-20 34°50.38'N 120°49.91'W 148

PJ-21 35001.23'N 120051.15'W 143 rail

PJ-22 34°55.25'N 120°49.93'W 143 •

PJ-23 34°56.33'N 120°49.90'W 143

I

Table A.3. Sampling dates of MMS Phase H Monitoring Program. I

Cruise Date I

1-1 October 1986

1-2 June 1987

1-3 May 1987 •

1-4 July 1987 | 2-1 October 1987

2-2 January 1988

2-3 May 1988 m

3-1 October 1988

3-2 May 1988 I

I

370 |
Table A.4. MMS Phase I - Locations of hard-substrate transects.

Station Beginning End Depth

Latitude Longitude Latitude Longitude (m)

1 A/B 34°24.454'N 120°01.876'W 34°24.464"N 120°00.878'W 69-73.5

1 C/D 34°24.076'N 120°00A43'W 34°24.184'N 120°01.480"W 73.5-78

2A/B 34°11.377"N 120°29.318'W 34°11.289"N 120°28.774"W 110-126

2 C/D 34° 10.984'N 120°28.094"W 34°10.780'N 120°27.554'W 120-123

4 A/B 34°27.539'N 120°40.364"W 34°28.162'N 120°40.t89'W 168-237

6 A/B 34°30.246'N 120°35.555'W 54-63

6 C/D 34Q30_42I'N 120°34.315'W 54-63

13 A/B 34°42".570'N 120°4"7.899'W 34°42.107"N 120°48.253"W 92-100

13 C/D 34°42.556'N 120°48.147'W 34°42.974'N 120°47.424'W 88.5-100.5

14 A/B 34°43.589"N 120°49.093"W 34°42.826'N 120°48.370'W 96-105

14 C/D 34°43.244"N 120°49.406"W 34°42.893'N 120°48.822'W 105-117

16 A/B 34°46.544"N 120°50.197'W 34°45.912'N 120°49.726'W 91.5-123

17 A/B 34°49.382'N 120°50.768"W 34°49.600'N 120°50.688'W 160.5-168

19 A/B 34°47.833'N 120°51.425'W 34°47.097"N 120°50.793'W 148.5-177

20 A/B 34°46.471YN 120°50.289'W 34°46.140'N 120°49.885'W 90-130.5

21 A/B 34°47.335'N 120°45.903'W 34°47.548"N 120°46.123"W 75-90

22 A/B 34°50.365'N 120°48.221'W 34°50.99(YN 120°48.365'W 114-115.5

23 A/B 34°49.868'N 120°47.393'W 34°50.003'N 120°47.480"W 93-102

25 A/B 35°05.662'N 120°47.562'W 35°06.036'N 120°47.652"W 64.5-72

26 C/D 35° 11.586'N 120°55.556'W 35°11.555"N 120°55.233'W 108-111

27 A/B 35°20.906'N 120°59.657"W 35°21.035'N 120°59.603'W 96-126

28 A/B 35°21.539'N 120°59.641"W 35°21.867'N 120°59.299'W 96-105

29 A/B 35°27.864'N 121°05.331'W 35°27.805"N 121°05.277'W 102-106.5

Table A.5. MMS Phase II - Locations of hard-substrate photosurvey stations.

Station Latitude Longitude Depth

(m)

PH-E 34°30.26'N 120°42.76"W 119

PH-F 34°30.81'N 120°42.36'W 105

PH-I 34°29.96"N 120°41.68"W 107

PH-J 34°29.82'N 120°41.82"W 117

PH-K 34°29.37"N 120°42.26"W 160

PH-N 34°29.2 I'N 120°42.05"W 166

PH-R 34°29.1 I"N 120°42.67"W 213

PH-U 34031.48"N 120':'43.5 I'W 113

PH-W 34031.52"N 120°45.86"W 195

371
I

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"_",... • 28 I

'_. • 27

I

", • 26

I

'', , • 25

I

!te • • 21 I

1§ go 20,

i • 14

_I"_°_ -, • '_ I

",,\ .,

'_ "7 6

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• 2 200 ............................ i

a4'_o'N I

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34 '_8'N

,_°.,,w ]7. In

Figure A.2 MaPand MonitoringSh°wing locatiOnSproghard rams.Of substrate stations from Phase I and Phase II MMS Reconnaissance I

I

372 I
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1

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I

The Department of the Interior Mission

I As the Nation's principal conservation agency, the Department of the Interior has responsibility

for most of our nationally owned public lands and natural resources. This includes fostering

sound use of our land and water resources; protecting our fish, wildlife, and biologicaldiversity;

i preserving the environmental and cultural values of our national parks and historical places;

and providing for the enjoyment of life through outdoor recreation. The Department assesses

our energy and mineral resources and works to ensure that their development is in the best

interests of all our people by encouraging stewardship and citizen participation in their care.

i The Department also has a major responsibility for American Indian reservation communities and for people who live in islandterritories under U.S. administration.

The Minerals Management Service Mission

I _'_AsabureauoftheDepartmentofthelnterior, theMineralsManagementService'slMMS)

primary responsibilities are to manage the mineral resources located on the Nation's Outer

Continental Shelf (OCS), collect revenue from the Federal OCS and onshore Federal and Indian

i lands, Moreover and distribute those revenues.

, in working to meet its responsibilities, the Offshore Minerals Management Program

administers the OCS competitive leasing program and oversees the safe and environmentally

sound exploration and production of our Nation's offshore natural gas, oil and other mineral

i resources. efficient, timely The MMS and accurate Royalty coManagement llection and Program disbursemment eetsofitsrevenue responsibilities from mineral by ensuring leasing the and

production due to Indian tribes and allottees, States and the U.S. Treasury.

I The responsive MMS strives to the topublic's fulfill itsconresponsibilities cerns and interests throughby thmaintai e general ning guiding a dialoguprinciples e with allof:potentially (1) being

affected parties and (2) carrying out its programs with an emphasis on working to enhance the

qua|ity of life for all Americans by lending MMS assistance and expertise to economic

I development and environmental protection.

I