I OCS Study

MMS 96-0042

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1 TAXONOMIC ATLAS OF THE BENTHIC FAUNA OF

THE SANTA MARIA BASIN AND

][ WESTERN SANTA BARBARA CHANNEL

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I INAL REPORT Volume 7 of 14

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The Annelida Part.4 m Polychaeta: Fiabelligeridae to Ampharetidae

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Edited by:

I James A. Blake

Brigitte Hilbig

I Paul H. Scott

Submitted by:

I Science Applications international Corporation

10260 Campus Point Drive

San Diego, California 92121

I For:

U.S. Department of the Interior

I Minerals Management Service Pacific OCS Region

770 Paseo Camadllo

i Camarillo, California 93010

Under Contract No. 14-35-0001-30484

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Minerals Management Service

I Pacific OCS Region July 1996

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DISCLAIMER I

This Service, reportU.S. hasDepartment been reviewedof by the Pacific Outer Continental Shelf Region, Minerals Mangement I

the Interior and approved for publication. The opinions, findings,

conclusions or recommendations expressed in this report are those of the authors, and do not

necessarily reflect the view of the Minerals Management Service. Mention of trade names does I

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

conformity with Minerals Management Service editorial standards.

TAXONOMIC DISCLAIMER I

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

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

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PROJECT ORGANIZATION

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I PROGRAM MANAGER

I DR. AND_W L. LLSSNER Science Applications International Corporation

10260 Campus Point Drive

I San Diego, California 92121

I DEPUTY PROGRAM MANAGER

I DR. JAMESA. BLAKE

ENSR Consulting and Engineering

I 89 Water Street

Woods Hole, Massachusetts 02543

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I PRO_--_r TECHNICAL OFmCER

MR. FRANKJ. MANAC, O

I U.S. Department of the Interior

Minerals Management Service, Pacific OCS Region

I 770 Paseo Camarillo

Camarillo, California 93010

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Co_r_ur_ AUrHORS

I DR. JAMESA. BLAKE

DR. BmGrrrE HmBIG

I DR. MAR',"E. P_RS_N

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ACCESS NUMBER: 30484.7 

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 

7. The Annelida Part 4 Polychaeta: Flabelligeridae to Ampharetidae. 

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; 1996. 

COMPLETION DATE OF REPORT July 1996. 

COSTS: FY 1989, $229,013; FY 1990, $1 50,000; FY 1 991 $1 50,000; FY 1993 $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, Polychaeta, Flabelligeridae, Acrocirridae, Fauveliopsidae, 

Capitellidae, Ampharetidae, California, Santa Maria Basin, Santa Barbara Channel, continental shetf. 

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 t 983. 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 7 are to continue coverage of the Polychaeta that was initiated in 

Volume 4. This volume continues the treatment of the so-called sedentary families. The following 5 families 

are treated in this volume: Flabelligeridae, Acrocirridae, Fauveliopsidae, Capitellidae, Ampharetidae. 

DESCRIPTION: Volume 7 continues treatment of the so-called sedentary families and includes the 

Flabelligeridae, Acrocirridae, Fauvetiopsidae, Capitellidae, Ampharetidae. The 5 chapters are organized 

into sections that include the morphology, taxonomic history, biology, keys of species, and descriptions of 

genera and species. Each species is fully illustrated with relevant characteristics labeled and identified. 
ACCESS NUMBER: 30484.7 

SIGNIFICANT CONCLUSIONS: Five families including 31 genera and 53 species of polychaetes are described 

and illustrated. The chapter on Fauveliopsidae includes detailed descriptions and new illustrations for 3 

species: Fauveliopsis glabra, F: armata, and F: magnus. Scanning electron micrographs are included for F: 

glabra; these are believed to the first SEMs published for a fauveliopsid. The chapter on Arnpharetidae 

includes 3 new species belonging to the genera Anobothnrs and Asabellides are described. A new genus, 

P seudampharete is established for Lysippe mexicana Fauc hald, 1 972. Sosanopsis wireni and Mexarnage 

corrugafa are newly recorded for California. 

STUDY RESULTS: The fauna of the Santa Maria Basin and western Santa Barbara Channel is evaluated 

from samples that were taken from depths of about 50 to 1,000 m. Organisms were collected from soft 

sediments using box cores and from rocks using manipulator arms of submersibles and remotely operated 

vehicles. The collections are organized into sets of vouchers from Phase I and I1 that have been made 

available to the team of investigators. Additional material from the bulk collections now archived with the 

Natural History Museum of Los Angeles County is also being examined. The total number of species treated 

in the entire Taxonomic Atlas (1 4 volumes) may exceed 1,300 species. 

Volume 7 includes 31 genera and 53 species of polychaetes in 5 families: Flabelligeridae (4 genera, 11 

species), Acrocirridae (1 genus and species), Faweliopsidae (1 genus, 3 species), Capitellidae (1 0 genera, 

16 species), Ampharetidae (1 5 genera, 25 species). Each chapter includes generic and species keys and 

usually includes some genera or species muring in adjacent waters. The chapter on Fauveliopsidae includes 

detailed descriptions and new illustrations for 3 species: Fauveliopsis glabra, E armafa, and E magnus. 

Scanning electron micrographs are included for F: glabra; these are believed to the first SEMs published for 

a fauveliopsid. The chapter on Capitellidae includes updated generic diagnoses for Decamastus, 

Heteromastus, Leiochrides, Mediomastus, and Notomastus. A new genus and species, Dodecamastus 

mariaensis, is described. Taxonomic problems related to the genera Neoheteromastus and Neomediomastus 

are discussed. The chapter on Ampharetidae is a major review of offshore genera and species. Fifteen 

genera are defined and taxonomic problems noted. Three new species belonging to the genera Anobothnrs 

and Asabel/ides are described. A new genus, Pseudampharete is established for Lysippe mexicana Fauchald, 

1972. The mrd of Sosanopsis wireni is the first for California; the range of Mexamage corrugata is 

extended to California from Mexico. 

STUDY PRODUCT: Blake, J.A. Hilbig, B., and P.H. Scott (Editors). 1996. Taxonomic Atlas of the Benthic 

Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Volume 7. The Annelida Part 4: 

Polychaeb: Flabelligeridae to Ampharetidae. A final report prepared by Science Applications International 

Corporation, for the U.S. Department of the Interior, Minerals Management Senrice, Pacific OCS Region, 

Camarilla, CA. OCS Study MMS 96-0042. Contract No. 14-35-0001 -30484. 

'P.l's affiliation may be different than that listed for Project Manager(s). 
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REPORT DOCUMENTATION !1. REPocr.O. =; =..==i_,_s k==.,_,..o.

PAGE MMS 96-0042

! 4. Title alt¢l Subtitle 5, Report Date

Taxonomic Atlas of theSanta Maria Basin and Western Santa Barbara Channel. Volume Juiy 1996

7. The Annelida Part 4 -- Flabetligeridae to Ampharetidae. 6.

7. Author(s) 8. Pefformir_ OrlPnization Rapt, No.

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

9. IselfornlJnll Oqplnizstion Name and A_lross )0. Project/Task/Work Unit No.

Science Applications International Corporation ,. co,,t,=_c)o, Qr, t(a) No.

10260 Campus Point Drive

San Diego, California 92121 fc)

14-35-001-30484

(G)

12. Sponsodnl OrlaniZatton _ma and Address 13. Type of Report & Pe_od Covered

Final 7 of 14 volumes, Mineral Management Service, U. S. Department of the Interior,

Pacific OCS Region, 770 Paseo Camarilla 9t89 to 7/96

Camarilla, California 93010 14

1S. Supplementary Notes

Contributing authors: Dr. James A. Blake, Dr. Brigitte Hitbig and Dr. Mary E. PatersOn.

14. Abstract {Limit: 200 words)

Volume 7 includes 31 genera and 53 species of polychaetes in 5 families: Flabelligeridae (4 genera, 11 species),

Acrocirridae (1 genus and species), Fauveliopsidae (1 genus, 3 species), Capitellidae (10 genera, 16 species),

Ampharetidae (15 genera, 25 species). Each chapter includes generic and species keys and usually includes some

genera or species occuring in adjacent waters. The chapter on Fauveliopsidae includes detailed descriptions and new

illustrations for 3 species: Fauvefiopsis gfabra, F. armata, and F. magnus. Scanning electron micrographs are included

for F. gfabra; these are believed to the first SEMs published for a fauveliopsid. The chapter on Capitellidae includes

updated generic diagnoses for Decamastus, Heteromastus, Leiochrides, Mediomastus, and Notomastus. A new genus

and species, Dodecamastus manaensis, is described. Taxonomic problems related to the genera Neoheteromastus

and Neomediomastus are discussed. The chapter on Ampharetidae is a major review of offshore genera and species.

Fifteen genera are defined and taxonomic problems noted. Three new species belonging to the genera Anobothrus and

Asabeflides are described. A new genus, Pseudampharete is established for Lysippe mexicana Fauchald, 1972. The

record of Sosanopsis wireni is the first for California; the range of Mexamage corrugata is extended to California from

Mexico.

17. Document AmCysts a. O_crtptors

Marine invertebrates, Annetida, Polychaeta, taxonomy, California, Santa Maria Basin, Santa Barbara Channel,

continental shelf.

k I_/Open-Ended Terms

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

c. COSATI field/Group

|9. Aveiblbillty Sttttment 19. Security Class (This Report) 2I. No. of Pai[e$

Unlimited Unclassified 164

20. Security Class (This Pqre) 22. Price

See ANS_Z39.18) See Instructkms on Reverse OPTIONAL FORM 272 (4-77)

(Formerly NTIS-35)

I Oepa_ma4_t of Commute
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Table of Contents I

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1. F_nLY FLABELUCEmV_ SAN JOSEPH, 1894 (by James A. Blake)

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

Morphology ..................................................................................................................................... 1 1

Biology ............................................................................................................................................ 2

Taxonomic History .......................................................................................................................... 3 m

Key to Genera and Species from the Santa Maria Basin ................................................................ 3 II

Descriptions of Species ................................................................................................................... 4

Brada viltosa (Rathke, 1843) ................................................................... 5 •

Flabeltigera affinis Sars, 1829................................................................. 8 I

Flabelligera commensalis Moore, 1909................................................ 10

Piromis hospitis Fauchald, 1972 ............................................................ 12 •

Pherusa capulata (Moore, 1909) ........................................................... 13 I

Pherusa inflata (Treadwell, 1914) ......................................................... 15

Pherusa neopapilIata Hartman, 1961 .................................................... 17 •

Pherusa papillata (Johnson, 1901) ........................................................ 19 l

Literature Cited ............................................................................................................................. 20

2. FAMILYACROCmRmAEBANSE,1969 (by James A. Blake) 1

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Introduction ................................................................................................................................... 25

Morphology ................................................................................................................................... 25 I

Biology .......................................................................................................................................... 26 l

Taxonomic History ........................................................................................................................ 26

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Key to Genera and Species of Acrocirridae from California ........................................................ 26 m

Description of Species ................................................................................................................... 27 l

Acrocirrus heterochaetus Annenkova, 1934 ......................................... 27

Literature Cited ............................................................................................................................. 29

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3. FAMmV FAU_LIOPSmAE HARTMAN,1971 (by James A. Blake and Mary E. Petersen) II

Introduction ................................................................................................................................... 31 1

Morphology ................................................................................................................................... 31 1

Taxonomic History ........................................................................................................................ 33

Key to Species of Fauveliopsis from California ........................................................................... 35 I

Description of Species ................................................................................................................... 35 l

Fauveliopsis glabra (Hartman, 1960) .................................................... 36 1

Fauveliopsis armata Fauchald and Hancock, 1981 ............................... 40

Fauveliopsis magna Fauchald and Hancock, 1981 ................................ 42 l[

Acknowledgements ....................................................................................................................... 44

Literature Cited ............................................................................................................................. 44

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4. FAMILY CAPITELLIDAEGRUIIE,1862 (by James A. Blake) I

Introduction ................................................................................................................................... 47

Morphology ................................................................................................................................... 47 I

Taxonomic History ........................................................................................................................ 48

Biology .......................................................................................................................................... 49

Key to Genera of Capitellidae from California ............................................................................. 54 •

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I Descriptions of Species ................................................................................................................. 55

Barantolla americana Hartman, 1963 ................................................... 56

i Capitella Capitella capitata ovincola (Fabricius, Hartman, 1947 1780)......................................................... sensu lato ..................................... 5861

Decamastus gracilis Hartman, 1963 ...................................................... 63

I Key to SDodecamastus pecies of Heteromastus mariaensisfromBlake, California new species ................................................................................... 6567

Heteromastusfilobranchus Berkeley and Berkeley, 1932 .................... 67

i Heteromastus filiformis (Clapar_de, 1864) ............................................ 69 Leiochrides hemipodus Hartman, 1960 ................................................. 71

Key to Species of Mediomastus .......................................................................... 73

i Mediomastus acutus Hartman, 1969 ...................................................... 74 Mediomastus ambiseta (Hartman, 1947) ............................................... 76

Mediomastus californiensis Hartman, 1944 .......................................... 78

i Key to Species of Notomastus ............................................................................ 80 Notomasms hemipodus Hartman, 1945 ................................................. 81

Notomastus magnus Hartmart, 1947 ...................................................... 83

Notomastus tenuis Moore, 1909 ............................................................. 85

I Neoheteromasms 1960 87 Hartman,

Neoheteromastus lineus Hartman, 1960 ................................................ 87

Neomediomastus Hartman, 1969 ........................................................... 88

i glabrus (Hartman, 960) ...........................................

Neomediomastus 1 89

Acknowledgments ......................................................................................................................... 90

Literature Cited ............................................................................................................................. 90

I 5. FAMn,YAMPHARE'rn)AEMALMC_RW1867 , (by Brigitte Hilbig)

Introduction ................................................................................................................................... 97

I Morphology 97 Taxonomic History ...................................................................................................................... 100

Distribution and Biological Notes ............................................................................................... 100

I Key to the Ampharetidae ............................................................................................................. 102 Description of Species ................................................................................................................. 105

Subfamily Ampharetinae ............................................................................................... 106

I Amage anops (Johnson, 1901) ............................................................. 106 Ampharete acutifrons (Grube, 1860) ................................................... 108

Ampharetefinmarchica (Sars, 1864) ................................................... 110

I Ampharete Amphicteis mucronata labrops Hartman, Moore,1961 ...................................................... 112 1923 .................................................... 114

Amphicteis scaphobranchiata Moore, 1906 ........................................ 116

i AAnobothrus mphisamythagracilis bioculata (Malmgren, (Moore,1866) 1906)............................................................................................ 112018

Anobothrus paleatus Hilbig, new species .......................................... 122

i Asabe AsabeUUides ides cacornuta lifornicaHilbig, Hilbig, newnewspecispecies es ............................................ ....................................... 124 126

Asabellides lineata (Berkeley and Berkeley, 1943) ............................. 127

I Eclys Glyphaippenostomum trilobatapal(Har lescens tman(,T19h6e69)l, ................................................... 1879) ......................................... 1302

Lysippe Iabiata Malmgren, 1866 ......................................................... 134

i Mexamage corrugata Fauchald, 1972 ................................................. 135

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Mugga wahrbergi Eliason, 1955 .......................................................... 138 I

Paralysippe annectens (Moore, 1923) ................................................. 140

Paramage Pseudampharete scutatamexicana (Moore, (1923) Fauehal........................................................ d, 1972), new combination ....... 14244 I

Samytha californiensis Hartman, 1969 ................................................ 146

Sosane Sosanopsis occidentalis wireni Hessle, (Hartman, 19171........................................................... 969) .................................................. 115048 I

Subfamily Melinninae .................................................................................................... 151

Melinna Melinna heterodonta Moore, 1923 ...................................................... 151 i

oculata Hartman, 1969 .......................................................... 153

Literature Cited ........................................................................................................................... 155

APPENDIX..................................................................................................................................... 159 I

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i List of Figures

I Figure 1.1. Brada villosa: A, entire anhnal, lateral view; B, anterior end, lateral view; C, papilla; D,

middle parapodium; E, neuroseta; F, notoseta. (A, C, F, after St0p-Bowitz, 1948; B, D, E,

I after Uschakov, 1955) ........................................................................................................ 6

Figure 1.2. Flabelligera affinis: A, entire animal, ventral view; B, neuroseta with papillae; C, notoseta.

(A, B after St_p-Bowitz, 1948; C, after Uschakov, 1955) ................................................. 9

I Figure 1.3. Flabelligera commensalis: A, entire animal, dorsal view; B, notoseta; C, neuroseta. (A

after Hartman, 1961; B-C after Moore, 1909) ................................................................. 11

i Figure 1.4. Piromis hospitis: A, anterior end, lateral view; B, parapodium, anterior view; C, anterior neuroseta; D-E, posterior neurosetae in different degrees of wear .................................. 13

I Figure 1.5. Pherusa C, capulata: A, anterior end, dorsal view; B, papilla and basally adhering sand grains; tip of neuroseta ............................................................................................................ 14

Figure 1.6. Pherusa inflata: A, entire animal, lateral view; B, anterior end, frontal view, with right palp

I and most branchial filaments removed showing location and arrangement of branchiae, prostomium, eye spots, and mouth; C, notoseta from middle body segment; D-H, neurosetae

from setigers 4, 5, 6, 7, and middle setiger. (after Hartman, 1961) ................................. 16

I Figure 1.7. Pherusa neopapillata: A, entire animal, lateral view; B, anterior end, dorsal view; C, anterior

end, frontal view, showing palps, branchial filaments, scars of same, prostomium, and oral

i Figure 1.8. Pherusa slit; D, neur papillata oseta.:(A, A,C,anterior D, afterend, Hartman, dorsal 1961; view; BB, , original) anterior ........................................ end, frontal view showing18

palps, branchial filaments, scars of same, prostomium with eyes, and oral slit; C, neurosetae

I from setiger 14. (A, original; B, C after Hartman, I961) ................................................ 19

Figure 2.1. Acrocirridae: A, anterior end of a representative acrocirrid, Acrocirrus frontifilis; B-G,

i Acrocirrus heterochaetus: B, anterior end, lateral view; C, sixth segment, anterior view; D, segments 13-15, lateral view; E, compound falciger; F, notoseta with inset showing detail

of fine serrations; F, acicular hook from segment 14. (A-D, after Banse, 1969; E-G, original)

............................................................... , .......... °................... ° ..................... ° ..... °....... ° ..... °. 28

I Figure 3.1. FauveIiopsis glabra (Santa Maria Basin, Sta. PJ-9), entire animal in ventral view ........ 32

• Figure 3.2. FauveIiopsis glabra (Santa Maria Basin, Sta. PJ-9): A, anterior end in lateral view showing

I modified setae of setigers 1-3 and following setigers; B, lateral view of setigers 8-12, showing

structure and location of genital papilla on setiger 10 ..................................................... 38

I Figure 3.3. Fauveliopsis setiger 2, anterior glabraview; (Santa C, setiger Maria4,Basin, posterior Sta.view; PJ-19):D, A, setiger entire30,animal posteriorin dorsal view; E,view; papillaB,

from anterior part of body; F, papillae from around pygidium ........................................ 39

i Figure 3.4. Fauveliopsis armata (holotype, LACM-AHF Poly 1160): A, entire animal in dorsal view;

B, setiger 4, posterior view; C, setiger 13, posterior view; D, setiger 20, posterior view; E,

setiger 26, posterior view ................................................................................................. 41

I Figure 3.5. Fauveliopsis magna: A, entire animal in left ventrolateral view (holotype, LACM-AHF

Poly 1149); B, setiger 2, posterior view (Sta. DS-2); C, setiger 8, posterior view (Sta. DSi 2); D, setiger 22 (holotype), posterior view; E, setiger 35 (holotype), posterior view .... 43
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Figure 4.1. BarantoUa americana: Anterior end in lateral view (after Hartman, 1969) .................... 57 i

Figure 4.2. Capitella capitata: A, anterior end, dorsal view; B, setigers 7-10 of male showing genital

spines; C, similar diagram of female showing genital pore; D, E, hooded hooks. (A-C, after •

Berkeley and Berkeley, 1952; D-E, after Hartman, 1947) ............................................... 59

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Figure 4.3. Capitella capitata (Tomales Bay): A, anterior end dorsal view; B, anterior end of another •

specimen, lateral view ...................................................................................................... 60 II

Figure 4.4. Capitella ovincoIa: A, anterior end, lateral view (Sta. 79); B, posterior thoracic and anterior

abdominal region, dorsal view; C, abdominal segments; D, genital spine; E, F, hooded •

hooks. (B-F, after Hartman, 1947) ................................................................................... 62 II

Figure 4.5. Decamastus gracilis (Sta. R-5): A, anterior end, lateral view; B, anterior abdominal segments, U

dorsal view; C, middle abdominal segment ..................................................................... 64 U

Figure 4.6. Dodecamastus mariaensis Blake, new species (Hototype, Sta. 62): A, anterior end, lateral

view; B, far posterior abdominal setiger with branchiae; C, hooded hook ...................... 66 ii

Figure 4.7. Heteromastusfilobranchus: A, anterior end, lateral view (Sta. R-3); B, far posterior branchial

bearing abdominal segment, dorsal view; C, detail of branchiae from same; D, pygidium, m

dorsal view; E, F, thoracic hooded hooks; G, H, abdominal hooded hooks. (B-D, after Ii

Berkeley and Berkeley, 1952; E-H, after Hartman, 1947) ............................................... 68

Figure 4.8. Heteromastus filiformis: A, anterior end, lateral view through first 5 setigers (Humboldt t

Bay); B, same specimen, setigers 6-11; C, far posterior setiger from same specimen, showing !

branchiae; D, E, thoracic hooks; F, G, abdominal hooks (D-G, after Hartman, 1947) ... 70

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Figure 4.9. Leiochrides hemipodus (Sta. 28): A, anterior end, lateral view, prostomium and first 8 I

segments (fh'st 7 setigers); B, same specimen, continuation of A, showing last 5 thoracic

setigers and first 2 abdominal setigers; C, far posterior branchial bearing parapodium; D, E, I

abdominal hooks .............................................................................................................. 72 i

Figure 4.10. Mediomastus acutus: A, anterior end, lateral view; B, winged thoracic capillary seta; C,

modified thoracic notoseta; D, bidentate hooded hook. (A, D, after Hartman, 1969; B, C, i

after Warren et al., 1994) ................................................................................................. 75 I

Figure 4.11. Mediomastus ambiseta: A, anterior end, lateral view; B, posterior end, lateral view; C, D. i

abdominal hooded hooks. (after Hartman, 1947) ............................................................ 77 R

Figure 4.12. Mediomastus californiensis: A, anterior end, lateral view; B, detail of anterior end, lateral

view; C, D, abdominal hooded hooks. (A, B, after Hartrnan, 1944; C, D, after Hartman, i

1947) ................................................................................................................................. 79 U

Figure 4.13. Notomastus hemipodus: A, anterior end, lateral view; B, prostomium and peristomium, i

dorsal view; C, middle abdominal segments, dorsal view; D, E, abdominal hooded hooks. I

(after Hartman, 1947) ....................................................................................................... 82

Figure 4.14. Notomastus magnus: A, anterior end, lateral view; B, prostomium and peristomium, dorsal IIII

view; C, anterior abdominal segments showing nephridial openings; D, far posterior

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segments, dorsolateral view, showing branchiae; E, F. abdominal hooded hooks. (after

Hartman, 1947) ................................................................................................................. 84 i

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I Figure 4.15. Notomastus tenuis: A, anterior end, lateral view; B, prostomium and anterior part of

peristomium, dorsal view; C, posterior setigers, lateral view; D, E, abdominal hooded hooks.

I (after Hartman, 1947) ....................................................................................................... 86

Figure 4.16. Neoheteromastus lineus: Anterior end, lateral view. (after Hartman, 1969) ................. 88

i Figure 4.17. Neomediomastus gIabrus: Anterior end, lateral view. (after Hartman, 1969) ............... 89

Figure 5.1. Schematic lateral views of anterior ends of genera and species treated in this chapter,

indicating distribution of branchiae, noto- and neuropodia, and specialized setae among

I segments 1 through 7 99

Figure 5.2. Amage anops: A, entire animal, dorsolateral view, first notopodium concealed by branchiae;

B, firstanteri abdominal or end, dsegments, orsal view,lateral schemaview; tic; C, E, ndetail otopodi ofunotoseta; m, anterioF-G, r view; thoracic D, lastuncini, thoracilateral c and

and frontal views. (lateral view from Johnson) .............................................................. 107

I Figure 5.3. Ampharete acutifrons: A, entire animal, lateral view; B, anterior end, dorsal view,

schelnatic;

C, palea; D, thoracic uncinus, lateral view; E, abdominal uncinus, frontal view; F, last

thoracic and first abdominal segments, lateral view; G, posterior end, lateral view. (D, E, G

I after Zottoli) ................................................................................................................... 109

Figure 5.4. Ampharetefinmarchica: A, entire animal, dorsolateral view; B, anterior end, dorsal view,

i schematic; C, palea; D-F, thoracic, anterior abdominal, and posterior abdominal uncini, lateral and frontal views; G, last thoracic and first abdominal segments, lateral view; H,

posterior end, lateral view; I, detail of pygidium. (D-F, H, I after Zottoli) ................... 111

I Figure 5.5. Ampharete labrops: A, entire animal, lateral view; B-C, anterior end, schematic, dorsal and ventral views; D, tip of palea; E, notoseta; F-G, dorsalmost and ventralmost thoracic uncini

from same fascicle; H, abdominal uncini, 3 different views ......................................... 113

I Figure 5.6. Amphicteis mucronata: A, entire animal, dorsal view; B, anterior end, schematic, dorsal

view; C, palea; D, detail of notoseta; E-F, thoracic and abdominal uncini, lateral and frontal

• views; G, notopodium, anterior view; H, abdominal neuropodium, anterior view; I, posterior

I _ end, dorsolateral view .................................................................................................... 115

I Figure 5.7. Amphicteis scaphobranchiata: A, entire animal, dorsolateral view; B, last thoracic and first

Ii abdominal segments, lateral view; C, posterior notopodium, anterior view; D, palea; E,

thoracic uncini, lateral and frontal view; F, abdominal uncinus; G, posterior end, lateral

view. (A, E, F modified after Uebelacker) ..................................................................... 117

I Figure 5.8. Amphisamytha bioculata: A, anterior end, dorsal view; B, same, ventral view; C, abdominal

segments, lateral view; D-E, thoracic uncini; F, abdominal uncinus. (A-C from Williams;

D-F from Hartman) ........................................................................................................ 119

i Figure 5.9. Anobothrus gracilis: A, entire animal, lateral view; arrows: dorsal ridges; B, anterior end,

dorsal view, schematic; C, tip of regular notoseta; D, tip of modified notoseta, unciniger 8;

i E-F, ........................................................................................................................................ thoracic and abdominal uncini, lateral and frontal view; G, posterior end, lateral view121

Figure 5.10. Anobothrus paleatus Hilbig, new species: A, entire animal, lateral view; arrows: dorsal

I ridges; end, view; C, same, view; D, palea; E, modified seta; F,

B, anterior frontal lateral

detail of tip; G, normal seta (D, E, G drawn to scale); H, thoracic uncinus, anterolateral

i view ................................................................................................................................ 123

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Figure 5.11. Asabellides californica: A, entire animal, lateral view; B, detail of buccal tentacle; C-D, t

first and third abdominal neuropodium; E, thoracic uncinus, frontal view; F-G, anterior and

posterior abdominal uncini, lateral views ...................................................................... 125

Figure 5.12. Asabellides cornuta: A, anterior end, dorsal view, bottom of buccal cavity extended; B, tip

of buccal tentacle, 4 rows of pinnae shown; C, anterior end, ventral view; D, notopodium;

E, abdominal neuropodium; F-G, thoracic and abdominal uncini, frontal and lateral views i

........................................................................................................................................ 27

Figure 5.13. Asabellides lineata: A, entire animal, dorsolateral view; B, tip of buccal tentacle; C, anterior n

end, lateral view; D, last thoracic and t-n-stabdominal segments, lateral view; E, thoracic _

uncinus, frontal view; F-G, abdominal uncini, frontal and lateral views; H, posterior end,

dorsal view. (Puget Sound specimens) ........................................................................... 128

I! Figure 5.14. Eclysippe trilobata: A, entire animal, dorsal view; B, anterior end, schematic, dorsal view;

C, anterior end, ventral view; D, palea; E, anterior end, lateral view; F, posterior notopodium;

G-H, thoracic and abdominal uncini, 3 different views ................................................. 131

i

Figure 5.15. Glyphanostomum pallescens: A, entire animal, lateral view (after Zottoli); B, anterior end,

lateral view, buccal tentacles extended; C-D, thoracic and abdominal uncini, lateral and •

frontal views ................................................................................................................... 133 |

Figure 5.16. Lysippe labiata: A, entire animal, lateral view; B, anterior end, dorsal view; C-D, thoracic

and abdominal uncini, frontal view. (C, D after Zottoli) ............................................... 135 i

u

Figure 5.17. Mexamage corrugata: A, entire animal, dorsal view; B, anterior end, lateral view; C, posterior

end, ventrolateral view; D-F, uncini, 3 different views ................................................. 137 •

!1 Figure 5.18. Mugga wahrbergi: A, entire animal, lateral view; B, regular notoseta; C, modified notoseta;

D-F, uncini, different views ........................................................................................... 139

i

Figure 5.19. Paralysippe annectens: A, anterior end, ventrolateral view (from Williams); B-C, thoracic i

uncini, frontal and lateral views; D, abdominal uncinus; E, posterior end, ventral view

(from Hartman) .............................................................................................................. 141 i

I Figure 5.20. Paramage scutata: A, entire animal, dorsal view; B, anterior segmentys, lateral view; C,

last thoracic and first abdominal segments, lateral view; D, superior thoracic uncinus, frontal

view; E-F, abdominal uncini, 2 different views ............................................................. 143 i

n

Figure 5.21. Pseudampharete mexicana: A, anterior end, ventral view; B, same, dorsal view; C-D, tips

of smooth and annulated branchiae; E-F, uncini, anteror and lateral views; G, posterior end, i

lateral view ..................................................................................................................... 145 n

Figure 5.22. Samytha caIiforniensis: A, entire animal, lateral view; B, thoracic unciniger, lateral view;

C-D, abdominal uncinigers, frontal and lateral views; E, posterior end, ventrolateral view. 'i

03, E after Hartman) ....................................................................................................... 147

Figure 5.23. Sosane occidentalis: A, entire animal, ventrolateral view; B, regular notoseta, tip; C, thoracic m

uncinus, frontal view; D-E, abdominal uncini, 2 different views; F, modified seta, tip; G, i

setiger 13, dorsum, anterior view ................................................................................... 149

Figure 5.24. Sosanopsis wireni: A, anterior end, lateral view; arrow: insertion of posteriormost branchia; 10

B, regular seta; C, modified seta; D-E, thoracic and abdominal uncini, frontal and lateral

views. (after an unpublished sketch by R.E. Ruff) ........................................................ 150

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I Figure 5.25. Melinna heterodonta: A, entire animal, lateral view; B, anterior end, lateral view, buccal

tentacles partly extended; C, same, dorsal view; D-F, thoracic uncini, 3 different views; GI H, abdominal uncini, lateral and frontal views .............................................................. 152 Figure 5.26. Melinna oculata: A, entire animal, lateral view; B, anterior end, dorsal view, branchiae

pulled back to show prostomium; C, same, branchiae in natural position; D-E, thoracic

i uncini, lateral and frontal views; F, uncinus from middle of abdomen, frontal view; G-H, uncini from far posterior segment, lateral and frontal views; I, posterior end, ventrolateral

view ................................................................................................................................ 154

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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.

Marie Murphy of the Santa Barbara Museum of Natural History provided graphic design and production ill

assistance for this volume. J

.I

List of Acronyms i[

ANSP Academy of Natural Sciences, Philadelphia.

I

BLM Bureau of Land Management. _

BRA Refers to a station designation from the MMS Phase I Reconnaissance: Benthic Rocky,

transect A/B. I

BRC Refers to a station designation from the MMS Phase I Reconnaissance: Benthic Rocky,

transect C/D. I

CAS California Academy of Sciences, Department of Invertebrate Zoology, San Francisco,

California, USA. I

LACM Natural History Museum of Los Angeles County, Los Angeles, California, USA. 1

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA. 111

MMS United States Minerals Management Service. I!

SCAMIT Southern California Association of Invertebrate Taxonomists. I

SBMNH Santa Barbara Museum of Natural History, Santa Barbara, California, USA. U

SDNHM San Diego Natural History Museum, San Diego, California, USA. 1

USNM United States National Museum. A historical designation for the National Museum of !1

Natural History (NMNH), Smithsonian Institution, Washington, D.C., USA.

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I 1. FAMILYFLABELLIGERIDAE SAINTJOSEPH, 1894

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i James A. Blake t

Introduction

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The flabelligerids are highly modified, sedentary, surface deposit feeders that are commonly called

I "iscage elongate, worms"thickened, because oand f theusually chractesomewhat ristic long,fusiform forwardlyindire shape. ctd The setaecuticle found inis many often gener ornamented a. The bod withy

numerous short to long mucus-secreting epithelial papillae. Silt and sand particles often adhere to the mucoid

i seOther cretions, providing some species with a covering of particles through which setae and papillae protrude. species secrete a thick, transparent mucous sheath.

The prostomium and peristomium retract into the first few setigers, sometimes making it difficult

I tThe o study perisctri omium tical cephalic bears eight strucortures. moreTbranchiae wo long, groo arising ved fromprostomi a membrane ai palps are or hood. presentParapodia and usedarein reduced. feeding.

Notosetae are capillaries, with those of setigers 1-6 often elongated and directed anteriorly forming the

I cephalic compoundcage. or pseudocompound Neurosetae includehooks, distinct or various crossbarred combinations capillaries, of these similarsetae. to nIfotatosetae, all possible, simple flabelligerids or bidentate,

should be examined alive for an unobstructed view of prostomial and peristomial features.

l Morphology

i The body may be subcylindrical and elongate, expanded anteriorly and tapering posteriorly, short

and fusiform, grub-shaped, or thick, flattened, and disk-shaped. Segments are generally similar throughout

I the body and not particularly well defined. The body has a glandular epithelium that is usually covered with papillae of various shapes and sizes. In the genus Flabelligera, the entire body is covered with a thick,

transparent mucous layer through which papillae and setae project. In some genera, a thin mucous matrix is

I impregnated with sand, silt, and debris, imparting a rough granular appearance to the body.

The head region of flabelligerids contains several complex structures that are more or less retractable

into the anterior setigers, becoming mostly hidden from superficial inspection. It is formed of an indistinct

1 prostomium that is fused with the peristomium and consists of a prostomial ridge that usually bears four l

eyes, a pair large grooved palps, two groups of ciliated, retractile branchiae arising from a curved dorsal

membrane, and a ventral mouth enclosed in upper, lateral, and lower lips. The pharynx is noneversible. The

i cephalic cage surrounds the head region and when present, is formed of capillary setae and sometimes elongated club-shaped papillae from setigers 1-6.

i

1ENSR,89WaterStreet,WoodsHoLe,Massachusett0254 s 3.

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Parapodia are biramous with podial lobes and tori usuaUy reduced, so that the only evidence of II

noto- and neuropodia are the setal bundles themselves. Podial lobes are sometimes prominent (genus

Flabelligera); dorsal and ventral cirri are absent. Notosetae are all crossbarred or smooth capillaries, t.

Neurosetae may be similar to notosetae or may include simple or bidentate hooks, compound or I

pseudocompound hooks, or various combinations of these. The pygidium is simple and lacks anal cirri.

Details concerning the anatomy and fine structure of flabelligerids may be found in Schlieper (1927:

Pherusaplumosa), Borodin (1929), and Spies (1973, 1975, 1977: FlabetIigera commensalis as Flabelliderma). II

Biology 1

I

Flabelligerids are generally thought of as burrowers that do not form a permanent tube (Fauchald l[

and Jumars, 1979), however, Day (1967) indicated that some species might form tubes. Flabelligerids have

been reported to crawl over the surface, live in U-shaped tubes, or inhabit burrows in rock. They are found

under stones, and in their burrows just below the surface of sand or mud. Some species such as Flabelligera I!

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commensalis, are commensals that live on the surface of sea urchins. The cephalic cage and the sandy crusts

that develop over the bodies of some species may serve as protection from predators.

According to Fauchald and Jumars (1979), most flabelligerids are motile surface deposit feeders, N

using their palps to gather food particles. These authors indicate that the beating cilia on the branchiae set up

currents that may assist feeding by presumably suspending particles that can be picked up by the palps and I

carried toward the mouth. Gut contents include diatoms, other unicellular algae, larger algal fragments, II

sponge fragments, foraminiferans, and detritus. Fauchald and Jumars (1979) suggest that some particle

sorting takes place. The commensal species, F. commensalis, uses its palps to feed on the fecal matter of the dl

host sea urchin. |

The only detailed account of a burrowing flabelligerid is that of Amor (1994) who studied the

ecology of Pherusa sp., that forms burrows in hard intertidal substrates in Argentina. The worms live in a m

burrow in the rock that is lined with aragonite. The hardened dorsal shield over the first three setigers serves II

as a plug for the burrow when the animal withdraws. The paired palps and branchiae play a role in feeding.

Food particles collected by these organs are moved by ciliary current to the mouth. Food resources include _t

suspended microalgae and detritus. I

Very little is known concerning reproduction and larval development of flabelligerids. The only

study dedicated to this subject is by Spies (1977), who studied F. commensalis at Catalina Island. Sexes of I

F. commensalis and other flabelligerids are separate. Externally, there is no evidence of sexual dimorphism !1

in 17.commensalis, except that sexually mature male gonads are white and those of females are green. Spies

described the morphology and ultrastructure of the ovaries and testes of F. commensalis. Sperm have a I

spherical head and midpiece consisting of "five spherical bodies..." presumably mitochondria. The form of I

these sperm clearly suggests that they are of the primitive type and should be spawned directly into seawater.

Spies was unable to obtain successful fertilizations in the laboratory, but found evidence of a mass spawning /

in the field in late December. Pelagic larvae having flabelligerid characteristics were abundant in plankton !1

samples taken at that time and Spies observed their morphology and provided descriptions of several stages

of development from an early 3-segment stage to a late, nearly juvenile 13-segment stage that exhibited ill

evidence of development of anteriorly directed setae. |

Studies of the biology of flabelligerids are needed to understand how these unusual polychaetes live

and contribute to the ecosystems they inhabit. The complex structures of the head are used for a variety of /

functions. For example, we know that the palps are used for feeding, the blood vessels of the palps and I!

branchiae function in respiratory exchange, nephridiopores opening into the dorsal membrane function in

excretion, and the buccal apparatus in general serves as a sensory organ. We know nothing about recruitment ill

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I and habitat selection. Spies (1977) observed that adults of F. commensalis disappeared from the sea urchin

hosts presumably after a mass spawning when the larvae he studied appeared in the plankton. Juveniles

_t subsequently appeared on the hosts, presumably as a result of settlement from those same larvae observed

I in the plankton.

I Taxonomic History

I There has never been a comprehensive monograph on flabelligerids. All revisions or reviews have been regional. For example, important treatments include: Haase (1915: Arctic and boreal species), Fauvel

(1927: French species), Stop-Bowitz (1948: Norwegian species), Day (1961, 1967: South African species),

i and Hartman (1961, 1969: Californian species). i According to Fauchald (I 977) there are 14 genera and 126 species of flabelligerids. Pettibone (I 982)

i indicates that there are 16 genera and about 130 species. The genera Fauveliopsis McIntosh and Ftabelligella

I Hartman, formerly included in the removed to the and Acrocirddae,

Flabelligeridae, were Fauveliopsidae

respectively by Hartman (1971) and Orensanz (1974).

i Day (1961) was the first to use the structure of the head region and the prostomial and peristomial structure to define genera and species. Typically, the oral structures are retracted and require some dissection

in order to observe all features. The position and number of the branchiae, size and shape of lips surrounding

i the mouth, form of the pulps, and location of the median ridge have proven to be critical taxonomic characters. These features, when combined with more readily observed setae, papillae, and body form provide the basis

on which genera and species can be distinguished.

i Fauveliopsis Hartman (1961, 1969) has provided detailed summaries of known species from California. When is removed, Hartman (1969) records six genera and 11 species from California. Two additional

species are recorded in the Santa Maria Basin reports (Lissner et aI., 1986; Hyland and Neff, 1988), and at

I least two additional genera and six species have been found in shelf and slope collections from off northern California (Blake, unpublished). It thus appears that as many as eight genera and up to 21 species may

actually occur in California.

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Key to Genera and Species from the Santa Maria Basin

I 1A. Body covered with a transparent, mucilaginous sheath through which papillae and setae emerge.

............................................................................................................................ Flabelligera affinis

I 1B. Body covered with papillae, sand grains, or silt, not atransparent sheath 2

i 2A. 2B. Ne Neurosetae urosetae from from seg segments ments posteri posterior or toto ccephalic ephalic ccage age inc all lunidentate ude bidentate ................................................. falcigers (Fig. 1.4) ............. 34

I 3A. Dorsum with each branchial filaments in 2

conspicuous paired papillae on segment; numerous, thin,

groups on narrow membrane; anterior postcephalic neurosetae all pseudocompound and bidentate,

; posterior neurosetae crossbarred, both bidentate and unidentate ............................ Piromis hospitis 1i 3B. Dorsum with tips of numerous papillae emerging between sand grains on each segment; branchial

filaments thick, limited to 4 pairs; postcephalic neurosetae pseudocompound and bidentate .........

i ............................................................................................................................... Pherusa capulata
I

4A. Cephalic cage absent or poorly developed, with a few long noto- and neurosetae present on setigers 1- I

=

2; postcephalic neurosetae long, pointed, crossbarred or pseudocompound ..................................... 5

4B. Cephalic cage well developed; postcephalic neurosetae short, curved falcigers ................................ 6 I

5A. Postcephalic neurosetae long, crossbarred, and pointed; body surface covered with elongate papillae, ill

usually with sand grains at base ................................................................................... Brada villosa il

5B. Postcephalic neurosetae include both crossbarred capillaries and pseudocompound setae, sometimes

with hooked tips; body surface usually with papillae and sand grains ............. Genus Diplocirrus 2 I

6A. Cephalic cage formed of numerous long, forwardly directed setae from first 1-4 segments; body ,,_

usually with some adhering sand grains ..................................................... Genus Pherusa ........... 7 II

6B. Cephalic cage covered with dorsal collar or hood, with setae best seen ventrally; dorsum densely

covered with papillae, usually lacking adhering sand grains; commensal with sea urchins •

Flabelligera commensalis II

7A. Papillae mostly limited to anterior margins of segments; dorsum of anterior end oblique, densely I

covered with sand grains .......................................................................................... Pherusa inflata !1

7B. Papillae mostly scattered over entire segment surface, not limited to anterior margins; dorsum of

anterior end not oblique, sand grains covering other parts of body ............................................... 8 I

8A. Dorsum of anterior setigers with long, thickened papillae and remainder of body with numerous Ii

globular, silt-covered papillae ......................................................................... Pherusa neopapiUata |

8B. Dorsum of anterior setigers with numerous very small, narrow to rounded papillae, remainder of

body with few scattered papillae .......................................................................... Pherusa papillata I

Descriptions of Species i

Seven flabelligerid species distributed among four genera were identified from MMS collections. ml

An additional species, FlabeUigera commensalis, is expected to occur in the area. !

Brada viIlosa (Rathke, 1843) Pherusa capulata (Moore, 1909)

Flabeltigera affinis Sars, 1829 Pherusa inflata (Treadwell, 1914) 1

!1 Flabelligera commensalis Moore, 1909 Pherusa neopapillata Hartman, 1961

Piromis hospitis Fauchald, 1972 Pherusa papillata (Johnson, 1901) /

I

! 2Severalspeciesof Diplocirrushavebeenencounteredincollectionsfromthe continentalslopeoffnorthernCalifornia

(Blake,unpublished).Itislikelythatmembersofthisdeep-watergenus_11alsobe foundtooccurofftheSanta

MariaBasin. I

4 i
I Genus Brada Stimpson, 1854

Type Species. Siphonostoma vilIosum Rathke, 1843.

I Diagnosis. Body short, expanded anteriorly, tapering posteriorly, with relatively few segments.

Cephalic cage absent, but notosetae of first segment usually longer than those of following setigers. Epithelial

i papillae but shorterof and variable thickerdensity than notosetae. and form. Branchial Notosetae filaments crossbarrednumerous, capillaries. filiform, Neurosetae in 2 lateral crossbarred groups. Nephridial capillaries

papillae present ventrally on setigers 3, 4, or 5.

I the collections. Remarks. A single, morphologically variable appearing species, Brada villosa, has been found in

iI Brada viUosa (Rathke, 1843)

Figure 1.1

Siphonostoma villosum Rathke, 1843:215, pI. 11, figs. 11-12.

.... Brada villosa: Malmgren, 1867:84.--Fanvel, 1927:121, fig. 43e-l.--Stop-Bowitz, 1948:33-37, fig. 9a-c.--

I Berkeley Berkeley, 1952:7, fig. (synonymy).--Uschakov,

and 5.--Pettibone, 1954:290-292

1955:310, fig. 115D-H.--Imajima and Hartman, 1964:302.--Hartman, 1965:174; 1969:281,5 figs.--

Hobson and Banse, 1981:58, fig. 1lb-c.--Milligan, 1984:47-15 to 47-17, figs. 47-9 and 47-10.

i Trophonia rugosa Hansen, 1882:38, pl. 7, figs. 9-12. Fide Pettibone, 1954.

Trophonia arctica Hansen, 1882:39, pl. 7, figs. 17-20. Fide Pettibone, 1954.

I Brada pilosa Moore, 1906:222, pl. 10. Fide Pettibone, 1954.

Stylarioides pluribranchiata Moore, 1923:222. Fide Pettibone, 1954.

t Bradapluribranchiata rada rugosa: St_p-Bowitz, : Hartman, 1948:37, 1969:279, fig. 10.5Ffigs. ide Pettibone, 1954.

i San Luis,Material Sta. R Examined. California, Santa Maria Basin, off Point Buchon, Sta. 19, 296 m (1); off Port -3 (3); Sta. 106, 492 m (16); off Point Sal, Sta. PJ-6 (1); Sta. PJ-7 (1); Sta. PJ-8 (1, 3); Sta.

R-7 (1); Sta. 34 (1); off Purisima Point, Sta. 45, 395 m (1).--Massachusetts, Georges Bank, numerous

i specimensDesccollected ription.asLength part of the 20-60Georges mm, width Bank Benthi 2-5 mmc Inf forau23-35 nal Monitoring setigers; Santa programMafia(1982- Basin1985), specimens 60-160 m. up

to about 27 mm long, 5 mm wide, up to 25 setigers. Body fusiform, slightly flattened ventrally, thickened,

I somewhat elongate, somewhat tapered posteriorly; clavate papillae appearing (Fig. 1.1B-C), characteristically longest dorsally, rough and usually hirsutewith(Fig. sand 1. grains 1A); atcovered base. Color with

in alcohol tan.

i of dorsolateral Head reprotuberan gion inclucdes es, small pair retra of thi ctileck pgrooved rostomiupalps, m, 2 grou ventral ps ofeversible numerousbuccal fdiformsiphon branchiae surrounding on pair

mouth. Setae of first setigerous segment slightly more elongate than on following segments, directed anteriorly

I (Fig. and neuropodia 1.1A). Pairdistinct, of nephridial wth long papillaefusiform lcatedpapillae near antero grouped v ntralround bordersetaeof setiger (F. 1.1B, 5 (Fig.D).1.1B). Notosetae Notopodi long, a

ringed capillaries, 2-5 per fascicle (Fig. 1.1F). Neurosetae from setiger 2, large, amber-colored, with sharp,

I transparent tips, 3-6 per fascicle (Fig. 1.1E).

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'"..,_ -_-". ' - ' _ '_'_ A

._i._,.:_,_ ..._,._.._.._. .,_...-- . .-. ,

o !

B

,|

.. ... _ ___ ,

nephridum I

Figure 1.1. Brada vitlosa: A, entire animal, lateral view; B, anterior end, lateral view; C, papilla; D, middle I

parapodium; E, neuroseta;F, notoseta. (A, C,F, after StVp-Bowitz, 1948;B, D, E, after Uschakov,

I955). i

Biology. Brada villosa is a widely distributed species occurring in various sediment types including

mud, gravel, stones, and rocks. The species has been found from the intertidal to continental slope depths, i

Remarks. Pettibone (1954), as part of a study of Alaskan Arctic polychaetes, examined the types of

both Bradapilosa Moore from Alaska andStylarioidespluribranchiata Moore from California and determined I

that both were synonymous withB, villosa. Pettibone observed that B. viUosa exhibits considerable variability i

in both the number of papillae and degree of sand encrustation on the body. She determined that specimens

having fewer papillae appeared to have worn them off, especially in larger specimens. The body surface m

beneath the papillae and sand encrustations is rugose and nodular and most frequently exposed on larger nN

specimens.Smallerspecimensappear"haLry"duetothedenseconcentrations ofpapillae.

Among the Santa Maria Basin MMS voucher collections are specimens that were identified as B. am

villosa andB. pluribranchiata. These specimens represent the extremes that were noted by Pettibone (1954). II

For example, specimens with little or no sand encrustation on the body have dense concentrations of papillae

on both dorsal and ventral body surfaces. These were identified as B. villosa. Specimens with dense sand m

encrustations have knobby lumps over their bodies that are apparent when the sand is removed, but surface !1

papillae are absent. These sand-encrusted and knobby forms were identified as B. pluribranchiata. Other

specimens are clearly intermediate between these two extremes (e.g., vouchers from Stations R-7 and Sta. I
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I 34 labeled as P. pluribranchiata and B. villosa, respectively) in that moderate sand encmstation results in

fewer papillae, but with a thickening of their bases. Presumably, these thickenings develop into the larger

' II non-papillated knobs found on heavily sand-encrusted specimens. Because all other features of these

Ii specimens, including setae, parapodial papillae, and form and arrangement of the head region are identical,

the synonymy of B. villosa and B. pluribranchiata, first proposed by Pettibone (1954), is supported. A

I parallel latter species speciesbearscouplet the same /synonymy type of innodular northernlumpsEurope on theinvolves body asBspecimens . villosa and referred B. rugosa to B. pIuribranchiata (Hansen). The

in the California materials. Pettibone (1954) also referred B. rugosa to synonymy with B. villosa. To lend

further support to these proposed synonymies, I examined a series of specimens ofB. villosa from Georges Bank, off Massachusetts. Exactly the same pattern of replacement of numerous small papillae with larger,

sand-encrusted knobby lumps was present, and there was absolutely no difference between these North

,LID Atlantic specimens and those from off California.

II Obviously, the papillate and non-papillate knobby forms of this species present considerable confusion

for biologists attempting to identify them because they superficially appear very different in their extremes. It

i isspecies likely throughout that sand accumulation its life because and abrasion only the have largesta steady specimens and cumulative in the collections effect onwere the body the non-papillated surface of this

knobby forms. The reduction in number of soft surface papillae and the development of enlarged bases of a

i few papillae during growth of the species, provides greater surface area for adhering sand particles. This appears to be an excellent protective mechanism.

Distribution. Widespread in the northern hemisphere, including the Arctic, both sides of the North

i Atlantic, 2,500 m. Mediterranean, Japan, and the eastern Pacific from Alaska to southern California; intertidal to

I Genus Diplocirrus Haase, 1915 ,

Type Species. Trophonia glauca Malmgren, 1867.

I Diagnosis. Body elongate, expanded anteriorly, tapered posteriorly. Cephalic cage weakly developed

or absent. Body surface papillate, often encrusted with sand or debris. Noto- and neurosetae articulated

(compound); neurosetae stouter than notosetae, sometimes with hooked tips. Head region with pair of grooved

I 3 oral and few thickened branchiae of 1 2 in 2

palps, lips, only a (lO) or kinds groups. Nephridial papillae

sometimes present on setiger 5.

Remarks. Diplocirrus was emended by Day (1961) to include species having 1or 2 kinds of branchiae few in number (10 or tess). The genus is closely related tolIyphagus Chamberlin, and according to Fauchald

(1977) is separated only on the basis of body shape. Species of Diplocirrus are elongate, thickened anteriorly

i and tapered posteriorly, whereas species ofllyphagus have short, flattened, nearly disclike bodies. Milligan (1984) considered Ilyphagus to be a synonym of Diplocirrus.

No species of Diplocirrus were encountered as part of this study. However, several species were

I identified from deep-water samples taken off northern California (Blake, unpublished). I.

!

! 7
!

II

Genus Flabelligera Says, 1829 i

Type Species. Flabelligera affinis Says, 1829.

Im

Diagnosis. Body soft, flaccid, translucent, covered with thick mucilaginous sheath supported by I

long, pedunculate papillae. Cephalic cage well developed, formed from long noto- and neuropodial capillaries

of setiger 1. Notosetae annulated capillaries. Neurosetae of setiger 2 and following segments include 1 or

more stout pseudocompound hooks. Branchial filaments numerous, arising in 2 groups of several irregular II

rows from thickened margins of cephalic hood.

Remarks. Two species, F. affinis Says (=F. infundibularis Johnson) and F. commensalis Moore m

occur in California waters. Q

Flabelligera affinis Sars, 1829. I

Figure 1.2

II

FlabelIigera affinis Says, 1829:31, pl. 3, fig. 16.--Fauvel, 1927:113, fig. 40a-f.--Stop-Bowitz, 1948:8-13, B

fig. la-b (synonymy).--Pettibone, 1954:289-290 (synonymy).--Day, 1961:505, fig. 8a; 1967:655-

656, fig. 32.I.a-f.--Hobson and Banse, 1981:58, fig. 1le. i

III

Flabelligera infundibularis Johnson, 1901:417, pl. 12,figs. 124-127.--Hartman, 1938:14; 1948:40; 1961:32;

1969:291, 5 figs.--Berkeley and Berkeley, 1952:7, figs. 1-4. Fide Pettibone, 1954. ii

Material Examined. California, off Point Sal, Sta. R-8 (1); off Purisima Point, Sta. BRA-14, from

a rock, 96-105 m (1).

Description. Large species, up to 130 mm long, 12-15 mm wide including mucilaginous sheath, I

with up to 70 segments; specimen from Sta. BRA- 14, 26 mm long, 6 mm wide, for about 50 setigers. Body

elongate, tapering anteriorly and posteriorly; covered with numerous papillae; papillae elongate, thin, flexible,

usually coiled, with large bulbous tips (Fig. 1.2B), variable in shape. Body usually covered with thick, •

transparent, mucous mantle or covering, penetrated by papillae and setae (Fig. 1.2A); generally colorless in W

alcohol.

Prostomium and peristomium retractile into anterior setiger, with pair of stout, grooved palps, 2 I

groups of numerous f'fliform papillae, 2 pairs of eyes, and protruding buccal siphon surrounding ventral

mouth. First segment with wide, funnel-like depression into which buccal segment may be withdrawn; 111

bearing circlet of elongate, ringed setae around margin, directed forward, forming cephalic cage; numerous II

elongate papillae with slender, straight peduncles and fusiform tips.

Notopodia conical, truncate lobes, with ringed capillary setae (Fig. 1.2C); with elongate papillae on II

posterior side up to three-fourths length of notosetae, similar in form to those associated with cephalic cage, II

but different from those on body. Neuropodia conical, each with 1 or 2 large, hooked, amber-colored

pseudocompound setae, ringed basally, tips dark to black (Fig. 1.2B). m

Biology. FIabelligera affinis is a widespread Arctic-boreal species, occurring on various sediment !|

types of mud, gravel, and rocks; sometimes specimens are found in the water.

Remarks. Pettibone (1954) considered F. affinis to be a widely distributed species, and as part of

her Alaskan Arctic study, synonymized it with the California species, F. infundibularis Johnson. I have not g

been able to find any differences between California specimens and ones from the North Atlantic; the

synonymy is thus supported.

I! Distribution. Widespread in the northern hemisphere, including the Arctic, both sides of the North

Atlantic, Japan, the eastern Pacific from Alaska to southern California. Also reported from the Mediterranean,

and subantarctic locations; low intertidal to more than 2,500 m. I

!

FlabeUigera commensalis Moore, 1909. I

Figure 1.3

! Flabelligera commensalis Moore, 1909,286-288, pl. 9,figs. 62-63.--Hartman, 1961:117-118, fi'ontispiece.--

Light, 1978:682-683. i

Flabelligera ha erens Chamberlin, 1919:16. Fide Hartman, 1961. 1

FlabeUiderma commensalis: Hartman, 1969:285, 3 figs.--Spies, 1973:465-490, pls.l-9; 1975:187-207;

1977:323-345, pls. 1-6. 1

Description. Body elongate, tapering posteriorly, with enlarged cephalic collar or hood concealing

cephalic cage (Fig. 1.3A); dorsal surface and parapodia covered with dense, small papillae, ventrum and l

sides mostly devoid of papillae; each papilla with short stalk and inflated tip; color in life: dorsum and sides II

reddish purple, yellow to orange with green tinge, colors similar to those of host sea urchin. Up to 50 mm

long, 3.2 mm wide, for 70-90 setigers. !ll,

Cephalic cage formed of numerous free setae from first setiger, sometimes concealed by collar into II

which head region retracts; with numerous thin, branchial filaments inserted on broad membrane over small

prostomium; latter with 4 small eyes; palps short, thick. Notopodia with few slender, crossbarred capillaries I

(Fig. 1.3B); each neuropodium with a single curved, pseudocompound hook (Fig. 1.3C). |

Remarks. Flabettigera commensaIis is commonly found on the surface of sea urchins. The species

was originally described inFlabelligera by Moore, butreferred to Flabelliderma (type species, F. essenbergae _l

(Hartman, 1961)) by Hartman (1969) on the basis that it lacks a mucilaginous sheath around the body. Light il

(1978), however, determined that the species actually has a rudimentary sheath and returned it to the genus

Flabelligera. I Distribution. Central and southern California, free-living in kelp holdfasts, commensal with sea

urchins, or with potychaetes; intertidal to shelf depths.

lm

Genus Piromis Kinberg, 1867 _l

Type Species. Piromis arenosus Kinberg, 1867. jw

Diagnosis. Body elongate, expanded anteriorly, fusiform, tapered posteriorly. Cephalic cage well

developed, formed by setae of first few segments. Surface papillate, sand-encrusted. Notosetae crossbarred /

capillaries. Neurosetae forming cage crossbarred capillaries on several anterior setigers; uni- or bidentate B

and simple, or pseudocompound falcigers posteriorly. Branchiae numerous, on single or double long, tonguelike peristomial membrane. Two pairs of prominent prostomial eyes. J

Remarks. Piromis was redefined by Hartman (1961), who combined it and Semiodera Chamberlin. tl

These genera had been previously separated on the basis of single or double branchial membranes,

respectively. Species are separated on the basis of papillae and shape of neurosetae. A single species, P. lJ

hospitis Fauchald, has been found in the collections.

I

I

!
I

i

| ,a ,halic collar

i

I

I

1

I

i

I

I

I

I

I

I

1

Figure 1.3. FlabeUigera commensalis: A, entire animal, dorsal view; B, notoseta; C, neuroseta. (A, after Haxtman,

I 1961; B-C, after Moore, 1909).

!

| 11
!

Piromis hospitis Fauchald, 1972 I

Figure 1.4 i

Piromis hospitis Fauchald, 1972:229-231, pl. 48, figs. a-e.

Material Examined. California, Santa Mafia Basin, off Point Buchon, Sta. R-I, 90 m (2); off I

Purisima Point, Sta. 52, 98 m (4); Sta. BRA-16, 91.5-123 m (1). g

Description. A large, robust species, largest specimen (Sta. R-3) 47 mm long, 6 mm wide across am

expanded anterior region for 50 setigers; other specimens, 10-15 mm long. Body greatly expanded in anterior 1

one-fourth, tapered posteriorly; body surface entirely covered with small sand grains enclosing papillae and

II

parapodia; setae protrude cleanly from body. Dorsum with pair of large, triangular sand covered papillae, II

with narrow, tapering tips on each segment, extending posteriorly along body in 2 rows (Fig. 1.4A); papillae

largest in anterior setigers, becoming smaller posteriorly. Ventrum of each segment with 2 small lateral

g

papillae, 2 mid-ventral papillae; another papilla located just ventral to notopodia and neuropodia; these m

papillae with slight basal enlargement, tapering to rounded tips. Additional papillae on postsetal lobes (Fig.

1.4B). qll

Head region with long prolongation inside branchial membrane; upper lip thick, lateral and lower lib

tips forming semicircle around mouth; branchial filaments numerous, in 2 groups, on narrow membrane. I!

First 3 parapodia directed anteriorly, with setae forming cephalic cage (Fig. 1.4A). Anterior notopodia

with large, triangular postsetal papilla, similar to papillae on dorsum; neuropodia with smaller postsetal i

papillae (Fig. 1.4B); with additional papillae ventral to setal fascicles; notopodia with 1-3 extra papillae 1

dorsal to setae. Neuropodia of middle and posterior segments with postsetal lobes or papillae forming

swollen outgrowth ventral to setal bundles, appearing as pouches. Notosetae crossbarred capillaries; 1/

neurosetae of anterior setigers include 5-6 bidentate multiarticulated hooks (Fig. 1.4C); setae of middle and |

posterior setigers include 5-6 crossbarred hooks, those in superior position bidentate, in inferior position

worn, appearing unidentate (Fig. 1.4D-E). •

Biology. Numerous hydroids are attached to the body surface and setae forming the cephalic cage U

of the large specimen from Sta. R-1. Fauchald (1972) also indicated the presence of hydroids on his Mexican

specimens. The specimens examined here come from inner shelf depths and occur in sands or silty sands. I

Remarks. The paired dorsal papillae are diagnostic for this species. The anterior neurosetae are

distinctIy multiarticulate and bidentate, whereas the posterior neurosetae are crossbarred and include both

bidentate and unidentate forms. However, the unidentate setae are clearly bidentate setae that have been I

worn.

Distribution. Eastern Pacific, central California to western Mexico; 90-230 m.

Genus Pherusa Oken, 1807

N

Type Species. Amphitrite plumosa O.E Mtiller, 1776. I

Diagnosis. Body inflated anteriorly, tapering to narrow posterior end. Cephalic cage well developed,

formed by noto- and neurosetae of first 1-6 setigers. Epithelial papillae usually present, integument often •

covered with sand and debris. Notosetae crossbarred capillaries. Neurosetae of cephalic cage crossbarred

capillaries; subsequent neurosetae include simple uni- or bidentate hooks. Head region with pair of grooved

palps, 2-3 cushion-shaped oral lips, and curved or semi-circular dorsal cephalic hood beating numerous, I

small branchial filaments; lateral margins of branchial arc sometimes spiraled inward. II

Remarks. A well-defined and widely distributed genus, with four species present in MMS materials.

i
I cephalic

cage

noS

I noS

;IC

II

Ii

!

i C D E_/

A B

I

I

Figure 1.4. Piromis hospitis: A, anterior end, lateral view; B, parapodium, anterior view; C, anterior neuroseta;

I D-E,posteriorneurosetaein different degrees of wear.

I Pherusa capulata (Moore, 1909)

Figure 1.5

I Trophonia capulata Moore, 1909:284-286, pl. 9, figs. 60-61.--Treadwell, 1914:212.

Stylarioides eruca: Berkeley and Berkeley, 1941:47. Not Clapar_de, 1870. Fide Hartman, 1969.

Pherusa capulata: Hartman, 1963:55; 1966:405; 1969:295, 2 figs.

Material Examined. California, Santa Maria Basin, offPort San Luis, Sta. R-1 (1).

I Description. A large species, recorded to 108 mm long, 5.5 mm wide, for about 135 setigers;

up

specimen from Sta. R-1 small, 6 mm long, 1 mm wide, with about 30 setigers. Body widest anteriorly,

tapering posteriorly. Body surface densely covered with sand grains, obscuring cuticle in anterior one-half

I of specimen from Sta. R-1 (Fig. 1.5A); papillae coated with sand grains through which elongate, blunt tips

protrude (Fig. 1.5B); approximately 5-6 long papillae on dorsum of anterior setigers; papillae of posterior

segments shorter. Color in alcohol light tan. Buccal region with pair of short, grooved palps and 4 pairs of

I longer, branchial filaments.

| "
I

Cephalic cage formed by long, forwardly directed noto- and neurosetae of setigers 1-2; setae of I

setigers 3-4 also projecting forward, but not as long. Notosetae long, crossbarred capillaries; with those

from notosetae, setigerbecoming 4 thinner. distinctly Neurosetaepseudocompound of setigers 1-3 same and asfalcigerous, notosetae; with from distinctive, setiger 4, neurosetae curved bidentate thicker than tips '1

(Fig. 1.5C).

bidentateRempseudocompound arks. Pherusa falcigers capulata instead is unusofualsetae amongthat the are California crossbarred Pherusa and unidentate. species in having long, I

Biology. Pherusa capulata occurs in coarse sediments having considerable sand, as evidenced by

the presence of sand grains that adhere tightly to the body. I

Distribution. Central and southern California, intertidal to upper slope depths.

!

!

/j

/IJ I

silt I

sand I

I

C

A B I

sand grains

I

Figure 1,5. Pherusa caputata: A, anterior end,dorsal view; B, papilla and basallyadhering sandgains; C,tip of I

neuroseta.

I

I

I

,4 |
!

i Pherusa inflata (Treadwell, 1914)

Figure 1.6

! Trophonia inflata Tmadwell, 1914:213-214, pl. 12, fig. 33.

Trophonia minuca Treadwell, 1914:213, pl. 12, fig. 32. Fide Hartman, 1952.

i Stylarioides dimissus Hartman, 1936:21.

Stylarioides inflata: Berkeley and Berkeley, 1941:46.--Hartman, 1951:98; 1952:71-74, figs. 1-2.

I Pherusa inflata: Hartman, 1961:33 1966:405; 1969:297-299, 5 figs.--Milligan, 1984:47- I 1 to 47-13, figs. 47-7-47 -8.

I Material Examined. California, Santa Maria Basin, off Port San Luis, Sta. BRA-25, 64.5-72 m (3); off Point Conception, Sta. BRA-2, 110-126 m (1).

Description. A moderately sized species, 40-60 mm long, 4-5 mm wide for about 50-70 setigers;

i from Santa Maria 20 1.8 wide for 48 thickened largest specimen Basin, mnl long, mill setigers. Body

anteriody, slightly narrower in median region, tapering to narrow posterior end; anterior end with welldeveloped cephalic cage; dorsum of anterior end oblique, flattened, covered with sand grains; surface of

I body with rows of papillae on anterior margins of each segment, remainder of surface somewhat rough, but lacking papillae (Fig. 1.6A); adhering sand groins limited to broad area on dorsum of a few anterior setigers.

Hartman (1952) reports a pair of large nephridial openings on ventrum of setiger 5. Color in alcohol light

I tan to opaque white, with anterior body sometimes appearing rusty, probably from adhering oxidized sediments.

Head region sometimes everted, with prostomium appearing as small lobe bearing pair of large

I eyes; palps large, thickened, with deep grooves, inserted at dorsolateral margins of prostomium (Fig. 1.6B);

branchial membrane with about 12 large branchial filaments along dorsal margin and 12-13 smaller filaments

laterally; filaments long, tapering, with blunt tips; mouth a narrow slit.

I Cephalic cage formed by long, transversely barred setae from setigers 1-2, notosetae generally

longer and more numerous than neurosetae. Notosetae from setiger 3 also long, curved, projecting forward,

i but not part of cage; about 5-6 per fascicle, becoming shorter, reduced to 3-4 per notopodium by setigers 4- 5 and continuing to posterior end; notosetae simple, thin, crossbarred capillaries. Neurosetae of setiger 3

similar to notosetae, then becoming shorter, thicker, darker, and falcate in subsequent setigers (Fig. 1.6DH), continuing with 3-4 falcate hooks per neuropodium to end of body; hooks with heavy crossbars, sometimes

I appearing weakly pseudocompound.

Biology. Pherusa inflata is typically found in rocky habitats in intertidal and nearshore environments,

associatedRemwitharkssponges and shells.

. Hartman (1952) examined the holotypes of Trophonia inflata Treadwell and T. minuta

Treadwell and established their synonymy. She also redescribed the species, as Stylarioides inflata, based

I uPherusa pon thebytypeHartman as well(1961 as new). Thematerial speciesfroismreadily centralidentified and southern by the Crows alifornia. of papillae The spe on ctheies anterior was referred marginsto

of each body segment.

I Distribution. Oregon to southern California; intertidal to 126 m.

i

!
I

i

t'J

I

?i7

,"'::3"}

_ /.

F _ ,,:

E _ i i

h

G _

+.

pa branchial

filaments

• _lJ

L_, ,-

__i '_ _ I

C _ _c

H

pr

._2_ ,, I

ora] lips_ _ "mouth i

Figure 1,6. brPheanchial rusa inflata:A, entireanimal,lateralview; B, anteriorend, frontalview,with rightpalp andmost i filamentsremoved showinglocationandarrangementofbranchiae,prostomium,eye spots,

and mouth; C, notoseta from middle body segment; D-H, neurosetae from setigers 4, 5, 6, 7, and

middle setiger. (afterHartman, 1961). I

16 |
!

Pherusa neopapillata Hartman, 1961

Figure 1.7

! Pherusa neopapillata Hartman, 1961:121-122, pl. 25, figs. 1-2, pl. 26, figs. 1-7; 1963:56; 1969:301, figs. 1-5.

i Pherusa sp. B. Lissner et al., 1986: Appendix D.

Material Examined. California, Santa Maria Basin, off Point Sal, Sta. PJ-2 (2); Sta. PJ-6 (2, 5);

Sta. PJ-10 (2); off Point Arguello, Sta. BRA-4, from rocks, 168-237 m (1).

Description. Moderate sized species, up to 45 mm long with 45-54 setigers; specimen from Point

Arguello 30 mm long, 2.8 mm wide, with 40 setigers. Body widest in first 4-5 setigers, narrowing slightly

i posteriorly; posterior end not tapered as in P. inflata (Fig. 1.7A). Body covered with numerous papillae, most dense dorsally, less so ventrally, most papillae covered with silt and appearing globular, providing

much of body with appearance of having adhering "mud balls;" individual papillae elongate, tapering distally

i to nipple-like tip, longest papillae on dorsum of first 3-5 setigers, 4-5 per segment (Fig. 1.7B); silted papillae appear globular with protruding narrow tip.

Head region with pair of thick, grooved palps, larger and thicker than 4 pairs of laterally attached

I branchial branchial fifilaments; laments (Fig. prostomium 1.7C); someti small,mconical, es with 2-3 withpairs 4 eyes. of additional short, stubby lobes associated with

Cephalic cage composed of long, forwardly directed crossbarred capillary noto- and neurosetae

i fend. rom Neurosetae setigers 1-3from . Notosetae setigerof4foshort, llowingthick, setigers dark, short, crossbarred thin, crossbarred falcigers 3-4capillaries, per fasciclecontinuing (Fig. 1.7D). to posterior

Biology. Pherusa neopapillata is associated with silty sediments as evidenced by the silt that adheres

i to the papillae on the body. The species occurs in deeper, outer shelf and upper slope sediments than the closely related P. papillata that ranges from the intertidal to inner shelf depths.

Remarks. Pherusa neopapiUata differs from P. papillata in having large, thick and elongate papillae

J on the dorsum of anterior setigers and rest of the body more or less densely covered with numerous, siltcovered papillae that appear globular. In contrast, the papillae ofP. papilIata are smaller, but more numerous

on the dorsum of anterior setigers; the remainder of the body is less densely covered with papillae and they

I are not covered with silt and do not globular.

appear

All specimens identified as P. neopapilIata in the MMS Phase I program are P. papillata, whereas

t a specimeDn identified as P. sp. B is P. neopapillata.

istribution. Southern California, Point Conception to the Mexican border in outer shelf and upper

slope depths; 70-566 m.

I

.!

| 17
I

i

_",,,'ii_ I

' ii!_ " cephalic

_,_--- oaoo

":"_'__:"":i"_/' I

_.._-_ noS

•_.!.. , _..._.._,.

_>_'_' ' I

___'": '_"_'_':"::_ " '" _( i

_:"'_1"

_.'..>_d_ ..

__f_ I

_i:---

'- _ilaments I

I

I

oral lips _ mouth

, |

Figure 1.7• Pherusa neopapitlata: A, entire animal, lateral view; B, anterior end, dorsal view; C, anterior end, I frontal view, showing palps, branchial filaments, scars of same, prostomium, and oral slit; D, neuroseta.

(A, C, D, after Hartman, 1961; B, original).

I
!

i Pherusapapillata (Johnson, 1901)

Figure 1.8

I

Trophonia papillata Johnson, 1901:416, pl. 12, figs. 122-123.

n Stylarioides Stytarioides pluapillata mosa:: Berkeley Berkeley and andBeBerkeley, rkeley, 1941:47. 1952:8-9,Not fig.M_l 6. ler, 1776.

Pherusapapillata: Hartman, 1961:33, 120-121, pl. 24, figs. 1-7; 1963:56; 1966:406; 1969:303, 6 figs.--

I Blake, 1975:222, fig. 277.

Pherusa neopapillata: Lissner et al., t986: Appendix D. Not Hartman, 1961.

I Material Examined. California, off Morro Bay, Sta. 6, 109 m (6); off Purisima Point, Sta. BRA- 16, 91.5-123 m (11); off Point Arguello, Sta. BRA-4, 168-237 m (1); off Point Conception, Sta. 79, 98 m

(1).--La Jolla, intertidal with coralline algae, coll. Aug 1964, L. Hollister (2).

i

I Figure 1.8. Pherusapapillata: A, anteriorend,dorsalview; B, anteriorend,frontalview showingpalps, branchial

filaments, scars of same, prostomium with eyes, and oral slit; C, neurosetae from setiger 14.(A,

i original;B, C after Hartman, 1961).

I

| 19
!

Description. A moderate sized species, to about 50 mm long, 3 mm wide, with about 70 setigers; I

Santa Maria Basin specimens to about 20 mm long, 2 mm wide, for about 45 setigers. Body thick through

about 25 setigers, then tapering posteriorly to blunt pygidium. Dorsum of In-st 6-10 setigers with dense

concentration of small papillae (Fig. 1.8A), remainder of body lightly covered with papillae, fewer on !1

ventrum; papillae short, narrow to conical, sometimes lightly coated with silt.

Head region with pair of thick, grooved palps and 4 pairs of laterally attached branchial filaments I_

(Fig. 1.8B); prostomium small, conical, with 4 eyes. I!

Cephalic cage mainly composed of noto- and neurosetae from setigers 1-2; setae long, anteriorly

directed, crossbarred, pointed spines; setae from setiger 3 also directed anteriorly, but not as long and not i

part of cephalic cage. Notosetae of following setigers short, thin, crossbarred capillaries, continuing to II

posterior end. Neurosetae from setiger 4 short, thick, dark, crossbarred falcigers, 3-4 per fascicle, sometimes

with narrow companion seta (Fig. 1.8C). I

m

Remarks. Pherusa papillata differs from P. neopapillata and other California species by having a

dense concentration of small papillae on the dorsum of some anterior setigers and few papillae over the rest m

of the body. All specimens of P. neopapillata identified as part of the MMS Phase I program are actually P. II

papillata.

Biology. Pherusa papillata has been recorded from the intertidal to upper slope depths (Hartman, Ill

1961, 1963), but appears to be most common from the intertidal to inner shelf depths where it occurs in I

sediments of mixed sand, rock, and silt.

Distribution. Alaska to southern California; intertidal to 560 m. I

Literature Cited I

Amor, A. 1994. Ecology of Pherusa sp. (Polychaeta, Flabelligeridae). In: Dauvin, J.-C, Laubier, L., and

D.J. Reish (eds.), Actes de la 4_me Conf6rence internationaledes Polych_tes. M6moirs du Mus6um i

National d'Histoire Naturelle 162:339-346. aIB

Berkeley, E. and C. Berkeley. 1941. On a collection of Polychaeta from southern California. Bulletin of the I

Southern California Academy of Sciences 40:16-60. II

Berkeley, E. and C. Berkeley. 1952. Polychaeta Sedentaria. Canadian Pacific Fauna 9b (Part 2): 1-139.

i

Fisheries Research Board of Canada. I !

Blake, J.A. 1975. Phylum Annelida: Class Polychaeta. In: Smith, R.I. and J.T. Carlton (eds.). Light's Manual:

Intertidal Invertebrates of the central California coast. University of California Press, Berkeley. Pp. i

151-243. II

Borodin, D. N. 1929. Vergleichende Histologie der Hautorgane bei den Chloraemiden (Polychaeta). Zeitschrift

ffir Morphologie und Okologie der Tiere 16:26-48, pls. 1-2. I

Chamberlin, R.V. 1919. New polychaetous armelids from Laguna Beach, California. Journal of Entomology

and Zoology Pomona 11:1-23. 1

Clapar_de, E. II 1870. Les Annrlides Chrtopodes du Golfe de Naples. Seconde partie. Mrmoirs de la Soci&6

de Physique et d'Histoire Naturelle de Gen_ve 20(2):1-225.

Day, J. H. 1961. The polychaete fauna of South Africa. Part 6. Sedentary species dredged off Cape coasts I

with a few new records from the shore. Journal of the Linnaean Society of Zoology 44: 463-560, 18

figs. I

|
!

i Day, J.H. 1967. A monograph on the Polychaeta of southern Africa. British Museum of Natural History,

Publication No. 656:1-878.

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, 69 pls.

I Fauchald,History K. 1977. Museum The polychae of Los tAngeles e wormsCounty, . Definitions ScienceandSekeriyess to28:the1-190. orders, families and genera. Natural

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.

Fauvel, P. 1927. Polych&es srdentaires. Addenda aux Ermntes, Archiannrlides, Myzostomaires. Faune de

i France 16: 1-494, 152 figs. Haase, P. 1915. Boreale und arktische Chloraemiden. Wissenschaftliche Meeresuntersuchungen 17:169-

228, pls. 1-2.

I Hansen, G.A. 1882. Annelida. Den Norske Nordhavs-Expedition 1876-1878. Zoologi. Oslo. Vol. 3:1-54, 7

pls., map.

I Hartman, O. 1936. Nomenclatural changes involving California polychaete worms. Journal of the Washington Academy of Sciences 26:31-32.

I Hartman,Zoology. O. 1938.Bulle Anntinotated of thelistMuseum of the types of Comparative of polychaetous Zoology, annelids Harvardin the University Museum85:1-31, of Comparative 3 pls.

Hartman, O. 1948. The polychaetous annelids of Alaska. Pacific Science 2: 1-58.

_1 Hartman, O. 1951. The littoral marine annelids of the Guff of Mexico. Publications of the Institute of

Marine Science 2:7-t24.

I Hartman,Pacific O. 1952. Science On the6:71-74. identity ofStylarioides inflata (Treadwell) and its extended distribution (Annelida).

i Hartman,34O.pls.1961. Polychaetous armelids from California. Allan Hancock Pacific Expeditions 25: 1-226,

Hartman, O. 1963. Submarine canyons of Southern California Part 17I. Systematics: Polychaetes. Allan

i Hancock Pacific Expeditions 27(3): 1-93, 4 figs.

Hartman, O. t965. Deep-water benthic polychaetous annelids from New England to Bermuda and other

l North Atlantic areas. Allan Hancock Foundation Publications Occasional Paper No. 28:1-378.

: Hartman,Final O. 1966. resultsQuantitative and conclusions. surveyAllan of theHancock benthos Pacific of the San Expeditions Pedro Basin, 19(2):southern 187-456.California Part 1I.

I Hartman, O. 1969. Atlas of the Sedentariate polychaetous annelids from California. Allan Hancock

Foundation, University of Southern California, Los Angeles, 812 pp.

I Hartman, O. 1971. Abyssal polychaetous annelids from the Mozambique Basin off southeast Africa, with a compendium of abyssal polychaetous annelids from world-wide areas. Journal of the Fisheries

Research Board of Canada 28:1407-1428.

I Hobson, K.D. and K. Banse. 1981. Sedentariate and archiannelid polychaetes of British Columbia and

Washington. Canadian Bulletin of Fisheries and Aquatic Sciences 209:1-144.

!
!

Hyland, J. and J. Neff (eds.). 1988. California OCS Phase II Monitoring Program: Year-one annual report, i

Report prepared for the U.S. Department of the Interior, Minerals Management Service, Pacific

OCS Region, Los Angeles, CA. Contract No. 14-12-0001-30262. Volume 1 (MMS 87-0115) and IN

Volume II (MMS 87-0116). n

Imajima, M., and O. Hartman. 1964. The polychaetous annelids of Japan. I-II. Allan Hancock Foundation,

Occasional Papers 26: 1-452, 38 pls. iw

Johnson, H. P. 1901. The Polychaeta of the Puget Sound region. Proceedings of the Boston Society of

Natural History 29: 381-437, 19 pls. t

Kinberg, J. 1867. Annulata nova. Ofversight af Kungliga Vetenskaps-Akademiens Frrhandtingar, Stockholm W

23: 337-357.

Light, W.J. 1978. Reexamination of the species referred to the genus Flabelliderma Hartman (Polychaeta: D

Flabelligeridae and Acrocirridae). Proceedings of the Biological Society of Washington 91:681-

690. I Lissner, A., Phillips, C, Cadien, D., Smith, R., Bemstein, B., Cimberg, R., Kauwling, T., and W. Anikouchine.

1986. Assessment of long-term changes in biological communities in the Santa Maria Basin and

Western Santa Barbara Channel. Phase I. Final Report submitted for the Minerals Management i

Service, U.S. Department of the Interior, Pacific OCS Office. Contract No. 14-12-0001-30032. 111

MMS OCS Study, MMS 86-0012.

Malmgren, A.J. 1867. Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scandinaviae hactenus m

cognita. Ofversight af Kungliga Vetenskaps-Akademiens F6rhandlingar, Stockholm 24:127-235.

Milligan, M.R. 1984. Family Flabelligeridae Saint Joseph, 1894. In: Uebelacker, J.M. and P.G. Johnson R

(eds.), Taxonomic guide to the polychaetes of the northern Gulf of Mexico. Vol. II. Barry A. Vittor m

& Associates, Inc. Mobile, AL. Pp. 47-1 to 47-20.

i

Moore, J.P. 1906. Additional new species of Polychaeta from the north Pacific. Proceedings of the Philadelphia i

Academy of Natural Sciences 58:2t7-260, pls. 10-12.

Moore, J.P. 1909. Polychaetous annelids from Monterey Bay and San Diego, California. Proceedings of the i

Philadelphia Academy of Natural Sciences 61:235-295, pls. 7-9. HI

Moore, J.P. 1923. The potychaetous annelids dredged by the U.S.S. Albatross off the coast of southern m

California in 1904. Proceedings of the Philadelphia Academy of Natural Sciences 75:179-259, pls. |

17-18.

Mfiller, O.F. 1776. Zoologiae Danicae Prodromus, seu Animalium Daniae et Norvegiae indigenarum

chamcteres, nomina, et synonyma imprimis popularium. Havniae. xxxii + 282 pp. I

Orensanz, J.M. 1974. Poliquetos de la provincia biogeographica Argentina. 10. (Acrocirridae). Neotropica i!

20:113-118. i

Pettibone, M.H. 1954. Marine polychaete worms from Point Barrow, Alaska, with additional records from

the North Atlantic and North Pacific. Proceedings of the United States National Museum 103:203- •

356.

Pettibone, M.H. 1982. Classification of Polychaeta. In: Parker, S.P. (ed.), Synopsis and Classification of m

Living Organisms. McGraw-Hill. Pp. 3-43. H

Rathke, H. 1843. Beitr/ige zur Fauna Norwegens. Nova Acta Leopoldina 20:1-264, 12 pls.

I
!

i Sars, M. 1829. Bidrag til S6edyrenes Naturhistorie. I. Bergen. 59 pp.

Schlieper, C. 1927. Stylarioides plumosus, eine monographische Darstellung. Zeitschfift fiir Morphologie

I und t)kologie der Tiere 7:320-383.

Spies, R.B. 1973. The blood system of the flabelligerid polychaete Flabelliderma commensaIis (Moore).

I Journal of Morphology 139:465-490. Spies, R.B. 1975. Structure and function of the head in flabelligerid polychaetes. Journal of Morphology

147:187- 207.

I Spies, R.L. 1977. Reproduction and larval development of FIabelliderma commensalis (Moore). In: Reish,

D.J. and K. Fauchald (Eds.), Essays on Polychaetous Annelids in Memory of Dr. Olga Hartman.

i University of Southern California Press, Los Angeles. Pp. 323-345, 6 pls. St_p-Bowitz, C. 1948. Les Flabellig6riens Norv6giens. Bergens Museums /_rbok 1946-1947,

Naturvitenskapelig rekke, No. 2:1-59, 12 pls.

I 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 Uschakov, P.V. 1955. Polychaeta of the far eastern seas of the USSR. Opredeliteli po Faune SSSR 56:1- 445. Academiya Nauk SSSR. (In Russian). (Translated by the Israel Program for

Scientific Translations, U.S. Dept. Comm. Wash. D.C. 419pp., 1965)

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iiIIIllIIIllIiIIIl _ i
!

i 2. FAMILYAcRocmamAr BANSE, 1969

I by

James A. Blake 1

I

Introduction

!

The Acrocirridae are a small family of sedentary polychaetes that are believed to be deposit feeders.

: i prostomium The three knisowreduced n generaandhabears ve species a pair woifthpalps an elloocnga atedte, onslethenderfrontal body margin; with numer eyesousmaysegbemepnresent ts. Theor

absent. The pharynx is unarmed and the small ventral buccal sac is only weakly eversible. Branchiae may be

I present compoundor absent. The body often bears numerous papillae. Setae usually include hollow looking neuropodial falcigers. Acrocirrids were formerly included among the cirratulids, but were removed to a

separate family by Banse (1969). They range from intertidal and shallow waters to abyssal depths.

I Morphology

I The head region of acrocirrids is formed by a small prostomium fused to the peristomium. The

prostomium is reduced, pentagonal, pointed anteriorly, and lies dorsal to the peristomium (Fig. 2.I A). The

I head attachedis retractile in Flabelligella, but not retractile in Acrocirrus and Macrochaeta. A pair of palps is frontally and two pairs of eyes are usually present. The mouth is subterminal; the unarmed pharynx

includes a weakly eversible sac. When present, f'diform branchiae begin on the second segment and usually

i iothers. nclude four pairs arising above the notopodia; the second pair is always inserted more dorsally than the

The body is roughly divided into thorax and abdomen, the boundary between the two occurring at

I the 13th segment (genus Acrocirrus), but is often not distinct. The peristomium usually overlies segment 1. The first three body segments are typically achaetous. Parapodia axe biramous, but reduced, with small setal

lobes; postsetal lobes, and interramal and subpodial papillae may be developed to varying degrees. Notosetae

I are capillaries, striated or spinous. Neurosetae are large, compound falcigers with a hyaline membrane below the tip of the blade. The cutting edges of blades of thoracic neurosetae are directed posteriorly,

whereas in abdominal segments the cutting edges of neurosetal blades are directed anteriorly. Some species

tl of Acrocirrus have modified acicular hooks on an anterior abdominal segment, usually 11 or 14. A pair of nephridia is present in the thoracic region and a nephxidiopore may sometimes be seen as

a short papilla below the branchiae on segment 3 (e.g., species of Acrocirrus). The pygidium generally lacks

I appendages, or small papillae may be apparent.

I _ENSR, 89 Water Street, Woods Hole, Massachusetts 02543.
i

Biology i

m

Despite their widespread distribution, acrocirrids are not commonly encountered. As a result, tittle

is known about their biology. They have been found under rocks in the intertidal zone and in soft sediments I

in deeper waters. Flabelligella appears to be limited to the deep sea, whereas Acrocirrus and Macrochaeta I

are found in shallower habitats. Acrocirrids are believed to be selective deposit feeders with food particles 1

carried to the mouth along ciliated grooves in the palps (Banse, 1969; Pettibone, 1982). dl_

Gonads are present throughout the abdomen (Okuda, 1934). Banse (1969)observed free floating

eggs in the coelomic cavity of anterior abdominal segments in some species of Acrocirrus. Nothing is i_

known concerning the reproductive biology or development of acrocirrids.

Taxonomic History I

Banse (1969) established the Acrocirfidae for the generaAcrocirrus and Macrochaeta, genera that I

had previously been included in the Cirratulidae. Banse also pointed out the close affinities of acrocirrids

with flabelligerids, and it is now quite clear that the two families share many affinities with one another, I

including compound and pseudocompound setae. Detailed anatomical studies of acrocirrids, however, have Ii

not been completed, precluding further phylogenetic comparisons with other polychaetes. Orensanz (1974)

transferred the deep-water flabelligerid genus FlabelligeIla Hartman to the Acrocirridae. I1

The most il detailed account that includes descriptions, illustrations, and keys of the then known

species of the family is by Banse (1969). Additional species have been described by Kudenov (1975, 1976)

and Banse (1979). These authors have provided updated keys. Wolf (1984) reviewed the Gulf of Mexico I!

acrocirrids. I

In the eastern Pacific, five species of Acrocirrus and one species of Macrochaeta have been reported

and documented (Banse, 1969, 1979; Kudenov, I975). At least one species of Flabelligella has been 1

discovered in slope sediments off northern California (Blake, unpublished). Acrocirrus crassifilis Moore D

and A. heterochaetus Annenkova have been reported from California (Moore, 1923; Banse, 1969; Hartman,

1969). I

Key to Genera and Species of Acrocirridae from California i

1A. Branchiae present from segment 2, usually numbering 4 pairs;epithelium smooth to densely papillate

........................ * ............ *---* ............................................................................................................... 2 iI

imp lB. Branchiae absent; epithelium densely papillate ................................................ Genus Flabelligella

2A. Palps at least as long as f'n-st10 segments, with bases close together; thoracic region always with 12 I

segments; body smooth, or with sparse papillae ........................................ Genus Acrocirrus ....... 3

2B. Palps short, with bases separated by at least width of both individual palps; number of thoracic 1

segments variable .............................................................................................. Genus Macrochaeta U

Segment 14 (setiger 11) with pair of large, modified acicular hooks; neurosetae from third branchial I

3A.

segment ..................................................................................................... Acrocirrus heteroehaetus

n

3B. Segment 14 (setiger 1!) lacking modified hook; neurosetae from fourth branchial segment l_

........................................................................................................................ Acrocirrus crassifilis II
!

i Description of Species

I A Acrocirrus heterochaetus Annenkova has been identified from the MMS Phase I

single species,

collections. The second species,A, crassifilis, was not encountered, but it is differentiated in the key (above).

I Genus Acrocirrus Grube, 1872

Type Species. Heterocirrus frontifilis Grube, 1860.

I Diagnosis. Body sparsely papiltated, with papillae sometimes arranged in rows; thorax always

consisting of 12 segments. Palps long, located close together. Branchiae from setiger 2, numbering 4 pairs.

I Promine neurosetaent papil simplela alwor compound ays presenthooded below seco hooks; nd pair sometimes of branchiae with large, on segment modified3. Notosetae acicular hook long,onannsegment ulated;

14 (setiger 11).

I Remarks. Acrocirrus is the largest genus of the Acrocirridae with 12 described species (Banse, 1979). At least two species, A. crassifilis Moore and A. heterochaetus Armenkova have been reported from

California (Banse, 1969). The latter species has been discovered in the MMS Phase I collections examined

I as part of this study.

i Acrocirrus heterocha F etus Annenkova, 1934 igure 2.1

I Acrocirrus heterochaetus Annenkova, 1934:326; 1938:183.--Hartman, 1948:38.--Uschakov, 1955:304.-- Banse, 1969:2605-2606, fig. ld, 4a-c.

Acrocirrus sp. A: Lissner et al. 1986: Appendix D.

I Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA-20, 90-130.5 m

(1). Western Santa Barbara Channel, Sta. BRC-1, 73.5-78 m (1).

I Description. Specimen from Sta. BRA-20 complete, but in 3 pieces, 21 mm long, 0.5 mm wide, for

68 setigers; reported with up to 130 setigers (Banse, 1969). Body with epithelium sparsely papillated, most

papillae concentrated in rows on parapodia (Figs. 11B-D). Color in life bright orange to yellow; in alcohol,

I dusky pigment on prostomium, peristomium, segments. Body

tan with and most of thoracic thickened

anteriorly, tapering to narrow posterior end.

i Prostomium blunt anteriorly, clearly separated from peristomium by groove; with middorsal crest separating origins of palps; palps reported as long as 15-20 segments (Banse, I969); pair of large, black

eyes and second, smaller pair sometimes visible (Fig. 2.1A, B). Branchiae numbering 4 pairs, each thick,

i about as long as 7-8 segments (Fig. 2.2A). Single papilla located dorsal to first branchia; rows of papillae associated with podial lobes of anterior setigers (Figs. 2.1B-C). Neuropodia well developed, consisting of

rounded lobe bearing several short papillae (Fig. 2.1C) and fascicles of 3-4 long, compound falcigers (Fig.

11E); cutting edges of neurosetae directed posteriorly through segment 12 (setiger 9), thereafter directed

I but with shorter blades than in of thorax. Acicular hook with anteriorly, setae neuropodial enlarged tip

present on segment 14 (Fig. 2.1D, G). Notosetae from segment 5 (setiger 2), each with narrow jointed,

almost camerated appearance (Fig. 2.IF). Pygidium a simple lobe with terminal anus surrounded by minute

I papillae.

!
!

Remarks. Acrocirrus heterochaetus belongs to a group of species with a modified spine on segment 14. I

Distribution. Bering Sea and Alaska to southern California, in rocky habitats, in shelf depths;

western Pacific, Sea of Japan, Sea of Okhotsk; shelf to middle slope depths. I

pa _ I

branchiae ,_ ' I

I

I

°:L _

I

i

I

Figure U,1. Acrocirridae: A, anterior end of a representative aerocirrid, Acrocirrusfrontififis; B-G, Acrocirrus

heterochaetus: B, anterior end, lateral view; C, sixth segment, anterior view; D, segments 13-15, I

lateral view; E, compound falciger, F, notoseta with inset showing detail of fine serrations; F, acicular

hook from segment 14. (A-D, after Banse, 1969; E-G, original).

I

!

_8 |
i Literature Cited

I Annenkova, N.P. 1934. Kurze Ubersicht der Polychaeten der Litoralzone der Bering-Insel (Kommandor- Inseln), nebst Beschreibung neuer Arten. Zoologischer Anzeiger 106:322-331.

•I Annenkodistribution]. va, N.P. 1938.Akademiia [PolychaetaNauk, of theLeningrad. northern paDal'nevostochnie n of Sea of Japanfilial, and Trudy their hoGidrobiologischeskoi rizontal and vertical

Ekspeditsii sin an 1934, AN SSSR, 1:81-230. (In Russian)

I Banse, K. Canada 1969. Acrocirridae n. ram. (Polychaeta Sedentaria). Journal of the Fisheries Research Board of 26:2595-2620.

I Banse, K. 1979. Acrocirrus columbianus and A. occipitalis, two new polychaetes (Acrocirridae) from the northeast Pacific Ocean.

Grube, A.E. 1860.Beschreibungen neuer oder weniger bekannter Anneliden. 4. Zahlreiche Gattungen. Archiv

I fiir Naturgeschichte 26:71-118, 3 pls.

Hartman, O. 1948. The polychaetous annelids of Alaska. Pacific Science 2:3-58.

I Hartman, O. 1969. Atlas of the sedentariate polychaetous annelids from California. Allan Hancock Foundation, University of Southern California, Los Angeles, 812 pp.

I Kudenov, J.D. 1975. Sedentary polychaetes from the Gulf of California, Mexico. Journal of Natural History 9:205-231.

Kudenov, J.D. 1976. Polychaeta from southeastern Australia 1. Acrocirridae Banse, 1969, from Victoria

I and New South Wales. Records of the Australian Museum 30:137-I49.

Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bernstein, R. Cimberg, T. Kauwling, T. and W. Anikouchine.

I Western 1986. Assessment of long-term changes in biological communities in the Santa Maria Basin and Santa Barbara Channel. Phase I. Final Report submitted for the Minerals Management

Service, U.S. Department of the Interior, Pacific OCS Office. Contract No. 14-12-0001-30032.

i Moore, MMS OCS Study, 86-0012. J.P. 1923. The polychaetous annelids dredged by the U.S.S. "Albatross" off the coast of southern

California in 1904. IV. Spionidae to Sabellariidae. Proceedings of the Academy of Natural Sciences

I Philadelphia 75:179-259.

Okuda, S. 1934. The polychaete genusAcrocirrus, from Japanese waters. Journal of the Faculty of Sciences,

i Hokkaido Imperial University, series 4, 2:197-209. Orensanz, J.M. 1974. Poliquetos de la provincia biogeografica Argentina. X. Neotropica 20:113-118.

i Pettibone, M.H. 1982. Classification of Polychaeta. In: Parker, S.P. (ed.), Synopsis and Classification of Living Organisms. McGraw-Hill. Pp. 3-43.

Wolf, P.S. 1984. Chapter 13. Family Acrocirridae Banse, 1969. In: Uebelacker, J.M. and P.G. Johnson

I (eds. Associates, ), Taxonomic Inc. Mobile, guide toAL.thePp. polchaetes 13-1 to 13.7. of the northern Gulf of Mexico. Vol. 2. Barry A. Vittor &

i Uschakov, P.V. 1955. Polychaeta of the far eastern seas of the USSR. Opredeliteli po Faune SSSR 56:1- 445. Academiya Nauk SSSR. (In Russian). (Translated by the Israel Program for

Scientific Translations, U.S. Dept. Comm. Wash. D.C. 419pp., 1965)

!
IlIIIIlIIIIIlIIIII

_°n
I 3. FAMILY FAUWUOPSlDAr HARTMAN, 1971

| uy

James A. Blake 1and Mary E. Petersen 2

!

i Introduction

The Fanveliopsidae are a small family of detritus-feeding, sedentary polychaetes comprising about

I 20thespec flabelligerid ies, severalgenus of whi Brada ch arewithi unnamed, n whichalltwoin the species genus(FFauveliopsis . gIabra and. FM. abrevis ny species ) weresuoriginally perficiallydescribed. resemble

However, these and all other fauveliopsids differ from flabelligerids in lacking head appendages (see below).

I Lhasikeresulted the flabelligerids, in a numbertheofprostornium species having and perist ther oposte miumriorcaends n be described retracted within as the the antefirst rior 3-4 end.setigers, which

Most fauveliopsids are less than 2 cm long. They are free-living or inhabit the empty shells of

I Bathysiphon scaphopod or gastropod molluscs, or the tubular tests of sand-agglutinating foraminiferans (genus ). They have been reported from depths ranging from about 20 m (e.g., Wolf, 1984; Riser,

1987) to nearly 7000 m (Levensteirt, 1970), and from a variety of sediment types: sand, mixed bottoms,

I reported mud, clay,fromandthediatomhigh ooarctic. ze. They are known from almost all of the world' s oceans but have not yet been

Very little is known about the biology of this family, and the only detailed morphological study

I published to date is that of Riser (1987), who investigated the histology of several species and described the internal morphology in connection with the description of F. arenicola, from New Zealand. Langhage

(1994) studied aFauveliopsis sp. close to (but not identical with) F. cf. adriatica of Katzmann and Laubier

I with the aid of light- and electron microscopy; the latter study has not yet been published. The family is currently being investigated by one of us (MEP), and a taxonomic review and partial revision will be published

elsewhere.

!

Morphology

i The body of fauveliopsids is elongate and cylindrical or club-shaped (Fig. 3.1) (spiraled like that of

a gastropod if f:txed in a gastropod shell), often slightly tapering and truncate at both ends, with from 16 to

I about 40 segments; a single species (F. arabica) has over 85 segments. The cuticle is smooth or minutely

rugose, usually with some fine papillae that are easily overlooked under lower magnifications; there are no

appendages anywhere except for a minute, mushroom- or pear-shaped interramal papilla between the notosetae

I and the neurosetae.

!

1ENSR, 89 Water Street, Woods Hole, MA 02543.

i 2Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen t3, Denmark.
I

There are usually 2-3 body regions recognizable either by body shape, changes in setation or both. I

The anterior region has 2-4 segments with modified setae, either tapering and weakly or strongly falcate, or

nearly straight and with blunt or irregular tips. Setae of this region are always somewhat heavier than those In

of the following segments. In fixed material the prostomium and peristomium are usually partly or completely II

retracted within the anterior region, and extended prostomia have only been observed a few times. The

pharynx is unarmed and probably not extensible. I

The middle and posterior segments usually form either one region with the simple parapodia described l

above and considered typical for the family, or two regions where posterior segments differ in having

multiple neurosetae and sometimes also multiple notosetae. In cases where middle and posterior regions are n

different, the posterior region is often somewhat inflated. One (or two) genital papillae, first described by

Riser (1987), are present immediately anterior to the anteriorly directed setae of one of the middle parapodia.

The papilla is usually low and somewhat flattened, and is easily overlooked, but at least in some species I

(e.g., Fauveliopsis glabra, Fig. 3.2B) it appears to be capable of extension and then resembles a true papilla. I

When only one papilla is present it appears to occur on the right side; when two are present they occur on

both sides of the same segment. The amount of variation in location of the genital papilla(e) is stillincompletely _i

known, but for the species treated here it appears to be a useful means of distinguishing between different l

taxa when used together with other features.

In some species living in mollusc shells or foraminiferan tests, a more or less thickened, smooth or n

rugose calluslike area, herein referred to as a ventral shield, has been observed on some (mainly) posterior II

segments. The function of this region is not yet fully understood, but it seems to occur in the area of greatest

contact between worm and shell (or test), along the ventral surface in contact with the columella in species IN

inhabiting gastropod shells, and may aid the worm in maintaining its position inside the shell; in some

species it is very conspicuous, in others (e.g., F. glabra) it appears to be present but is inconspicuous and

easily overlooked. The ventral shield will be considered in more detail in connection with a review of the •

family (M.E. Petersen, in prep.). This is the first report of this feature in the literature. II

Pampodia are biramous throughout, but protruding parapodial lobes are rudimentary or absent except

in posterior segments of some species, and setae are few, typically with two in each ramus except in anterior n

and posterior segments: a tong, slender, smooth pointed (capillary) seta outermost and a much thicker, i

sigmoid acicular spine innermost. Anterior and posterior segments may have greater numbers of setae, with

those of the anterior segments differing in structure from those of the following ones. i

I

i

I

I

I

Figure 3.1. Fauveliopsis glabra (SantaMaria Basin, Sta. PJ-9), entire animal in ventral view. i
!

I The pygidium is terminal and either unadorned or surrounded by a few small papillae. At least in

some species it is partially retractable within the last segment (e.g., see fig. 1 in Katzmann and Laubier,

I prostomium, 1974); the abhas ilitycaused to retract a number the pygidium, of misinterpretations together withof the the pygidial orientationpapillae of the anworm. d the usually retracted

Sexes appear to be separate, and observed variation in oocyte size (Riser, 1987) suggests that

I belong development to thismay familybehave eitheryetdirect ben (lecithotrophic) recognized. or with a planktonic stage, but no larvae that might

I Taxonomic History

I details The most recent review of famity characters of Fauveliopsidae is that by Wolf (1984); for additional see this. Reviews of described species have been provided by Katzmann and Laubier (1974, includes

key to 12 known species and possibly distinct taxa), Amoureux (1982, 13 species and unnamed taxa compared

I by number of segments), and Hartmann-Schr6der (1983, table with distribution of capillaries and acicular setae in 11 species).

The family Fauveliopsidae was erected by Hartman (1971) for ten species in four "flabelligeridI like genera", the main differences being lack of a cephalic cage and lack of the retractile oral branchiae and tentacles characteristic of Flabelligeridae. Three of the original four genera have since been referred elsewhere:

Flabelligella Hartman, 1965, 2 species, by Orensanz (1974) referred to Acrocirridae; Bruunilla Hartman,

i 1971, 1 species, by Pettibone (1979) referred to Polynoidae; Flota Hartman, 1967, 2 species, by Fauchald (1977) and Wolf (1984) considered uncertain status, by Santos and Silva (1992) referred to Acrocirridae.

The Fauveliopsidae were placed in the order Fauveliopsida, by Fauchald (1977), and treated as a member of

I Flabelligerida, together with Acrocirridae andFlabelligeridae, byPettibone (1982) and George and Hartmann- Schrfder (1985).

The type genus, FauveIiopsis Mclntosh, 1922, is thus the only recognized genus in the family,

I which presently contains 16 named species and several unnamed ones. The type species, F. challengeriae Mclntosh, 1922, is a fairly large species (17 mm long), described from a single complete specimen from

1950 fathoms (3569 m) depth in the Antarctic Ocean. It was found in a hard brownish "tube" of blackish to

I brownish sand grains, probably the test of a sand-agglutinating foraminiferan of the genus Bathysiphon Sars. Mclntosh (1922) compared the species to several other taxa, includingBrada and Trophonia (= Pherusa)

of the Flabelligeridae, but did not place it in this family although he (Mclntosh, 1922:7) concluded that "On

I the whole, Fauveliopsis appears to approach the Chloraemidae." The genus was characterized, inter alia, by a rudimentary prostomium, a blunt cephalic region with bristles on each side, and dorsal and ventral bristles

with a pear-shaped papilla between them. The species was figured in some detail and the epithelium was

I described as pale, iridescent and without papillae; it is shown as completely smooth.

The genus and species were not mentioned again until Hartman (1966a) published the second of her

Antarctic papers, by which time she (Hartman, 1960, 1965) had described two other species, both as

I flabelligerids of the genus Brada: B. glabra Hartman, 1960, from southern California, redescribed herein, and B. brevis Hartman, 1965, from the North Atlantic. Reviewing Antarctic records seems to have made

Hartman aware of the affinities of these worms, as the next (fourth) species described by her (Hartman,

I 1967) was placed in the genusFauveIiopsis: F. brevis Hartman, 1967 (=Homonym ofBrada brevis Hartman, 1965), from the Antarctic. Brada glabra was fu'st transferred to Fauveliopsis by Hartman (1969), and later,

together with B. brevis Hartman, 1965, again by Hartman and Fauchald (1971, genus emended); both papers

I treated the genus as a flabelligerid. It was first later that year that Hartman (1971) erected the family Fauveliopsidae and referred the AntarcticBrada brevis to Fauveliopsis, renaming the homonymF, brevipodus

Hartman, 1971 (later corrected to F. brevipoda by Fauchald and Hancock, 1981).

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From 1970 the number of species grew rapidly, with new species described in papers by Levenstein i

(1970): F. hartmani, from the western Pacific, Japan and Kurile-Kamchatka Trenches, corrected to F.

hartmanae by Levenstein (1971/72); Hartman (1971 ): F. arabica, from the Arabian Sea, formally described i

in Hartman (1975); Hartman and Fauchald (1971): F. scabra, the northwest Atlantic; FauchaId (1972a): F. II

brattegardi, western Norway; Fauchald ( 1972b): F. rugosa, western Mexico; Katzmann and Laubier (1974):

F. adriatica and F. fauchaldi, the Adriatic Sea; Fauchald and Hancock (1981): F. armata and F. magna, •

northeast Pacific off central Oregon; Amoureux (1982) F. cabiochi, northeast Atlantic; Hartmann-Schr6der l

(1983): F. olgae, northeast Atlantic; and Riser (1987): F. arenicola, New Zealand, southwest Pacific. Another

three species were distinguished in part, but not named in Hartman (1971): Fauveliopsis sp., off northeast lib

Africa; in Katzmann and Laubier (1974): F. cf. adriatica, Adriatic Sea; and in Wolf (1984): Fauveliopsis |

sp. A, northern Gulf of Mexico. Finally, additional information onF. challengeriae was provided by Hartman

(1978). Most of the above taxa and a new species of Fauveliopsis from the Faroe Islands (collected by the i

BIOFAR Project) will be treated in more detail elsewhere, and a new genus will be proposed for F. cabiochi B

and related species (M.E. Petersen, in prep.). Other species from deep water off California will likewise be

treated separately (Blake, in prep.). i

Many descriptions of fauveliopsids have placed a great deal of emphasis on the number of segments i

as a means of separating species. However, while a few species appear to have segment constancy, most do

not, and of those that do, there appear to be several species with the same number of segments. It is seldom •

possible to identify any species on a single character, and fauveliopsids are no exception. Accurate II

identifications require careful observation of the entire animal, not just one aspect of it.

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34 B
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I Key to Species of Fauveliopsis from California

I 1A. Genital papillae paired, on posterior edge of setiger 6, anterior to setae of parapodia 7; segments 1- 4 with modified setae, with up to 4-5 thick + 4-5 thin setae per ramus; cuticle smooth, completely

transparent, shiny, without transverse striations; body of similar width throughout ........................

I ........................................................................................................................... Fauveliopsis brevis _

lB. Genital papilla single, on right side of setiger 10, 11, 12, or 13 just anterior to right parapodium of

following setiger; segments 1-2 or 1-3 with modified falcate setae; cuticle rough, opaque, only

I transparent in posterior half of body, with numerous fine transverse striations and minute papillae

(latter not always visible without staining); body tapering at both ends or inflated posteriorly .... 2

I 2A. (Note three choices) Genital papilla small, buttonlike to slightly elongate, on posterior part of setiger

10, just anterior or dorsal to notosetae of parapodium 11 (rarely paired and present on left side also);

i modified neurosetae of setiger 1 large, recurved; neurosetae of setigers 2 and 3 normal falcate; posterior segments with multiple neuro- and notosetae; ventral shield inconspicuous, elongate oval,

broadly rounded anteriorly; specimens usually in shells of gastropods or scaphopods ...................

I ........................................................................................................................... Fauveliopsis glabra

I 2B. (Note three choices) Genital papilla small, buttonlike, on posterior part of setiger 11, just anterior to

anterior to notosetae of setiger 12; modified notosetae of setiger 1 curved, not recurved falcate;

I nandeurosetae slender ofcompanion setigers 2-3seta also in each normaramus; l falcventral ate; posterior shield absent; setigersknown with only singlefromheavyshellrecurved of scaphopod spine

.......................................................................................................................... Fauveliopsis armata

I 2C. (anterior Note three to notosetae choices) ofGenital setigerpapilla 13 or large, 14; modi conspicuous, fied notosetaeon posterior of setigers part 1-2ofstrongly setiger curved, 12 or 13,spines just

of following segments gently curved; posterior setigers with a single heavy curved spine in notopodia

I with notoaciculars slender cthi omcker panionthansetaneuroaciculars; and 2-3 heavyventral spinesshield in neuropodi inconspicuous; a with habitat slender unknown companion............ setae,

........................................................................................................................... Fauveliopsis magna

I

Description of Species

I Remarks. One species, Fauveliopsis glabra, is present in the MMS samples from the Santa Maria

Basin, it is likely that this is the only species found in continental shelf depths and the upper slope off

I from Californi deepa. Inwater conjunoffctioOregon, n with were this stud examined y, the tand ypes compared of 17. armatawithandtheF.material glabra, ofbothF. original glabral

.y Additional described

specimens taken from box cores off northern California in slope and rise depths were also examined as well

I ataken s additional by the DSRV specimeAlvin ns .from the Gorda Ridge hydrothermal fields off northern California and Oregon,

I

I _A single specimen of F. brevis was identified from samples takenfrom hydrothermal ventfields on the Gorda Ridge.

This specimen was compared with the holotype of F. brevis (Hartman, 1965) from the North Atlantic; no

differences were observed.The speciesis distinctive in the sleek, shiny nature of the cuticle that is reminiscent

of some opheliids.This is the first record ofF. brevis fromthe Pacific; the species will be described in detail in

I a separatepaperdealing withLhepolychaetesfromhydrothermalvents of theGorda Ridge(Blake,inpreparation).
!

The results of these observations indicate that at least four species occur in the offshore sediments I

of California and Oregon.

Fauveliopsis armata Fauchald and Hancock, 1981" lower slope depths off Oregon. I

Fauveliopsis brevis (Hartman, 1965): deep water off northern California, associated with

hydrothermal vents. Not described in this chapter.

_mt

Fauveliopsis glabra (Hartman, 1960): shelf and slope depths of California; the only species found •

in the Santa Mafia Basin. U

Fauveliopsis magna Fauchald and Hancock, 1981: lower slope depths of Oregon and California. 1

Ill[

Fauveliopsis glabra (Hartman, 1960)

Figures 3.1-3.3 I

Brada glabra Hartman, 1960:129-130, pl. 14, figs. 1-2; 1963:54; 1966b:398; (?)1965:172-174. I

[] Fauveliopsis glabra: Hartman, 1969:283-284, 2 figs (first use of new combination but not indicated as such

or commented); 1971:1422.--Hartman and Fauchald, 1971:116-117 (stated to be new

combination).--Levenstein 1972:173, table 2, fig. 1--Fauchald and Hancock, 1981:37 (key), 38, •

51 (table 1).--Amoureux, I982:192 (list).--Hartmann-Schr6der, 1983:183 (table 2).--Wolf,

1984:48-1 (review).--Riser, 1987:213, 215, fig. 11.

Fauveliopsis sp. A.: Steirdaauer and Imamura, 1990: Appendix F-1. I

Material Examined. California: off San Francisco, Gulf of the Farallones, R/V FarnelIa, Sta. J-13,

U.S. Navy 103 Disposal Site Survey, 37°40.012_r, 123°27.45'W, 15 Aug 1990, 2770 m (3, ZMUC POL- 1

739); R/V Wecoma, U.S. Navy 103 Disposal Site Survey, Sta. 13-2,37°40.36'N, 123°27.96'W, 19 Jul 1991,

2701 m (1); Sta. B-19, 37°31.39'N, 123°24.78'W, 23 Jul 1991, 2425 m (1); R/VPointSur, EPA 102Survey,

Sta. 4-12, 37°10.78'N, 123°09.87'W, 17Sep 1991, 1480 m (3); Sta. 4-15, 37°12.42'N, 123°15.81'W, 19Sep I1

1991, 1730 m (1, regenerating); Sta. DS-3, 37°09.86'N, 123°22.71'W, 24 Sep 1991, 2755 m (I).---Santa

Mafia Basin, 135-165 m, MMS Phase II Monitoring Surveys: Cruise 1-1 (Oct 1986), Stas. PJ-1 (3, LACMAHF); PJ-2 (2, LACM-AHF); PJ-3 (1, LACM-AHF); PJ-4 (18, LACM-AHF); PJ-5 (29, LACM-AHF); PJ- •

6 (1, LACM-AHF); PJ-7 (3, LACM-AHF); PJ-8 (10, LACM-AHF); PJ-9, rep. 1 (15, S13MNH); rep. 2 (6,

USNM); PJ-11 (9, LACM-AHF); PJ-13 (5, LACM-AHF); PJ-19 (1, LACM-AHF). Cruise 1-2 (Jan 1987),

Sta. R-2 (1, LACM-AHF); Sta. PJ-6 (2, LACM-AHF); Sta. PJ-7 (1, LACM-AHF); PJ-23 (1, LACM-AHF). []

Cruise 1-3 (May 1987), Stas. R-2 (3, LACM-AHF), PJ-8 (9, LACM-AHF); PJ-9 (29, ZMUC POL-736, 737, I

738); PJ-11 (7, LACM-AHF); PJ-22 (1, LACM-AHF); PJ-23 (6, LACM-AHF). Cruise 3-1 (Oct 1988): PJ1(5, LACM-AHF).----San Nicholas Basin, off San Clemente Island, R/V Velero/VSta. 6341, 15 Aug 1959, []

1670 m, olive green clay, holotype (LACM-AHF Poly 0525). l

Description. Body more or less club-shaped (Fig. 3.1, 3.3A), rounded, truncate anteriorly, either

straight or coiled, denoting collection from scaphopod or gastropod shell; one elongate 34-setiger specimen •

from Gulf of the Farallones with body fusiform, thickest at setiger 16, then narrowing posteriorly, 8.4 mm

long, 1.3 mm wide; specimen from Sta. DS-3 coiled, with 37 setigers. Santa Mafia Basin specimens up to 6

mm long, 0.6 mm wide across setiger 3, 1.1 mm wide across posterior section, with 36 setigers. Atlantic •

populations reported to be up to 8 mm long, with up to 47 setigers (Hartman and Fauchald, 1971). Color in

alcohol dusty whitish or yellowish to light tan, sometimes with black, dusty intersegmental pigment; oocytes

pinkish, visible through body wall. I

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I Prostomium usually invaginated, causing some contraction of setiger 1 (Fig. 3.3A). Body narrow

anteriorly, with well defined segmental annulations through setigers 11-13, becoming swollen and saelike

i posteriorly, numerous, fine, withtransverse segmentalridges furrows (multian poorlynulate defined or rugose), (Fig. 3.1, providing 3.3A). dull Bodyappesurface arance,relatively at low magnifications smooth, with

dusty, most pronounced on anterior half of body but present throughout most of ventral surface; in stained

i individuals and in profile, transverse ridges interrupted at more or less regular intervals by minute, distally pointed micropapillae, about 6-6.5 m high (Fig. 3.3E). Cuticle appearing thick anteriorly, thin and stretched

posteriorly, becoming transparent, with internal organs visible through body wall (Fig. 3.3A). Largest

i specimens with weakly developed ventral groove in posterior one-third of body; smaller specimens (<4 mm long, 30 segments) with flattened area on ventrum on about setigers 13-18 or 15-20. Ventral shield poorly

developed, inconspicuous, elongate oval, broadest anteriorly, present midventrally on posteriormost segments

i as an irregularly shield-shaped smooth, transparent area. Last few segments of posterior region with minute, low, rounded papillae ventrally and slightly larger ones dorsally.

Body with anterior, middle and posterior setal regions. Parapodia slightly dorsolateral throughout;

I, protruding parapodia aparapodial rising directly lobesfromwellbody developed wall; setae only from in posterior midbodyregion, to endwithof worm setae ofarising anterior fromandsmooth, most middle shiny

parapodial welts resembling pockets and extending laterally as true parapodia in posterior setal region.

I Anterior with setal region comprising setigers 1-3, with modified setae (Fig. 3.2A): notopodia of setiger 1 each 2 (or 4) weakly falcate, slender, subequal spines; neuropodia of setiger 1 usually with 2 (occasionally

4) falcate spines; upper spine thick, broad, and strongly hooked; lower one straighter and slender, about

i one-third as wide as upper one; noto- and neuropodia of setigers 2-3 each with 2-4 slender, subequal falcate spines similar to those of notopodia of setiger 1. Middle setal region beginning on setiger 4, extending to

about setiger 22; apart from occasional doubling, all parapodia with single aeicular spine and capillary seta

i in each ramus, with spines initially being more slender than those of setigers 1-3 (Fig. 3.2B, 3.3C), then becoming shorter, thicker and more curved from setiger 14 or 15 to 22. Posterior setal region beginning

about setiger 23 and characterized by segments with multiple neumsetae and sometimes also multiple

notosetae, i.e., 2-3 acicular spines and 2-3 capillaries in each ramus (Fig. 3.3D). Neuroaciculars of anterior

I region usually larger or stronger than notoaciculars; neuroaciculars of middle and posterior regions slightly smaller and more slender than notoaciculars.

i Interramal papilla stalked, close to notosetae on setigers 1 and 2, shifting to more medial position between rami of setiger 3; papillabecoming smaller, more sessile from about setigers 6 to 10, then becoming

closer to notosetae.

I transversely Genital ovalpapil weltlaonusposterior ually retracted part of(extended setiger 10,andimmediately papillalike anterodorsal on figured specimen) to (appears andto bevisible behind)as

anteriorly oriented setae of right parapodium 11 (Fig. 3.2B). A single female specimen (May 1987, Sta. PJI 9, leftRepside3,asfrowell. m gastr Larger opod females shell) withwithooposte cytesrior120half x 135of coelom tamin diameter filled withhaseggs similar100-112 papillalx(mretra in cdiameter. ted) on the

Pygidium bluntly rectangular or circular flattened dorsal lobe bending ventrally, forming cover

I over anus (Fig. 3.1); dorsal border sculpted, with 8 papillae or lobes (Fig. 3.3F); anus terminal.

Biology. Specimens were often coiled around the columella of gastropod shells, or inside empty

scaphopod shells. The gut of the largest specimens is filled with f'me silt and a few coarser particles including

i foraminiferans.

The presence of what appear to be two genital papillae on one specimen with oocytes is not easy to

explain; other specimens with oocytes have only one papilla and do not appear to differ morphologically.

I The difference in sizes in this and other is considered indicative of slight oocyte specil21ens not specific

differences.

II The maximal size of P. glabra is larger on the slope (8.4 mm long) than on the shelf (6.0 mm long).

i

I 37
I

Figure 3.2. Fauvetiopsis glabra (Santa MariaBasin,Sta.PJ-9). A, anterior end in lateralview showingmodified ,:_

setae of setigers 1-3 and following setigers; B, lateral view of sefigers 8-12, showing structure and I

location of genital papilla (gen P) and interramal papillae (int P) on setiger 10.

I

Remarks. Although Hartman and Fauchald (1971) indicate the combination Fauveliopsis glabra

to be new, Hartman had already used this combination, without any comment or indication that it was new,

in her Atlas of Polychaetes (Hartman, 1969).

In the original and subsequent descriptions of this species by Hartman (1960; 1969), the orientation I

of the animal was reversed in the text description (Riser, 1987), but the figures appear to be correct, the i

location and size of the small interramal papilla in fig. 2 being typical of posterior segments, not anterior

ones. The figured middle segment, stated to be from setiger 16, is in good agreement with the present

material. The narrow end with three setigers beating paired falcate spines in the noto- and neuropodia is the

anterior end. The inflated, saclike end is the posterior end. The maximal number of setigers appears to be 36

in the present material. According to Hartman (1960; 1969), the maximal number of setigers was 33. The

interramal papilla is close to the notosetae in setiger 1, shifting to a middle location on setiger 2 and continuing I

posteriorly as described above.

By comparison with most Brada, in which genus this was originally placed, the cuticle is smooth, I

but it is not as smooth as originally described; the anterior half of the animals tend to appear dusty, even

when cleaned of any loose sediment particles. What appears to be a partial sloughing off of the cuticle of

specimens in good condition suggests that the surface layer may be shed. i

Although the basic pattern of the middle region (setigers 4 to about 22) is one capillary and one

acicular seta per ramus, the presence of many segments with extra setae, without any apparent pattern to

their distribution, suggests some variability in this character.

Riser (1987) was the first to draw attention to the genital papilla(e), and suggested that their number

(1 or 2) and location might prove to be useful specific characters. From the material we have observed to I

date, both treated here and in preparation, this assumption appears to be correct. |

38 II
I

I_piHa

100 pm

]

I ,

I rm,armmal loqple

i .......... '" fQ&

I

I Figure 3.3. Fauveliopsis glabra (Santa Maria Basin, Sta. PJ-19). A, entire animal in dorsal view; B, setiger 2, anterior view; C, setiger 4, posterior view; D, setiger 30, posterior view; E, papilla from anterior part

of body; F, papillae from around pygidium.

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Fauveliopsis armata Fauchald and Hancock, 1981 _1

Figure 3.4

II

Fauveliopsis armata Fauchald and Hancock, 1981:37-38, pt. 7, figs. g-i. II

Material Examined. Oregon, continental slope, Sta. AD6, NAD17, 44°33.5'N, 12504.6'W, 2000 1

m, coll. 6 June 1963, A.D. Carey, holotype (LACM-AHF Poly 1I60). I

Description. Holotype with body more or less club-shaped, rounded truncate anteriorly (Fig. 3.4A), lll

straight, denoting collection from a scaphopod; 4.0 mm long, 0.7 mm wide and with 28 setigers. Color in 1

I

alcohol light tan.

Prostomium invaginated, but with rounded lobe apparent on anterior margin (Fig. 3.4A). Body •

narrow anteriorly, with segmental annutations and furrows poorly def'med, and apparent only on anterior |

borders of noto- and neuropodia of setigers 3-12; subsequent segments swollen, with no evidence of segmental

furrows, until last 4-5 setigers, where podial lobes better developed. Body surface, with moderate numbers ,•

of randomly dispersed, minute papillae on anterior setigers (1-6); papillae becoming sparse over following |

segments, then becoming more numerous and arranged in transverse rows on last 5-6 setigers; anterior

papillae elongate, narrow, apically pointed, becoming low andnipplelike on rest of body. Numerous transverse I

ridges apparent, especially on anterior segments; cuticle appearing thick anteriorly, thin and stretched l

posteriorly, with digestive tract visible through body wall (Fig 3.4A). Ventral shield not apparent.

Body with anterior, middle and posterior setal regions. Parapodia slightly dorsolateral throughout; •

protruding parapodial lobes weakly developed from setiger 4, becoming more prominent over following I

segments and well developed in last 6 setigers (Fig. 3.4A). Noto- and neurosetae consisting of single heavy

medial spine and accompanying narrower spine throughout; some neuropodia from middle body segments •

with extra spine intermediate in thickness to main spine and companion seta. Anterior setal region comprising 1

setigers 1-3, with modified gently curving, pointed spines about twice thickness of those of setiger 4 (Fig.

3.4B) and following segments (Fig. 3.4C-D); last 6 setigers with large, blunt-tipped recurred spines (Fig. •

3.4E). I

InterramaI papilla stalked on anterior segments (Fig. 3.4B-C), becoming sessile and smaller by

about setiger 15 (Fig. 3.4C-D), continuing to posterior end; papilla attached on inner border of enlarged I

notopodia on last 6 segments. I

Genital papilla short, globular on posterior part of setiger 11,immediately anterodorsal to anteriorly

oriented setae of right parapodium 12 (Fig. 3.4A).

Pygidium a bluntly flattened terminal lobe bearing 8 or more minute papiUae.

Biology. The type specimen was likely taken from the inside of an empty scaphopod shells. No 1

gametes were observed in this specimen. 1

Remarks. Superficially, Fauveliopsis armata closely resembles F. glabra in body shape, setal

configuration, and sparse papfllafion of the body surface. However, F. armata has 2 setae per ramus throughout •

the body instead of multiple setae in posterior neuro- and sometimes also notopodia, a body with indistinct I

segmental furrows instead of very distinct ones in anterior setigers, the genital papilla arises from the posterior

right side of setiger 11 instead of setiger 10, acicular spines in far posterior parapodia are strongly recurved 1

instead of more gently curved, and the acicular spines and companion setae are mostly limited to one per I

ramus instead of up to 2-4 spines in some posterior rami. The number of segments is not important in

separating these two species because the range has not been established for F. armata. The holotype, the Is

only known specimen confirmed at this time, has 28 setigerous segments, whereas F. glabra has up to 36 l

setigers in California examples and has been reported with up to 47 in Atlantic specimens (Hartman and

Fauchald, 1971). The latter specimens, however, have not been examined as part of this study and confirmed 1

as belonging to the same species. I

° I
I Figure 3.4. Fa4,posterior uveliopsiview s armata ; C, s(ethoigloterype 13,,poLACsterior M-AHF viewPoly ; D, setiger 1160).A, 20,ent posterior ireanimviewalin; E, dosetig rsalvierew26,; Bpo, sestertigiorer

view.

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Fauveliopsis magna Fauchaid and Hancock, 1981 I[

Figure 3.5 l

Fauveliopsis magna Fauchald and Hancock, 1981:38. mR

Material Examined. Oregon, Cascadia Abyssal Plain, Sta. AD9, NAD21, 44°36.4'N, 125°24.8'W, n

2800 m, coll. 13 Aug 1962, A.D. Carey, holotype (LACM-AHF Poly 1149).---California, Gulf of the

FaraUones, R/V Wecoma, U.S. Navy Site Selection Survey, Sta. B-12, 37°34.28'N, 123°28.29'W, 22 Jul

1991, 2700m (2, ZMUC POL-740); R/V Point Sur, EPA 103 Survey, Sta. DS-2, 37°10.50'N, 123°20.52'W, 1

23 Sep 1991, 2394m (3).

Description of Holotype. Holotype with 41 setigers, 8.0 mm long and 1.0 mm wide (11.0 mm long

and 1.2 mm wide, Fauchald and Hancock, 1981); body elongate, narrow, slightly twisted, but generally II

straight except for curved posterior region (Fig. 3.5A); body narrowest anteriorly, somewhat dorsoventraily w

flattened (setigers 1-8), then somewhat thicker and more cylindrical over long middle region, widest in n

curved far posterior region consisting of last 12 setigers, this part expanded, curved toward ventrum (Fig. 'l

3.5A). Ventral shield weakly developed on curved ventrum. Body with numerous fine transverse ridges

w

apparent, especially on anterior segments; cuticle appearing thick throughout; internal organs not apparent n

through body wall. Body surface, with numerous minute, narrow papillae. Color in alcohol light tan. II

w

Prostomium invaginated, but with rounded lobe apparent on anterior margin; with weak ventral

notch. Setae of first 3 setigers mostly broken off; California specimens with 2 enlarged, recurved notosetae n

and 4 similar neurosetae on first 2 setigers (Fig. 3.5B). Segmental furrows well defined through first 20 n

setigers, less so over last 20 setigers. Parapodia of setigers 1-3 directed anteriorly, poorly developed; parapodia

of setigers 3-8 weakly developed with low podiai lobes, but with well developed interramal papilla shifted

slightly toward notopodium (Fig. 3.5C); interramal papillae stalked with expanded tip on anterior segments, II

becoming sessile and smaller by about setiger 20 (Fig. 3.5D), continuing to posterior end; papilla attached

on inner border of enlarged notopodia on last 20 or so segments. Podial lobes better developed from setiger

9 to about setiger 20, then becoming large and very prominent through posterior half of body. Posterior 1

parapodia with large, noto- and neuropodia bearing prominent acicular spines and small sessile interramal

papilla shifted to location high on notopodium (Fig. 3.5E). n

Body with anterior, middle and posterior setal regions. Noto- and neurosetae consisting of single 1

heavy medial spine and accompanying narrower spine in anterior and middle setigers; notosetae generally

thicker than neurosetae throughout, both gently curved, not recurved; posterior parapodia of last 12 setigers II

with spines more numerous and thicker (Fig. 3.5E); notosetae consisting of single thickened, curved spine 1

and companion spine about one-half thickness; neurosetae with 2-3 thick spines and 1 thin, pointed seta.

Genital papilla large, elongate, on posterior part of setiger 12 (Fig. 3.5A), immediately anterodorsal •

to the anteriorly oriented setae of right parapodium 13. I

Pygidium a rounded terminal lobe bearing 8-10 minute papillae (Fig. 3.5A).

Notes on California Specimens. Specimens from Farallones Sta. B-12 with bodies elongate, truncate, I

clavate anteriorly, tapering posteriorly, usually with slight inflation anterior to last 5 setigers; larger specimen

14.5 mm long, 1.0 mm wide, 40 setigers, extended genital papilla on posterior edge of setiger 12, as shown

for holotype, ventral shield inconspicuous, all anterior setae broken, parapodia of middle and posterior II

segments with 2 notosetae, parapodia of last 12 segments with multiple neurosetae; smaller specimen 7.0

mm long, 1.5 mm wide, 28 setigers, regenerating anterior to genital papilla, ventral shield narrow hourglassshaped, rounded anteriorly. Specimens from Sta. DS-2 with more or less club-shaped bodies, largest with m

40 setigers, 7.0 mm long, 1.0 mm wide; 2 smaller specimens with 35 and 41 setigers, respectively, latter

regenerating. Genital papilla located on posterior margin of right setiger 13 on all specimens. First 2 setigers

with heavy modified recurved neurosetae. Posterior of body expanded, curved ventrally, inflated dorsally. 1

42 I
I

i

i

I

I

Figure 3.5. Fauvetiopsis magna. A, entire animal in left ventrolateral view (holotype, LACM-AHF Poly 1149);

I B, setiger 2, posterior view (Sta. DS-2); C, setiger 8, posterior view (Sta. DS-2); D, setiger 22 (holotype), posterior view; E, setiger 35 (holotype), posterior view.

I

i 43
I

Remarks. Fauveliopsis magna is distinctive in having a higher number of body segments (35-41) •

than most known species. Locally, F. magna resembles F. glabra in setal configuration, but differs in

having the genital papilla arising from setiger 12-13 instead of 10, by having a more or less uniformly •

elongate body rather than one which expands in the middle, by having the small surficial papillae present I

over most of the body instead of limited mostly to the anterior and posterior ends, and by having up to 41

body segments instead of 26-36. The species resembles the type species of the genus, F. challengeriae, in •

general appearance but differs in having the genital papilla on setiger 12 or 13 rather than on 11, in the more I

rounded shape of the parapodia, and in having the multiple neurosetae in posterior segments rather than 2

setae per ramus throughout the body. •

The known depth range of P. magna is limited to the lower slope from about 2400 to 2800 m, l

whereas, P. glabra exhibits a wide range of depths from the continental shelf to the lower slope (135-2755 m).

I

Acknowledgements

I

Prof. Dr. Wilfried Westheide, Universit_t Osnabriick, is thanked for making available a copy of the i

thesis by Petra Langhage. Ms. Leslie Harris of the Los Angeles County Museum of Natural History is

thanked for loaning types of key species of fauveliopsids, i

Literature Cited I

Amoureux, L. 1982. Ann61ides Polych_tes recueillies sur la pente continentale de la Bretagne _ l'Irlande,

Campagne 1973 de la "Thalassa" (suite et fin) avec la description de quatre esp_ces nouvelles pour l

la Science. II. Inventaire taxonomique annot6 de toutes les Polych_tes Srdentaires. Cahiers de

Biologie Marine 23:179-214.

aa

Fauchald, K. 1972a. Some polychaetous annelids from the deep basins in Sognefjorden, western Norway. i

Sarsia 49:89-106.

Fauchald, K. 1972b. Benthic polychaetous annelids from deep water off western Mexico and adjacent areas i

in the eastern Pacific Ocean. Allan Hancock Monographs in Marine Biology 7:575 pp.

Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families and genera. Natural •

History Museum of Los Angeles County, Science Series 28:1-188. I

Fauchald, K. and D.R. Hancock. 1981. Deep-water polychaetes from a transect off central Oregon. Allan

Hancock Foundation Monograph 11:1-73. l

n

George, J.D. and G. Hartmann-Schrrder. 1985. Polychaetes: British Amphinomida, Spintherida and Eunicida.

In: Kermack, D.M. and R.S.K. Barnes (eds.): Synopses of the British Fauna (New Series), No. •

32:221 pp. Published for The Linnean Society of London and The Estuarine and Brackish-Water 1

Sciences Association by E.J. Brill/Dr. W. Backhuys, London-Leiden-Krln-Copenhagen.

Hartman, O. 1960. Systematic account of some marine invertebrate animals from the deep basins of southern i

California. In: Hartman, O. and J.L. Barnard: The benthic fauna of the deep basins off southern

California. Part II. Allan Hancock Pacific Expeditions 22:69-216.

Hartman, O. 1963. Submarine canyons of Southern California Part TIT.Systematics: Polychaetes. Allan i

Hancock Pacific Expeditions 27(3): 1-93, 4 figs.

!
!

I Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other North

Atlantic areas. Allan Hancock Foundation Publication, Occasional Paper 28:378 pp.

I Hartman, O. 1966a. Polychaeta Myzostomidae and Sedentaria of Antarctica. Antarctic Research Series 7:158 pp.

I Hartman,Final O. 1966b. results Quan and ticonclusions. tative surveyAllan of theHancock benthosPacific of the Expeditions San Pedro Basin, 19(2):187-456. southern California Part II.

Hartman, O. 1967. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises, chiefly

I from Antarctic seas. Allan Hancock Monographs in Marine Biology 2:1-387.

Hartman, O. 1969. Atlas of the sedentariate polychaetous annelids from California. Allan Hancock Foundation,

I University of Southern California, Los Angeles, California. 812 pp. Hartman, O. 1971. Abyssal polychaete annelids from the Mozambique Basin off southeast Africa, with a

compendium of abyssal polychaetous annelids from world-wide areas. Journal of Fisheries Research

I Board, Canada 28:1407-1428.

Hartman, O. 1974 [1975]. Polychaetous annelids of the Indian Ocean including an account of species collected

I byof the members speciesoffrom the Internati India. oJnal ournal Indian of theOcean Marine Expediti Biological ons, 1963-'64 Association and a catal of India ogue16(1): and bibliography 191-252.

Hartman, O. 1978. Polychaeta from the Weddell Sea Quadrant, Antarctica. Paper 4 in Biology of the Antarctic

I Seas VI. Antarctic Research Series 26:125-223.

Hartman, O. and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England to Bermuda

i and other North Atlantic areas. Part 1I. Allan Hancock Monographs in Marine Biology 6:1-327. Hartmann-Schrfder, G. 1983. Zur Kennmis einiger Foraminiferengeh_iuse bewohnender Polychaeten aus

dem Nordostatlantik. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut

I 80:169-176.

Katzmann, W. and L. Laubier. 1974. Le genre Fauveliopsis (Polych_te sddentaire) en Mrditerrange.

i Mikrofauna Meeresbodens 50:1-16. Langhage, P. 1994. Morphologische Untersuchungen an einer Fauveliopsis-Art (Annelida, Polychaeta).

Unverrffentlichte Diplomarbeit, Biologie, Universit_t Osnabriick. 80 pp.

I Levenstein, R.Ya. 1970. New and rare species of the abyssal genus Fauveliopsis Mclntosh (Polychaeta,

Annelida) and the peculiarities of its distribution. Transactions of the P. P. Shirshov/nstitute of

• • Oceanology 88:227-235. [In Russian, English summary]

| Levenstein, R.Ya. 1971/72. Ecology and zoogeography of some Polychaeta representatives of the abyssal

Pacific. Proceedings of the Royal Society of Edinburgh 03) 73(18): 171-181.

I Mcintosh, W.C. 1922. Notes from the Gatty Marine Laboratory, St. Andrews. No. XLIV. 1. On new and

rare Polychaeta, Gephyrea, etc., from various regions. Annals and Magazine of Natural History,

ser. 9, Vol. 9:1-30, pls. 1-3.

I Orensanz, J.M. 1974. Poliquetos de la provincia biogeografica Argentina. X. (Acrocirridae). Neotropica

20(63):113-118.

I Pettibone, M.H. 1979. Redescription of Bruunilla natalensis Hartman (Polychaeta: Polynoidae), originally

referred to Fauveliopsidae. Proceedings of the Biological Society of Washington 92(2):384-388.

I
!

Pettibone, M.H. 1982. Annelida. In: Parker, S. (ed.): Synopsis and Classification of Living Organisms. Vol. I

2:1-43. McGraw Hill Book Company, Inc. New York.

Riser, N.W. 1987. A new interstitial polychaete (Family Fauveliopsidae) from the shallow subtidal of New I

Zealand with observations on related species. Bulletin of the Biological Society of Washington

7:211-216.

Santos, P.J.P. dos and V.M.A.P. da Silva. 199211993. Macrochaeta westheidei n.sp.: first record of I

Acrocirridae for the Brazilian coast (Annelida; Polychaeta). Nerftica, Curitiba 7(1-2):7-12, Ed. da

UFPR. I

Steinhauer, M. and E. Imamura. 1990. California OCS Phase II Monitoring Program Three Year Annual

Report Volume I. Submitted to the U.S. Department of the Interior, Minerals Management Service,

Pacific OCS Region, Under Contract No. 14-12-0001-30262. I

Wolf, P.S. 1984. Chapter 48. Family Fauveliopsidae Hartman, 1971. In: Uebelacker, J.M. and P.G. Johnson

B(eds.): arry A. Taxonomic Vittor andGuide Associates, to the Polychaetes of the Northern Gulf of Mexico. Vol. 6:48-1 to 48-4. IInc., Mobile, Alabama.

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| 4. FAMILYCAPITELLIDAEGRUBE, 1862

| by

James A. Blake 1

I

i Introduction

Capitellids are burrowing, earthworm-like potychaetes that have a short, conical or pointed

i prostomium and an elongated body divided into two distinct regions. There are no prostomial or peristomial appendages and branchiae are rare. In general, the anterior or thoracic region bears capillary setae whereas

the posterior or abdominal region bears hooded hooks. The numbers of segments in these regions, the detail

I and initiation of the hooks, and presence or absence of other structures such as branchiae, nuchal organs, or genital hooks, have served to establish a highly complex genetic hierarchy.

Capitellids are among the most commonly encountered polychaetes in marine soft sediments. They

i from the intertidal to the and often the dominant of benthic infaunal range deep sea are components

communities especially in sediments that are organically enriched. Because of their accessibility and

importance in sedimentary environments, capitellids have been the subject of numerous ecological studies.

I

Morphology

I

Capitellids range from very small (1-2 mm), threadlike forms to large robust earthworm-like forms

exceeding 20 cm in length. The body bears a small prostomium that lacks appendages. It is divided into a

I well-defined and thickened thorax with a defined number of segments and a long, narrow,

sometimes tatteredappearing abdominal region that has a variable number of segments. Parapodia are simple tori, but better

developed in abdominal segments.

i Sensory structures include a pair of eversible nuchal organs (ciliated pits when not everted) located

near the posterior border of the prostomium. Eyes are present or absent on the prostomium; when present

'i eyes part oftenthed toprostomium be small, suandbdermal, are sometimes and appearrecessed as pairedintosing theleperistomium. or multiple spLateral ots dorsola senseterally organsonandtheneph posteri fidioarl

openings are often present along the body.

I number The of thoracic thoracicsetigers, region theincludes presencethe/abachaetous sence of setae peristomium on the peristomium, and up to 20andsetthe igrodistribution us segments. of setal The

types (capillaries and hooks) are critical taxonomic characters that distinguish one genus from another.

i Unfortun development ately, ofthe structure of the thoracic region changes during development, and because the postlarval so few capitellids is known, juveniles of species in one genus can be misidentified as

belonging to another genus if critical characters have not yet developed.

l sometimesTheenlarged abdominalandregi swollen on mayasbecompared modified tointhose some ofgentheera.thoracic Parapodiregion. a, especial Branchiae, ly the neifuropo present, dia, are

always on abdominal segments. Branchiae may be simple or branched and relxactile or non,retractile. When

i prese a caudal nt, branchi cirrus. ae serve as diagnostic characters for some genera and species. The pygidium normally bears

i 1ENSR,89WaterStreet,WoodsHole,Massachusetts02543.

I 47
!

The setal arrangement in thoracic segments is probably the most important taxonomic character •

used to def'me the genera. The various patterns can be illustrated diagrammatically, and this has been attempted

by several workers (Hartman, 1947; Ewing, 1984a). Readers are referred those papers and that of Fauchald •

(1972) for diagrams and tables of setal organization of the capitellid genera. The presence of capillaries and I

mixed fascicles in abdominal segments is also important, and most generic summaries emphasize the transition

between thorax and abdomen as being important. The large genital hooks are usually found in two of the last in

thoracic setigers. |

The structure of capitellid setae, especially of the hooded hooks may be quite elaborate. Typically,

the hooks have a relatively straight shaft with a distinct node or manubrium, usually at the point were the

hook emerges from the body. The main fang is usually thick and bent at aright angle. This fang is surmounted I

by few to many apical teeth in 1-4 rows, of variable complexity. A large, inflated hood typically surrounds

the teeth; the hood opening is small or large, depending upon species. The structure of these hooks and their fl

number in a fascicle may be critical species-level characters. m

Taxonomic History I

The capitellids represent one of the oldest groups of polychaetes known to science. The early history I

of the family was reviewed by Eisig (1887). The most important modern work on capitellid morphology and

systematics is that of Hartman (1947). In this work Dr. Hartman established the modern framework and

definition for the genera, provided details of the true nature of the hooded hooks, and reviewed or tabulated _l

most of the then known species. She directly defined and treated 11 of the 21 known genera and described J

and carefully illustrated 18 species, including six that were new to science.

Over the next 30 years, numerous additional taxa were described; Fauchald (1977) reported that the I

number of genera had increased to 36. Fauchald provided a key to these genera, a table outlining their W

critical characters, and brief diagnoses of each. Additional genera have subsequently been added including:

Amastigos Piltz, 1977, Dodecaseta McCammon and Stull, 1978, Nonatus Amaral, 1980, and OctocapiteUa I

Brown, 1987 among others. Clearly, with the increased number of genera that are still based largely upon m

thoracic segmental and setal distributions, the entire family is in need of detailed taxonomic review and

systematic analysis. Two important faunal accounts of capitellids are by Hartman (1969) for California and t.

Ewing (1984a) for the Gulf of Mexico.

One of the major problems with capitellid systematics is that the generic definitions are based upon

numbers of thoracic setigers, numbers of thoracic setigers with capillaries, and the first appearance of i

hooks. While these arrangements can be mapped and charted, it is clear from developmental studies, that

development of setal patterns in capitellids begins with only a few anterior setigers bearing capillaries. With

continued growth, subsequent setigers become modified and the pattern changes. During development through I

juvenile phases it is likely that a single species could pass through stages that approximate several genera !

until its final configuration is achieved. To prove this point, Fredette (1982) demonstrated that juveniles of ±

H. fiIiformis exhibited fewer capillaries on their thoracic setigers than adults in that the initial number of

thoracic setigers with capillaries is three. With growth, hooks present on setigers 4-5 are replaced with

capillaries. This same pattern has been observed by me in development of Neomediomastus and

Neoheteromastus in specimens from the continental slope. For the most part, these sorts of problems with I11

the capitellid genera have been ignored and the proliferation of genera has continued.

Hutchings and Rainer (1981) reviewed the taxonomic history the type species of Heteromastus, H.

filiformis, from the Mediterranean. Among other things these authors established a neotype, but from a i

locality in Egypt rather than the original location from the French coast. These results define the characters U

of the type species, H. filiformis, and permit comparison with specimens from distant localities. The genus

Capitella has been reviewed by Warren (1976a, 1991). Capitella capitata has been determined to represent I

|
i

I a suite of sibling species (Grassle and Grassle, 1976) and this discovery has stimulated considerable followon investigation (see below for details). The genus Mediomastus has been recently reviewed by Warren et

i aI (Hartman) . (1994). This and Mgenus . californiensis presents someHartman difficultihave es tobeen Northreported Americanas dominan workerstbecause components two species, from both M, ambiseta the east

and west coasts. There has, however, never been a comprehensive comparison of these widespread populations

i toNotomastus determine wasif they redefined are indeed by Ewing the same (1982), specie buts there or perhaps as beenclosely no reviewrelatedof this siblinggenusspecies. since The Hartman's genus

monograph in 1947. Ewing (1984b) emended the definitions of Mastobranchus and Peresiella and described

I new species of both genera from the Gulf of Mexico.

I Biology

Infaunal Community Interactions. Most of the common capitellids are found in offshore habitats,

I tolerant althoughofsome a wideof range the best of temperatures known speciesandocsalinities. cur in nears These hore species and estuari are often ne hacolonizers bitats and ofaredisturbed apparentlyor

stressed habitats. For the most part, capitellids inhabit soft sediments.

I Capitellids are among the most common polychaetes found in benthic infaunal samples. In some situations, species of Capitella and Mediomastus are dominants. For example, M. ambiseta was by far the

most abundant invertebrate in the Santa Mafia Basin MMS stations in the 90-170 m depth range (Blake,

i Massachusetts 1993). In New Bay, England oftenwaters, reachingM.densities californiensis of moreis than the m5000 ost abindividuals undant capite m-2l(Blake lid in ietnner al.,shelf 1987).depths In Newin

England, Mediomastus ambiseta has also been well documented as a colonizing species following episodes

I of positive disturbance response(Santodareas ers etthat al.,have 1980;been Grasorganically sle and Grassle, enriched. 1985;OnSoffshore cott et al.,disposal 1987).mounds The species and athasadjaa cvery ent

control sites, M. ambiseta has been found to exhibit distinct seasonal patterns with peak densities occurring

i inApril the and periodMay, from1983, December the population throughofMarch benthic(Scinfauna ott et alon., 1the 987). disposal Fllowing mounddisposal was reduced of dredged to near material, zero. Byin

December, 1983, however, normal patterns of population growth were apparent, although there were fewer

I specimens (Sanders etpalrese ., 1980) nt on the or mesocosm disposal mexperiments ound itself. The (Grassle se resuandlts dGrassle, iffer from1985) thosewhere in shallow, M, ambiseta nearshore populations habitats

undergo rapid and explosive growth between June and September, with reproduction declining in the fall.

i The that the offshore offshoreresul populations ts from themight disposal represe mounnt dsa different thus indicspecies ate a fall-winter (i.e., possiblypattern M. californiensis in reproduction, orMsuggesting .fragilis).

It will be necessary to examine specimens from these various localities to confirm or reject this hypothesis.

I indicator"In estuarine because ofenvironments, its dominanceCainpitelIa heavilycapitata stressedwas and one of organically the first species enrichedto besediments designated(Reish, a"pollu1955, tion

1957; Gmssle and Grassle, 1974; Pearson and Rosenberg, 1978). A direct relation of C. capitata with

I poll fromutiplankton on, howeappears ver, is dotubebtfuthel. Orga pimary nic enrichme cause forntsuocfcess sediments of C. capitata from waste . Whendispoorganic sal or content natural fluof xethes

sediments is high, the worms appear to reproduce rapidly and build up large populations (Grassle and

I Grassle, natural habitat 1974). (James There isandno evGibson, idence 1980). that theHowever, species respo sublethal nds to effects toxic cowere ntaminants, demonstrated such asinmetals laboratory in its

bioassays (see below). Warren (1977) determined that high organic content was the most important factor

controlling the distribution of C. capitata. Particle size may directly affect the distribution of the worms because of its relationship to organic carbon.

I

i I 49
i

!1

Capitellids are considered to be opportunistic recruits to disturbed or otherwise stressed sediments I

that have been defaunated. One of the best documented examples of faunal succession following an

environmental perturbation is that of the West Falmouth Oil Spill (Sanders et al., 1980). Following the spill, i

most of the benthic infauna was killed. In order of most opportunistic, the following species recolonized the ii

site: 1) Capitella capitata, 2) Polydora cornuta (=P. ligni), 3) Syllides verrilli, 4) Micropthalmus aberrans,

5) StrebIospio benedicti, and 6) Mediomastus ambiseta. The predominance of C. capitata in this sequence []

follows a typical pattern whereby capitellids and spionids build large populations following disturbance. A |

similar result has been obtained by Zajac and Whitlatch (1982a, b) in a Connecticut estuary. These patterns

are also well documented on disposal mounds (Scott et al., 1987) where fleshly deposited sediment provides _lm

an open site for recolonization. !1

Feeding and Burrowing. Capitellids build tubes or burrows at or near the surface of soft sediments.

Tubes are either horizontal or vertical, often extending 15cm in depth (Fauchald and Jumars, 1979). Capitellids I_

frequently live in black, anoxic muds, and the tubes permit the worms to maintain contact with the overlying B

water and obtain necessary oxygen through their burrow irrigation activities. Capitellids are non-selective

deposit feeders that feed by everting a papillose, saclike pharynx. Very little work, however, has been dR

conducted on the feeding activities of even the most common species. Fauchald and Jumars (1979) postulate []

that Capitella capitata, a subsurface deposit feeder capable of considerable mobility, may be relatively nonselective in its ingestion of sediment, whereas less opportunistic species of the genera Notomastus and []

Dasybranchus may be more selective. Information on its feeding may be found in Fauchald and Jumars !1

(1979).

Laboratory studies have documented that small individuals of Capitella can attain daily weight- j

specific growth rates as high as 21% (Tenore and Chesney, 1985). Because detrital components of sediments

are included in their diets, capitellids have become convenient experimental animals to biologists interested

in the role of detritus in feeding ecology, physiology, and biochemistry of the deposit feeders (Tenore, i

1977a, b; 1981; 1983; Phillips and Tenore, 1984). []

Sibling Species of Capitella spp. The taxonomic concept of CapiteUa capitata has changed

considerably since the discovery that genetically distinct, but morphologically similar sibling species actually U

comprised this "cosmopolitan" species (Grassle and Grassle, 1976; Grassle, 1980). At present, at least 11 m

sibling species have been separated from Capitella capitata sensu lato. Grassle and Grassle (1976) found

subtle morphological differences between 6 different siblings as well as differences in reproduction, larval

development, and in electrophoretic mobilities of alleles at selected enzyme loci. These latter genetic am

differences indicate great genetic distances between species. More recent ultrastructuraI studies of sperm

morphology and setae have further demonstrated the specific differences (Eckelbarger and Grassle, 1987a) !1

between the siblings as have studies of chromosomes (Grassle et al., 1987). Table 4.1 provides details of II

morphological and reproductive information on the sibling species complex.

Because of the morphological similarity of the various sibling species, they cannot be reliably i

identified in routine ecological investigations. Cortsequently, many workers have minimized or ignored the i

problem of Capitella sibling species by continuing to refer to specimens as C. capitata without qualification

(Warren, 1976a, Pearson and Rosenberg, 1978; Rhoads et al., 1978; Ewing and Dauer, 1981; Boesch, 1982; i

Tsutsumi and Kukuchl, 1983, 1984; Chapman and Fink, 1984). Other workers have recognized the problem, 31

but have been forced to cite Capitella spp. or CapiteUa capitata-complex when sorting field specimens

(Hannon, 1981; Zajac and Whitlatch, 1982a, b; Foumier and Levings, 1982). In contrast, those biologists i

conducting laboratory studies of living organisms have been able to use the numerical system of species II

designations established by J.P. Grassle (Tenore, 1981; Bhup and Marsden, 1982; Phillips and Tenore,

1984; Forbes and Lopez, 1987). I

!1 The concept of C. capitata as a pollution indicator needs to be revised following the results of these

studies because recruitment patterns and tolerance levels for the individual sibling species have not been

established. In any studies of bioaccumulation or toxicity of Capitella capitata, the identity of the individual •

sibling species will need to be determined. m

|
I

I Table 4,1. Morphology and life history traits in six sibling species of Capitelfa 2.

I

I Trait Species I la H jonesi OlD i ma ORLEANS !

Prostomium Broad, Sharply Long, Broad, Broad, ?

i Peristomium Short triangular Par conical triangular triangular triangular tly fused Partly fused Partly fused Distinct, i ?

very long :

i I

I Tail: Shape Plain ! Plain Lobed Flared Lobed tj .

i . I

Dorsal cleft Absent Absent Present Present Present ?

I Hooded Hooks:

i

Formula of rows 3 (3-5) 3 (0-5) 3 (0-3) 3 (3-4) [ ? ?

i

I & teeth 2 (5) 2 (4-6) 2 (3-4) 2 (4-5) _ _ 9 9

• I " "

[

1 (5) I (5) 1 (4) 1 (4-5) [ ? ?

I

No. c¢ Genital Spines: ]

+ SD Set. 8 6.44 ± 0.88 6 6.38 ± 0.96 ? i 6 7.43 +_0.79

x± SDSeL 9 3.77±0.44 4.20-*-0.45 3.92+0.27 ? 2.16±0.41 4_+0.41

Body Weight (mg) 3-12 10.1 12.0 14 : 1.7 ?

I Diplo Chromosomes id No. 20 20 26 14 26 20

i Sex o_,Q,Po'IQ &,q_,?c:/? c_, ?,r'c_/9 Pcr/9 c_/9 c:,9,Pc¢/?

i

Egg diameter 0am) 260 x 180 75 230 50 250 ?

I

No. Eggs/brood 30-400 200-2000 30-400 200-1000 30-50 ?

I

[

I SpermNu(Iclnn) ear length 3.10 ± 0.22 5.20_+0.14 !: 4.95 ±0.15 ? 6.10_+0.24 4.62_+0.27

Acrosome length 1.38_+0.06 1.16±0.06 0.94±0.05 ? 1.35+0.04 1.15±0.04

l Middle Piece 2.60+0.25 2.93±0.45 i 2.73 ±0.15 ? 3.61_+0.21 3.60±0.19

Larval Development: !

i Mod Prototroch e / LWecellithdevel. otro. Pl Waeaktotro. ll devel. LWeecillthdeve otro.l. Planktol Poorly dreov. el. Direct ? LWeeUcithot Oevro.el.

Teiotroch

I Neurotroch Body Cilia Narrow Absent band Larg Present e patch BAbsent road band Present Small patch ? Narro Absentw band

Pygidial bands Present Present Present Absent ? Present

I '

Time planktonic Several hrs Several days 6-24 hrs < 2 wks None ?

i 2Modified from Grassle and Grassle (1976); Grassle et al., (1987); Eckelbarger and Grassle (1987a;1987b).

I 51
[]

Capitellids as Test Organisms in Bioassay and Bioaccumulation. Despite the problems with ,m

taxonomy, Capitella cap•tara has been used extensively as a test organism to determine lethal and sublethal

effects of different pollutants. Laboratory experiments addressing effects of metals and detergents on the I

larvae of Cap••ella cap•tara in France and California have revealed sublethal effects that result in abnormal il

larvae (Foret, 1972; Reish et al., 1974; Reish, 1977). The abnormal larvae were produced using the metals

chromium, copper, and zinc and detergents at concentrations below the LC-50 levels that were lethal in the iN

normal bioassay tests. These results indicate that the production of abnormal larvae may be a more sensitive II

measure of the effects of a toxicant than a 96-hr LC-50 test on adults. Adult specimens have also been used

in bioassays to evaluate increased levels of nutrients and reduced levels of dissolved oxygen and salinity a

(Reish, 1970). n

Reproduction and Larval Development. Capitellids are typically sexually dimorphic, with males

(and rarely females) of species such as CapiteUa cap•rata readily identified by the presence of heavy genital i

spines on segments 8 and 9; females typically bear genital pores in the same area. Several of the sibling II

species of Capitella exhibit hermaphroditic phases in their life history (Table 1). George (1984) provides

detailed observations on a hermaphroditic species of Cap••ella. []

A II detailed account of the development of Capitella capitata was published by Eisig (1887, 1899).

The larvae described by Eisig are derived from yolky eggs that develop in the tube of the female for the first

10-14 days followed by another 7 days as a planktotrophic larva. A good modern account of reproduction,

larval development, and culture methods for Capitella cap•tara may be found in Reish (1974, 1980). Sperm II

are transferred to the female with fertilization occurring either internally or at the time of discharge. The

female deposits its fertilized eggs around the inner surface of the tube. The larvae develop within this egg III

mass. The eggs are about 250 prn in diameter and are incubated and kept free of bacteria and other fouling II

organisms by undulations of the female which produces water movement in the tube. Reish recorded a

range of 150-250 eggs per egg mass. The eggs remain in the tube until they become trochophores at which m

time they are released, approximately 4-6 days after egg laying. Reish recorded both planktonic and direct |

forms of larvae suggesting he may have been dealing with more than one of the sibling species identified by

the Grassles. Trochophores are ovoid in shape and bear a prominent prototroch and telotroch. []

Metatrochophores have segments and rudiments of set•e; this stage settles to the bottom and forms a tube. l

Reish's worms complete their life cycle in 4-6 weeks under laboratory conditions. His larvae either

develop directly or have a brief pelagic phase followed by a rapid settlement. They appear to correspond to

the larvae of sibling species Type II of Grassle and Grasste (1976). []

Warren (1976b) studied a population of C. cap•rata in Devon (U.K.) for an entire year (January

1973 to February 1974) and assessed size distribution, reproductive condition, and spawning. She determined If.

that females with ripe oocytes were nearly always present; immature ones were present as well. Most males

contained ripe sperm. Females with fertilized eggs or larvae in masses in their tubes were found in most AL

samples taken during the year of study suggesting that at some level, reproduction was continuous in the i

population. Warren noted that some larvae were large and had no ciliary rings suggesting some direct U

development in that population.

Considerable variability has been found in the reproduction and larval development of six sibling i

species of C. cap•tara studied by J.P. Grassle (see Table 4.1). Egg diameters range from 50 p_m(C. jones•)

_m

to 250 pm (I_. The range of egg diameters is clearly correlated to the type of development: those species

with the smallest eggs have planktotrophic larvae, while those with the largest are either Iecithotrophic or IN'-"

direct in their developmental pattern. The larvae of planktotrophic species may remain pelagic for 2 or more

41

weeks, while the lecithotrophic species are pelagic for only a matter of hours (Grassle and Grassle, 1976). IBm

account of development in Capitella giardi (Mesnil) was published by Day (1936). B

An excellent

This species produces large ovoid eggs that vary from 290-370 pm across their widest diameters. The

development is direct with a non-pelagic larval stage developing that is gradually transformed into a juvenile

capitellid. Metamorphosis is marked by the disappearance of ciliary bands, i
I

t George (1984) described the larval development of a hermaphroditic species of Capitella from a

mangrove swamp in Florida. This species, later named C. caribaeorum by Warren and George (1986), is

I completely hermaphroditic, with all specimens developing genital spines. George (1984) suggests that self fertilization takes place. Large yolky eggs are laid in tubes in a mucous sleeve around the body of the adult.

The entire larval development is completed in the tube, despite the fact that metatrochophores have ciliary

] i during bands and development apparently ofarethecapable juveniles. of swimming. A completeGeorge life cycle noted(eggthattohooded egg) canhooks be completed are replacedin by6.5capillaries weeks.

Grassle and Grassle (1985) have observed spawning of Mediomastus ambiseta in the laboratory.

Females diameter have of 75 their lam areanterior releasedabdomen into thefilled water.with Mature eggsmales that are bearfree clusters in theofcoelom. sperm inSmall dorsolateral eggs having parts ofa

the anterior abdominal segments. Natural fertilization has not been observed, but Grassle and Grassle (1985)

I have in 3 days strippedat 15°C. gametesThese fromtrochophores the adults anddeveloped the resultingintofertilization metatrochophores producedinswimming 8 days. Thetrochophore first settlement larvae

occurred at day 13 with the 13-setiger stage. At 18 days early settling juveniles had a caudal filament. These

I larvae results were clarlyfound support to bethecapable field observations of producingof rapid another rproduction generationandinbuild-up 74 days ofwhen populations. held at 15° To Cdate, . These this

has been the only successful attempt at culturing M. ambiseta in the laboratory (J.P. Grassle, personal

communiWilson cation). (1933) described the larval development ofNotomastus latericeus and later some aspects of

the influence of substratum on larval metamorphosis (Wilson, 1937). Wilson artificially fertilized the eggs,

I producing into a classicwell-developed trochophore with trochophores apical tufaft,twide er twoprototroch, days. The eggs and telotroch. are about The 150 development }amin diameterof and segments developis

fairly rapid and the first three setigers have capillaries with those following having hooks. Metamorphosis

iswithmarked mud orbysandtheunderwent disappearance metamorphosis of ciliary bands. several Choice days earlier expethan riments ones demonstrated maintained in clean that larvae glass containers provided

(Wilson, 1937).

i North Carolina. BookhoutEggs(1957) wereprovided obtained afrdetailed om jelly-like account masses of theanddevelopment reared in theoflaboratory Dasybranchus to a pre-adult caducusstage. from

Three days after deposition, trochophores emerge from the egg mass. The larvae are pelagic and lecithotrophic,

i remaining settlement, pelagic the juveniles for about construct 7 days after fragile, whichmucous they seek tubesthewithbotloosely tom andattached undergo sand metamorphosis. grains. Following

Lecithotrophic larvae ofCapitella sp. I have been experimentally manipulated to alter their settlement

I Butman et al. (1988) determined that the larvae could delay metamorphosis in the absence of

response.

sediment for up to five days at 15° with no deleterious effects. In a subsequent study, Pechenik and Cerulli

(1991) postponed metamorphosis for 9 days at 20°C and determined that although survival was significantly

I reduced, there was no effect on post-settlement growth, maturity, and fecundity.

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Key to Genera of CapiteHidae from California i

1A. Thorax with 8 setigers; capillary setae entirely absent, setae hooded hooks; first 2 sefigers 3-4 times

as long as setigers 3-8; transition from thorax to abdomen abrubt ...................... Genus Amastigos 3 U

lB. Thorax with 9 or more setigers; capillary setae present .................................................................. 2

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2A. Thorax with 9 setigers, genital spines present in setigers 8-9 ................................. Genus CapiteIla

2B. Thorax with 10 or more setigers, genital spines absent .................................................................. 3 I

3A. Thorax with 10 setigers ................................................................................................................... 4 am

3B. Thorax with 11 or more setigers ..................................................................................................... 5 !1

4A. (Note 3 choices) Capillary setae present in setigers 1-4; setigers 5-10 with hooks .......................... 1

........................................................................................................................... Genus Mediomastus w

4B. (Note 3 choices) Capillary setae present in setigers 1-6; setigers 7-10 with hooks .......................... in

..................................................................................................................... Genus Neomediomastus n

4C. (Note 3 choices) Capillary setae present in setigers 1-10 ................................... Genus Decamastus

m

5A. Thorax with 11 setigers ................................................................................................................... 6 N

5B. Thorax with 12 or more setigers ..................................................................................................... 8

I

6A. Thoracic setae all capillaries .............................................................................. Genus Notomast_¢

6B. Thoracic setae capillaries and hooks ............................................................................................... 7 i

7A. (Note 3 choices) Capillary setae present in setigers 1-5; setigers 6-11 with hooks only .................. 1

.......................................................................................................................... Genus Heteromastus II

7B. (Note 3 choices) Capillary setae present in setigers 1-6; setiger 7 with mixed capillaries and hooks

or with hooks only; setigers 8-11 with hooks only ............................................... Genus Barantolla a

i

7C. (Note 3 choices) Capillary setae present in setigers 1-7; setiger 8 with capillaries in notopodia and

hooks in neuropodia; setigers 9-11 with hooks only .................................... Genus Neohetermastus

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8A. Thorax with 12 setigers ................................................................................................................... 9

8B. Thorax with 15 or more setigers ................................................................................................... 11 t

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3Amastigos acutus Piltz, 1977is known from intertidalsandy beaches in southernCalifornia. 'if

4The genus Dodecaseta McCammon and Sa93, 1978,established for a shelf species found in muddy sediments of the II southernCalifornia Bight, differs from Notomastus only in having capillary notosetae on the f'u'stabdominal

setiger. The presence of abdominal capillaries is not a generic-level character in capitellids (e.g., Mediomastus

ambiseta) andin my opinion,Dodecaseta should be referredto synonymywith Notoraastus. i

n
I

I 9A. Capillary setae present in setigers 1-12; branchiae present or absent; setiger 1 biramous or uniramous

....................................................................................................................................................... 0

I 9B. Capillary biramous ............................................................................................................................................ setae present in setiger 1-10, setigers 11-12 with hooks; branchiae present; setiger 1

Genus Dodecamastus

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10A. Abdominal notopodial hooks of posterior setigers change to acicular spines; pygidium forms anal funnel

10B. Abdominal with 2 ventralsetaelobes;all thoracic hooded setae hooks;allthoracic capillaries;setaebraall nchiae capillaries absent except

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

for hooks GenusinScyphoproctus neuropodia of5

setiger 11-12; branchiae present .......................................................................... Genus Leiochrides

I 11A. Thorax with 15 setigers .................................................................................. Genus Dasybranchus 6

I 1B. Thorax with 17-18 setigers ............................................................................... Genus Anotomastus 7

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i Descriptions of Species

A total of 15 species of Capitellidae in nine genera were encountered as part of this study, including

i one nearshore species (Heteromastusfiliformis) that was added. An additional species, Mediomastus acutus is also added because of its presence in adjacent areas and because it may yet turn up in the study area. One

new genus and species, Dodecamastus mariaensis are described. The following species are included in this

i chapter. Barantolla americana Hartman, 1963 Mediomastus Leiochrides acutus hemipodus Hartman, 1969 Hartman, 1960

Capitella capitata (Fabricius, 1780) Mediomastus ambiseta (Hartman, 1947)

I Capitella ovincola Hartman, 1947 Mediomastus californiensis Hartman, 1944

Dodecamastus mariaensis Blake, new Notomastus hemipodus Hartman, 1945

i Decamastus genus, new species Notomastus magnus Hartman, 1947 gracilis Hartman, 1963

Notomastus tenuis Moore, 1909

Heteromastusfilobranchus Berkeley and

I Berkeley, 1932 Neomediomastus glabrus Hartman, 1960 Heteromastusfiliformis (Clapar&te, 1864) Neoheteromastus lineus Hartman, 1960

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i 5A single species,ScyphoproctusocufatusReish, 1959isknown from southern Californiain shallow,nearshore sediments.

Two speciesof Dasybranchus are recorded from California:D. glabrus Moore, andD. tumbricoidesGrube. SeeHartman

(1969)for keys and descriptions.

7 A single species, Anotomastus gordiodes Moore, occurs in intertidal habitats along the California coast. The large

i numberof thoracic setigersreadily differentiatesthis speciesfrom other eapitellidsthat occur in California.
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Genus Barantolla Southern, 1921 'l_

Type species: BarantolIa sculpta Southern, 1921, by monotypy.

II

Diagnosis. Prostomium oval to conical, with terminal palpode; eyes present or absent. Thorax with

achaetous peristomium, 11 setigers; neurosetae present or absent on setiger 1. Notosetae of setigers 2-6 with

capillaries only; setiger 7 with mixed fascicles or with hooks only. Neuropodia with capillaries only through i

setigers 6-8, rest with hooks only. Branchiae present or absent, when present with 3 or more lobes. Pygidium II

without appendages.

Remarks. Barantolla is a relatively small genus. A single species, B. americana Hartman has been R

reported from California. i

BarantoUa americana Hartman, 1963 I

Figure 4.I

I

Barantolta americana Hartman, 1963:59-60; 1969:357-358, fig. 1. J

Material Examined. California, Western Santa Barbara Channel, off Point Conception, Sta. 79, 98

m (1, USNM). wj

Description. A small species, up to 30 mm long, 1.2 mm wide, with more than 100 segments; ,h

MMS specimen small, incomplete, 1.4 mm long, 0.35 mm wide, with 15 setigers. Body elongate, relatively i

smooth, with weak areolations in thoracic region. Thorax consisting of 12 segments, 11 setigerous. 16

Prostomium short, conical retracted into peristomium; eyes absent (Fig. 4.1). Peristomium about 1- W

V2times as long as following segments, without setae. First setiger reported as biramous; MMS specimen |

lacking notosetae. First 6 thoracic notopodia with capillaries only; setiger 7 with mixed capillaries and

hooded hooks; setigers 8-t 1 with hooks (Fig. 4.1). First 8 neuropodia with capillaries; hooks in setigers 9- n

1i. Hooded hooks with main fang surmounted by a row of small apical teeth; hood expanded, i

w

Biology. The species was originally reported from three offshore California canyons in depths from

260-976 m (Hartman, 1963). The present record is from 98 m. Additional specimens from deep water off BI

northern California have been found. II

Remarks. The present specimen differs from the original description in that neurosetae are not

present on setiger 1.However, this specimen is ajuvenile. A different species of Barantolla occurs in Puget I

Sound and S.E. Alaska (R.E. Ruff, personal communication). i

Distribution. Off California in shelf and slope depths; Washington (J. Orensanz, unpublished list). 11

Genus Capitella Blainville, 1828 I_

Type species: Lumbricus capitatus Fabricius, 1780. I

i

Diagnosis. Prostomium conical; eyes present or absent. Peristornium achaetous ring. Thorax with 9

segments, all setigerous; capillary setae in both rami of setigers 1-3, 1-4, 1-6, or 1-7; following thoracic l

segments lacking capillaries, with hooded hooks, or mixed fascicles; genital spines in notopodia of setigers I

8-9 in males of all species and females of some species. Abdomen with hooded hooks in both rami; capillaries

absent. Branchiae absent. Pygidium without appendages. I

Remarks. Systematics of the genus CapiteUa have become very complex in recent years owing to !1

the discovery of biological sibling species among specimens that formerly were referred to the type species,

C. capitata (Fabricius). As part of a series of papers, Grassle and Grassle (1976), Grassle et al. (1987), and t8

Eckelbarger and Grassle (t 987a, b) established morphological, reproductive, developmental, and genetic n
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"pr !

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jm Figure 4.1. Barantolla americana. Anterior endin lateral view (after Hartman, 1969).

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criteria on six sibling species (see earlier discussion; Table 4.1). All of these siblings are associated with

i organically enriched sediments and most have relatively wide distributions. At times, more than one species has been found in a single location. Because of the difficulty of recognizing these siblings in routine benthic

surveys, it has become common practice to refer to Capitella capitata as C. capitata spp. complex.

I a series ofPredating species the andbiological subspeciesstudies of C. bycapitata the Grassles and closely and others, relatedOlga species. Hartman In Hartman's recognizedscheme, and established the stem

species of C. capitata had capillaries in both rami of setigers 1-6 and mixed capillaries and hooks in setiger

i 7. Three California subspecies had different arrangements of capillaries and hooks in the thoracic region: C.

capitata tripartita Hartman, 1961b came from the San Pedro Basin and had capillaries in the rami of setigers

1-3 and mixed fascicles in setigers 4-7; C. capitata ovincola Hartman, 1947 (originally described as a full

i species) was taken from squid egg masses in shelf and canyons of central California and had capillaries in both rami of setigers I-4 and mixed fascicles in setigers 5-7; and C. capitata oculata Hartman, 1961b came

from the LaJolla Canyon and other shelf and slope locations in southern California and had capillaries in

I both mmi of setigers 1-7, no mixed fascicles, and also had eyes. CapiteiIa jonesi Hartman, 1959 from Florida has capillaries in setigers 1-3 and hooks in 4-7.
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Warren (1976a) reviewed the known species of Capitella and recognized nine species-level I

categories. Among these were updates on the status of Hartman's subspecies and the new synonymy of

Capitellides Mesnil, 1897 with Capitella. Warren regarded the structure of the hooded hooks and the setal J

arrangements on the thoracic region to be the most important taxonomic characters. According to Warren's II

scheme, C. capitata, the stem species is characterized as having capillaries on setigers 1-6 and mixed fascicles

on setiger 7. This follows the definition of Eisig (1887) and Hartman (1947; 1969) and on this basis, other II

species and subspecies are be distinguished. Warren (1976a) reestablished C. ovincola as a full species and II

also elevated C.tripartita to full species status. However, Warren synonymized C. capitata oculata with C.

capitata because the setal formula fell into the range of variability found for the stem species and the j

presence of eyes alone were not considered sufficient justification for retention of a separate species. In a |

later paper, Warren (1991) referred the genus Capitomastus Eisig to synonymy to CapiteUa. Following this

revision, Warren went on to list 14 valid species and subspecies of Capitella. In light of the discovery of li

biologically distinct sibling species in the CapiteIla capitata-complex, there is need of another review of the II

taxonomy of the genus. It appears likely that the type species of Capitetla, C. capitata (Fabricius) as defined

by Eisig (1887), Hartman (1947, 1969), and Warren (1976a) is divisible into 5-10 or more distinct species II

(see Table 4.1 for some of these). At least one of the Grassle's subspecies, HI, can be referred to C.jonesi II

Hartman, 1959, originally described from Florida (J.P. Grassle, personal communication). Reports by Pearson

and Pearson (1991) from Scotland and Wu et al. (1991) from China clearly identify different biological I

species of CapiteIla, bringing some urgency to a resolution of the taxonomic status of the Capitella capitata |

sibling species complex.

Specimens labeled as CapitelIa capitata in the MMS Phase I collections are here referred to C. /

ovincola because capillaries are present on setigers 1-4 and hooks are present on setigers 5-7. il

Capitella capitata (Fabricius, 1780) sensu lato I

Figures 4.2 and 4.3

Capitella capitata: Eisig, 1887:849, pls. 27, 32, figs. 21-23.--Fauvel, 1927:154-155, fig. 55.--Hartman, I

1947:404-405, pl. 43, figs. 1-2; 1969:361-362, figs. 1-5.--Berkeley and Berkeley, 1952:100-101,

figs. 206-208.--Reish, 1974:189-193, figs. 1-5 (adults and larvae).--Lissner et al., 1986, Appendix I 

D (in part). II

Capitella capitata capitata: Warren, 1976a: 198-201, pl. 1.

m

Material Examined. California: Tomales Bay, White Gulch, intertidal, Sta. TS-1, sample 3-3, 5 1

Jul 1962, coll. C. Ford (4, JAB); same location, Sta. TS-1, sample 4-2, 21 Jul 1966, coll. S. Dederian (3,

JAB); Walker Creek Delta, coll. 23 Jul 1969, J.A. Blake (15, JAB).---Off Point Arguello, Sta. 65, 107 m (1,

USNM). 11

Description. A large species, recorded up to 100 mm long, 2 mm wide, with 90 or more segments;

California (Tomales Bay) specimens smaller, largest 50 mm long, 1.2 mm wide, with 54 segments. Body II

elongate, thoracic region broadest, partially inflated with narrow segments; abdominal region narrower, II

segments longer. Color in alcohol light tan.

Prostomium broad, triangular, shovel-shaped with dorsal depression in Tomales Bay specimens ,J

(Fig. 4.3A, B), reported as sharply conical to broad, triangular in other accounts (Fig. 4.2A); eyes usually el

absent (Tomales Bay; Point Arguello), when present, small, paired spots. Peristomium short, inconspicuous

in Tomales Bay specimens, reported larger in other accounts (see Table 4.1). t

II

Thoracic setigers with reduced podial lobes and capillaries in both rami of sefigers 1-6; setiger 7

with capillaries or mixed fascicles of capillaries and hooded hooks; parapodia of setigers 8-9 with only

hooks; males and sometimes females with enlarged opposing notopodial genital spines on setigers 8-9 (Fig. I

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I

i

i r

enital spines

I - A D i ,E

I

Figure 4.2. CapiteUacapitata. A, anterior end,dorsalview; B, setigers 7-10 of male showing genital spines; C,

similar diagram of female showing genital pore; D, E, hooded hooks. (A-C, after Berkeley and

I Berkeley, 1952;D-E, after Hartman, 1947).

I 4.2B); spines numbering 6 or more on setiger 8, 2-4 on setiger 9. Females with large genital pore apparent

dorsally on posterior border of setiger 8 (Fig. 4.2C). Hooks reported by Hartman (1947) to have large main

i fang D, E);surmounted by 3 teeth in straight row, with individual teeth well separated from one another (Fig. 4.2 recent studies suggest numbers of teeth highly variable, with 1-3 rows and 1-5 teeth per row (Table

4.1).

I where notopodia Abdominalbecome segments elevated, with podial forminglobesdistinct becoming channel swollen, on mid- inflated, dorsumbestof developed body; setaemore hooded posteriorly hooks.

Pygidium simple collar.

I Methyl Green Staining Pattern. Tomales Bay specimens with overall turquoise color; abdominal podial lobes light against darker segmental stain; no other staining pattern apparent.

Biology. Capitella capitata is associated with organically enriched sediments and this is true for

I most of the various siblings that have been identified (J.P. Grassle, personal communication). Grassle and Grassle (1976) report up to 5 of their sibling species in a single sample.

I Table 4.1 provides a summary of reproductive, developmental, and genetic differences between the different siblings ofC. capitata thus far studied. See Biology section at the front of this chapter for additional

information about C. capitata.

, El Remarks. Capitella capitata as here defined is restricted to species having capillaries in both rami

i I of setigers I-6 and either capillaries, hooks, or mixed fascicles on setiger 7. Within this very broad

classification resides considerable variation. Five of the sibling species identified by the Grassles reside

i here (Table 4.1) and other variants have been identified (J.P. Grassle, personal communication).

I 59
I

I

r

pr

i • " -"N

• . ':i no

*.

I

:':: . ' . • ,,::

;.'. ,. /,i" !. II

=:""" "" rle '" " "

."_"• •' ;- m

• , . "_:J I

_ _ _ in i j • .- -,, .:.:!;'"

" "" ' " A B II

• " _ 500 pm

!

Figure 4.3. Capitella capitata (Tomales Bay). A, anterior end dorsal view; B, anterior end of another specimen,

lateral view. I

!

The Tomales Bay specimens agree with sibling species I (Table 4.1) in the nature of the broadshaped prostomium and short peristomium. However, final resolution on the identity of the newly discovered I

sibling species of CapiteUa capitata awaits morphological and biological studies on specimens from the |

type locality at Naples. There appear to be sufficient morphological criteria in addition to the reproductive

and genetic criteria to justify formal descriptions of all of the various siblings identified by the Grassles. II

The Point Arguello specimen has a conical prostomium that is not flattened as in the Tomales Bay I

specimen. Further, the podial lobes on the abdominal region are not well-developed. The specimen is a

female lacking genital spines with large, oval eggs (321-388 pm long × 187-212 _ma wide) in the body •

cavity. This offshore specimen is quite likely a different species from the Tomales Bay form but not described m

in detail here due to lack of suitable material.

Distribution. Stem species with wide, cosmopolitan distribution; siblings may be more restricted, i

60 |
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I Capitella ovincola Hartman, 1947

l Figure 4.4

CapiteIla ovincola Hartman, 1947:405-407, pl. 44, figs. 1-6; 1955:170.--Warren, 1976a:202-203, pl. III.

i Capitella capitata: ovinc Lissner ola: etHartman, al., 1986,1961a:125; Appendix D1969:365 (in part). -366, Notfigs. Fabricius, 1-5. 1780.

i MDescrip aterialtionEx. amBody ined.elongate, California, slender, Western up toSanta 60 mmBarbtong, ara Channel, 1.5 mm wide, Sta. 79, for9850mor(2,more SBMNH). segments;

MMS specimens small, incomplete, up to 5 mm long, 0.25 mm wide, with 23 segments. Thoracic region

d(Fistinctly ig. 4.4C).larger Colorthanin aalcohol bdomenlight ; toraci tan.c segments narrow, crowded (Fig. 4.4A); abdominal segments longer

Prostomium small, conical, slightly depressed, longer than wide, recessed within peristomium (Fig.

4.4A); eyes absent. Peristomium small, distinct ring between prostomium and setiger 1, fused with setiger 1.First 1-4(5) setigers with capillaries; setiger 5(6) with capillaries and hooded hooks; setigers (6)7-9 with

hooks; genital spines on setigers 8-9 (Fig. 4.4B, D), with 8-i0 spines on setiger 8, 6-8 on setiger 9. Hooks

i with long main fang surmounted by 4-5 apical teeth (Fig. 4.4E, F). Abdominal segments biannulate, with swollen podial lobes. Setae hooded hooks. Pygidium simple

I

[ I collar with mid-dorsal notch (Hartman, 1947).

II Biology. This species was originally reported from squid egg masses.

Remarks. Both specimens were identified as C. capitata in Lissner et aI. 1986. Capitella ovincola

i differs 6. According from C. capitata in having capillaries limited to the first 4 setigers (sometimes 5) instead of the first to Warren (1976a), the main fang of the hooks is longer in C. ovincola.

Distribution. Off California in squid egg masses and soft sediments.

I Genus Decamastus Hartman, 1963 Emended

Type Species. Decamastus gracilis Hartman, 1963, by original designation.

Diagnosis. Prostomium short, conical; eyes absent. Thorax with achaetous peristomium, 10 setigers;

i neurosetae present on setiger 1. Noto- and neurosetae of setigers 1-10 capillaries only, or last 1-2 setigers with few hooded hooks in neuropodia; abdominal setigers with hooded hooks. Branchiae absent. Pygidium

without appendages.

i gracilis. RemHowever, arks. they Specime exhibit ns variability from the Santa in the Maria presenceBasin /absence agreeofwell hoodedwithhooks the type in thespecie thoracic s, Decamastus neuropodia.

In the original description all thoracic setae were said to be capillaries and that is the case with most of the

i specimens examined here. However, a few specimens have hooded hooks in the neuropodia of the last 1-2 thoracic setigers. These specimens are clearly the same species because of the distinctive dorsal lobe on the

first few abdominal setigers and the identical methyl green staining pattern. They are small and it is likely

i that the specimens with hooks are a transitional growth phase that will eventually give way to all capillaries

in the thorax. Nevertheless, it is necessary to emend the generic definition to reflect the presence or absence

of hooded hooks in the last few neuropodia.

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Figure 4.4. Capitella ovincola. A, anterior end, lateral view (Sta. 79); B, posterior thoracic and anterior abdominal I

region, dorsal view; C, abdominal segments; D, genital spine; E, F, hooded hooks. (B-F, after Hartman,

1947). I

I

_ JJ
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I Decamastus gracilis Hartman, 1963

Figure 4.5

I Decamastusgracilis Hartman, 1963:61-62; 1969:375-376, figs. 1-2.

i Decamastus sp. B: Lissner, et al. 1986, Appendix D.

Materials Examined. Sta. 71 (1, USNM, voucher labeledDecamastus sp. B); Sta. 85 (2, SBMNH);

Sta. 86 (15, USNM) Sta. R-5 (6)

I Description. Santa Mafia Basin specimens from Stations 86 and R-4 small, about 10 mm long (2

mm for 10 thoracic setigers; 8 mm for 40 or more abdominal setigers), 0.4 mm wide across thorax. Types

recorded up to 47 mm long, 1.1 mm wide across thorax, with 50 or more segments (Hartman, 1963; 1969).

I Color in alcohol light tan.

Prostomium short, conical, inconspicuous, beating minute palpode (Fig. 4.5A); eyes absent; nuchal

I organs papillae.notThoapparent. racic segment Peristomium s narrow,noshort, t separable setigersfrom5-7 firs smooth, t setiger; with everted poorly developed proboscis wi segmental th short,furrows, smoth

setigers 8-10 with deep furrows, biannulate with posterior half glandular. Abdominal segments at least

I twice with notopodial as long aslobes thoracic; (Fig.first 4.5B). 5-6 abdominal segments with dorsal medial lobe on posterior edge in line

Thoracic parapodia inconspicuous, setae emerging from furrows; setae consisting of capillaries or

i with if present, hoodedwithhooks short,incurved last 1-2main neuropodia; fang surmounted capillariesby 8-10 numerous per fascicle, small teeth, unilimbate hood large, with inflated. clear sheath; Abdominal hooks,

parapodia with fast 5-6 setigers bearing broadly elevated dorsal flange with fascicles of 5-6 notosetae on

I left rounded and right lobesides; on dorsolateral notopodia ofsidemoreofposterior segment,segments continuinglower, downlesssides conspicuous; and ontoneuropodia ventral surface with elevated, as long,

lateral flanges beating 25 or more hooded hooks (Fig. 4.5C). Pygidium unknown.

I Methyl Green Staining Pattern. Prostomium stained; last 2-3 thoracic setigers with encircling speckled bands; abdominal segments with stain concentrating in intersegmental grooves, especially in ventrum

of far posterior setigers.

I Remarks. The elevated mid-dorsal flange bearing the notosetae on some anterior abdominal segments is diagnostic. The presence/absence of hooded hooks in the last 1-2 thoracic neuropodia may be growth

related, with hooks being eventually replaced by capillaries.

I Another, undescribed species, occurs in S.E. Alaska (R.E. Ruff, personal communcation). In this

species, the prostomium does not stain with methyl green, it lacks the dorsal notopodial flange or lobe, and

nuchal slits and eyes are present.

l Distribution. Central and southern California in shelf and slope depths; also Washington 03. Hilbig,

personal communication).

I Genus Dodeeamastus Blake, new genus

i Type Species. Dodecamastus mariaensis, new species. Diagnosis. Prostomium short, conical; eyes absent. Thorax with achaetous peristomium, 12 setigers;

neurosetae present on setiger I. Noto- and neurosetae of setigers 1-10 with capillaries only, setigers 11-12

hooded hooks in noto- and neuropodia; abdominal setigers with hooded hooks. Branchiae present. Pygidium without appendages.

i
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!

Figure 4.5. Decamastus gracilis (Sta. R-5). A, anterior end, lateral view; B, anterior abdominal segments, dorsal

view; C, middle abdominal segment, m

II

|

Remarks. Dodecamastus is related to Heteromastides, Leiochrides, Scyphoproctus and Leiochrus w

in having 12 thoracic setigers. Of these only Heteromastides has some of the thoracic setigers with capillaries

(1-3) and others with hooks (4-12). In the new genus, the first 10 setigers have capillaries whereas 11-12

have hooks. Furthermore, the new genus differs fromHeteromastides and the others listed in having distinct J

dendritic branchiae in posterior setigers.

Only a single species, Dodecamastus mariaensis is known. 1

!
!

i Dodecamastus mariaensis Blake, new species

I_ Figure 4.6

Decamastus sp. A. Lissner et al. 1986: Appendix D.

I Material Examined. California, Santa Maria Basin, off Point Estero, Sta. 5, 585 m, holotype

(USNM); off Point Arguello, Sta. 62, 582 m, paratype (USNM).

Description. Large species, holotype incomplete, with 1 large fragment and 2 smaller ones, far

i posterior segments missing, totalling segments, cm long, mm across

about 120 11 2.5 wide anterior thoracic

segments; paratype smaller, broken into 4 parts, including far posterior segments totalling about 145 segments,

8.5 cm long, 2.0 mm wide across anterior thoracic setigers. Color in alcohol light tan.

i Prostomium short, broadly rounded on anterior margin, recessed into peristomium (Fig. 4.6A);

eyes absent; nuchal organs not apparent. Peristomium consisting of single achaetous segment, clearly distinct

i from fast setiger; everted proboscis enlarged, saclike, basal part with numerous posteriorly directed papillae,

distal part smooth (Fig. 4.6A). Thoracic segments crowded, biannulate, with well-developed segmental

furrows. Abdominal segments at least twice as long as thoracic. Thoracic parapodia inconspicuous, setae

am emerging from simple grooves in furrow separating biannulations of each segment; setigers 1-10 with

|

capillaries; setigers 11-12 with hooded hooks. Abdominal parapodia swollen, with setae encompassed by

elongate flange bearing fascicles of noto- and neuropodial hooded hooks; hooks numbering 15-20 in anterior

t setigers; posteriorly, notopodial setae becoming fewer, obscured in far posterior setigers when notopodia become modified and branchiae develop; neurosetae increasing to more than 100 in long row on flange

(Fig. 4.6B). Thoracic capillaries long, thin, with narrow adhering sheath visible on both sides; hooded hooks

of thoracic and abdominal segments similar, with short, thick, pointed fang surmounted by about 3 rows of

I apical teeth, teeth closest to those furthest shortest 4.6C); hood inflated; node

fang longest, away (Fig. large,

poorly developed on shaft. Branchiae first present from middle segments, consisting of pair of arborescent,

ImM branched structures with numerous filaments arising from thick base on posterior margin of segment (Fig.

l_, 4.6B). Pygidium unknown.

Methyl Green Staining Pattern. Prostomium, peristomium, and first setiger with stain

i differentiating reticulated cuticular pattern; following 10 thoracic setigers staining uniformly, with nopattern; setiger 12 staining intense turquoise, differing from all other segments on body. Abdominal segments clearly

differentiated from thoracic setigers by distinctive staining pattern of dense speckles (glands) surrounding

setal flanges; fewer speckles (glands) scattered over rest of segment. Far posterior abdominal segments with

I branchiae not staining, but with large elongated glands densely staining in patches anterior to branchiae.

Remarks. Dodecamastus mariaensis is the only species currently assigned to the genus. Locally,

i the species may be mistaken for Decamastus because the first 10 setigers bear only capillaries. However, there are actually 12 thoracic setigers, the last two bearing only hooded hooks. The abdominal segments can

be readily identified by the development of a long, lateral setal flange. The posterior, branched branchiae

are also Locally, diagnostic, Dodecamastus and not knowmariaens n for anyis species is readilyof Decamast distinguished us. from other species by the conspicuous

methyl green staining pattern where the twelfth thoracic segment stains an intense turquoise, unlike any

I other segme D nt on the body. istribution. Known only from upper slope depths of central California, ca. 580 m.

!

i

|
!

!

!

A !

/

Figure 4,6. Dodecamastus mariaensis Blake, new species (Holotype, Sta. 62). A, anterior end, lateral view; B, J

far posterior abdominal setiger with branchiae; C, hooded hook.

i

66 !
!

I Genus Heteromastus Eisig, 1887

Type Species. Capitellafiliformb Claparbde, 1864, designated by Eisig, 1887.

I Diagnosis. Prostomium short to long, conical, sometimes tapering; eyes present or absent. Thorax

with achaetous peristomium, 11 setigers, setigers 2-6 with capillaries, setigers 7-12 with hooded hooks;

I abdominal cirrus setae hooded hooks; neurosetae present on setiger 1. Branchiae present. Pygidium with terminal .

Remarks. The genus was def'med and detail sof the hooks illustrated by Hartman (1947). Hutchings

I and Rainer (1981) selected a neotype for Capitellafiliformis, unfortunately, from a non-type locality.

One species, Heteromastusfilobranchus Berkeley, was present among collections in the Santa Mafia

Basin. A second species, H. filiformis, occurs in nearshore, estuarine habitats (Blake, 1975) and was present

materials provided by Keith H. Woodwick from northern California.

among

I Key to Species of Heteromastus from California

l 1A. Branchiae with multiple filaments, from about setiger 30-50; body robust .....................................

Ii ................................................................................................................ Heteromastus filobranchus

lB. Branchiae small, single, not with multiple filaments, from setiger 100 or more posteriorly; body

I slender, fragile ............................................................................................. Heteromastusfiliformis

Heteromastusfilobranchus Berkeley and Berkeley, 1932

i Figure 4.7

l Heteromastus Hartman, filobranchus Berkeley and Berkeley, 1932:671-674, figs. 5-7; 1952:103-104, figs. 211-212.-- 1947:428-429, pl. 53, figs. 1-4; 1961a:35; 1963:62; 1969:379-380, figs. 1-5.

I off PointMate Arguello, rial ExSta.mi62,ned582. Cal m,ifo1(USNM).--Westem rnia, Santa Mafia Basin, SantaoffBarbara Point San ChaLnnel, uis Sta Sta.. R83,444 -3,00 m,m1, 1(SBMNH). (USNM);

Description. Body long, robust anteriorly, coiled, fragile posteriorly; transition between thorax and

I abdomen obscure, but may be denoted by color and reduction in size of notosetal fascicle on setiger 12 (Fig. 4.7A). A large species, reported up to 150 mm long, 2 mm wide across thorax, with more than 250 setigers.

Color in alcohol light tan anteriorly, grey posteriorly.

I Prostomium small, conical (Fig. 4.7A); eyes absent; shallow nuchal organs present. Peristomium single, broad uniarmulate, areolated ring; everted proboscis inflated, with minute papillae. Thorax with

peristomium and first 4 setigers areolated; setigers 1-2 more or less uniannulate, following setigers 3-11

I distinctly biannulate, with deep incision from which setal fascicles emerge. Abdominal segments soft, fragile, becoming long, cylindrical with fine, transverse folds. Branchiae beginning on setiger 30-50; occurring

singly at first, then with 12-15 filaments in row (Fig. 4.7A, C), emerging just posterior to notopodium.

i Setigers 1-5 with capillaries, setigers 6-11 with hooded hooks (Fig. 4.7A). Thoracic hooded hooks with 3

• apical teeth (Fig. 4.7E, F), smooth shafts and long hoods; abdominal hooks with 3 apical teeth, distinct node

on shaft, shorter hood (Fig. 4.7G, H). Pygidium with terminal filament (Fig. 4.7D).

Methyl Green Staining Pattern. Peristomium and thoracic setigers staining uniformly green; anterior abdominal segments with speckles around podial lobes. Pygidial ring, not filament staining.

I

. 67
II

Figure 4.7. Heteromastusfitobranchus. A, anterior end,lateralview (Sta_R-3); B,far posterior branchialbearing i

abdominal segment, dorsal view; C, detail of branchiae from same; D, pygidium, dorsal view;E, F,

thoracic hooded hooks; G, H, abdominal hooded hooks.(B-D, after Berkeley andBerkeley, 1952;EH, after Hartman, 1947). II

!

68 1
I Remarks. The vouchers from Stations 62 and 83 are both large, robust specimens that agree well

with published accounts. The specimen from Station R-3 is small, probably a juvenile.

l to southernDistribution. California; Eastern shelf anPad cific slope, Sdepths. .E. Alaska (R.E. Ruff, personal communication); British Columbia

I Heteromastus filiformis (Clapar_de, 1864)

Figure 4.8

I Capitellafiliformis Clapar&le, 1864:49-50, pl. 4, fig. 10.--Eisig, 1887:841-843.

Heteromastusfiliformis: Eisig, 1887:839-840.--Fauvel, 1927:150-I52, fig. 53 a-i.---Hamnan, 1947:427-

I 428, pl. 52, figs. 1-4; 1969:377-378, figs. 1-5.--Hutchings and Rainer, 1981:373-380 (Synonymy).

Material Examined. California, Humboldt Bay, 21 June 1962, coll. K.H. Woodwick, intertidal, in

i mud, very anaerobic (10, LACM-AHF). Description. A moderate sized species, up to 100 mm long, 1 mm wide, with about I50 setigers;

California specimens 60 mm long, 0.6 mm wide, up to 160 setigers. Color in alcohol brown. Prostomium

I small, conical; absent (Fig. 4.8A); with eversible nuchal Peristomium a single achaetous ring;

eyes organs.

eversible proboscis inflated, with papillae. First 3 setigers areolated; setigers 1-2 more or less uniannulate,

following setigers 3-11 distinctly biarmulate, with deep incision from which setal fascicles emerge (Fig.

i 4.8A, B). Abdominal segments solid, robust, crowded, biannulate, somewhat cylindrical in cross section, becoming shorter in far posterior region wtih intersegmental lines greatly constricted dorsally. Branchiae

from setiger 90-120; each gill short, broad basally, tapering rounded lobes projecting posteriorly (Fig. 4.8C).

I Setigers 1-5 with capillaries, setigers 6-11 with hooded hooks. Thoracic hooded hooks with 6-7 apical teeth in transverse row (Fig. 4.8D, E); abdominal hooks with 3 apical teeth (Fig. 4.8F, G). Pygidial ring with

crenulate terminal margin and short ventral terminal filament.

i Methyl Green Staining Pattern. Prostomium and peristomium not staining; thoracic segments

ran,

staining solidly green, with those of setigers 5-9 staining most intense; abdominal segments with band of

speckles on posterior half and thin longitudinal band on either side of midventral line. Crenulate margin of

I pygidium retains stain.

Biology. A dominant species in intertidal muds subject to low oxygen conditions. The species

builds thin, delicate tubes. As part of studies of recruitment into benthic communities in Virginia, Fredette

I (1982) determined that juveniles Heteromastus filiformis initially only setigers

of had 3 thoracic with

capillaries. With growth, the fourth and fifth setigers replaced hooks with capillaries, achieving the adult

i and generic condition in 15 days in the laboratory and a width of 200 [am.

, Remarks. Heteromastusfiliformis is readily differentiated from the offshore species H.filobranchus

in having small, single-lobed branchiae arising from posterior setigers instead of multiple branched branchiae

i arising frHutchings om more ante and rio Rainer r setigers (1981). re.described this species based upon specimens from the Mediterranean.

The branchiae are small, inconspicuous, and only single-lobed. This probably accounts for why Hartman

i (1969) assDumed branchiae were absent in this species. istribution. Widespread in the Atlantic and Pacific; Australia, Victoria to Queensland;

Mediterranean.

!

!
II

70I
!

I Genus Leioch rides Augener, 1914

Type Species. Leiochrides australis Augener, 1914.

I Diagnosis. Prostomium short, conical; eyes present or absent. Thorax with 13 segments, flu'st

achaetous, 12 setigerous; setigers 1-12 with capillaries, neumsetae absent on setiger 1, with or without

hooded absent. Pygidium hooks in neuropo withoutdiappendages. a of last 1 or 2 setigers. Abdominal setae hooded hooks. Branchiae present or

Remarks. Species of the genus Leiochrides are poorly known. Four species have been referred to

I this genus: L. australis Augener, 1914 from Australia, L. africanus Augener, 1918 from west Africa, L.

paUidior (Chamberlin, 1918) from Pacific Grove, California (referred to this genus by Hartman, 1947 by a

process of elimination), and L. hemipodus Hartman, 1960 from deep basins off california_ Chamberlin's

I species should be considered incertae sedis because it was incompletely described and not illustrated. Two specimens of what at first appeared to be an undescribed species of Leiochrides were present in the MMS

Phase I collections from the Santa Maria Basin. However, the types ofL. hemipodus were examined by L.H.

i Harris of the LACM and it is apparent that the MMS specimens can be referred to that species.

i Leiochrides hemiFigure podus4.9Hartman, 1960

i Leiochrides hemipodus Hartman, 1960:136; 1969:381-382, fig. 1.

g

Leiochrides sp. Hartman, 1963:62.

I Material Examined. California, Santa Maria Basin, Sta. 28, 603 m, 2 (USNM).

Description. Material consisting of 2 specimens, both broken into numerous parts, but complete.

Both specimens long, threadlike, with more than 100 setigers; anterior fragments measuring 5.3 mm long

I from of to end of thoracic 1.0 wide; total unknown due to

tip prostomium region, mill length fragmented

nature of specimens. Type collections larger, 30-40 mm long, 1-2 mm wide, with more than 60 setigers

(Hartman, 1969). Thoracic segments wider than long, covered with light areolation; setiger 6 with band of

I glands on anterior edge, differentiating with methyl abdominal segments longer than wide. Pygidium

green;

simple, with 2 lobes. Color in alcohol light tan.

i Prostomium short, conical; eyes absent; nuchal organs not apparent (Fig. 4.9A). Peristomium achaetous, merging with setiger 1; everted proboscis with short, rounded papillae (Fig. 4.9A). Setiger 1with

notosetae only; following thoracic setigers with capillaries in notopodia; neuropodia of setigers 2-10 with

i capillaries, setigers 11-12 with hooded hooks (Fig. 4.9B). Thoracic hooks with pair of apical teeth surmounting main fang, these usually surmounted by single, unpaired tooth; additional minute teeth possibly present, but

difficult to resolve with light microscopy; hood large, inflated, with small opening through which main fang

protrudes. Anterior abdominal segments with fascicles of hooded hooks in both noto- and neuropodia;

I middle and posterior abdominal with palmately branched branchiae arising from notopodial

segments

locations (Fig. 4.9C), and setae apparently absent, leaving only neuropodial hooded hooks. Abdominal

hooks with similar tooth arrangement as those of thoracic region, but with smaller hood (Fig. 4.9D, E), more

i tightly adhering to shaft.

Methyl Green Staining Pattern. Stain imparts distinct, dense, speckled pattern on all thoracic

i setigers; setiger 6 with smooth, green band on anterior border; remainder of body not staining.
I

I

/he I

ne

1

10 ,I

J --1 O0 i_m-

'D JJ

A B |'

I

Figure 4.9. Leiochrides hemipodus (Sta. 28). A, anterior end, lateral view, prostomium and first 8 segments 1

(first 7 setigers); B, same specimen, continuation of A, showing last 5 thoracic setigers and first 2

II abdominal setigers; C, far posterior branchial bearing parapodium; D, E, abdominal hooks.

i

Remarks. Leiochrides hemipodus has previously been reported from deep basins of southern m

California (Hartman, 1960; 1969), but is more widely disu'ibtued in the region according to L.H. Harris (in li

ha.). Hartman (1960, 1969) reported that there were capillaries in all of the thoracic setigers. But, in contrast

m

to the original description, L. hemipodus actually has neuropodial hooded hooks on thoracic setigers 11-12 Ak

or 12.The present specimens have hooks in the neuropodia of setigers 11-12. The type collection is variable, 1

with two of the pamtypes having hooks in 11-12 and the holotype in 12 according to L.H. Harris who has

examined the type collection and other specimens from Dr. Hartman's collections.

Distribution. Central and southern California, in shelf and slope depths, and offshore basins in time i

silty muds.

!

!

!
!

I Genus Mediomastus Hartman, 1944

Type Species. Mediomastus californiensis Hartman, 1944, by original designation.

I Diagnosis. Body small to large. Prostomium conical, sometimes elongate, pointed; eyes present or

absent. Thorax with achaetous peristomium and 10 setigers; setigers 1-4 with capillaries in noto-and

i neuropodia; sometimes capillaries setigers 5in-10notopodia with hooded and hooded hooks inhooks bothinrami. neuropodia. Abdominal Modified setigerspaddlelike with hooded setae hooks sometimes and

present. Branchiae absent. Pygidium with midventral cirrus.

I Remarks. The genusMediomastus was established by Hartman (1944) for M. californiensis Hartman, i 1944, a common capitellid found in intertidal zones along most of the eastern Pacific from Washington to

central California. Another common California species, M. ambiseta (Hamaan, 1947), was originally referred

I to a separate genus, Capitita Hartman, because capillary notosetae were found in posterior abdominal segments. These two species and M. acutus Hartman, 1969, from continental shelf depths are common in

• California. Of these, only M. ambiseta is present in the MMS collections.

i Warren et al. (1994) revised the Mediomastus and recognized nine valid species. These

genus

authors redefined the genus, established new synonymies, artd named three new species. Despite this study,

considerable question remains about the identification of Mediomastus in American waters. Both M. ambiseta

and M. californiensis are reported from the Atlantic and Pacific coasts where they are often dominant

J

components of offshore benthic communities.

i As of this writing it is not clear whether the widespread occurrence of M. ambiseta and M.

californiensis around North America represents the same species that were originally described from

California. A careful comparison of California materials with those from the Atlantic and Gulf coasts has

! Ill yet to be conducted.

I

i

i Key to Species of Mediomastus

1A. Prostomium acutely pointed; some thoracic capillaries short, paddlelike, with smooth tips ...........

.......................................................................................................................... Mediomastus acutus

I lB. Prostomium small, conical; thoracic capillaries limbate, with distinct wings, not spatulate ......... 2

I 2A. Posterior abdominal notopodial capillaries present; thoracic hooks long, with straight shaft, no constriction, abdominal hooks shorter, with distinct shoulder and constriction on shaft; body rarely

exceeding 14 mm in length, usually smaller ................................................ Mediomastus ambiseta

i 2B. Posterior abdominal notopodial capillaries absent; thoracic and abdominal capillaries more or

less

similar, both with long, straight shaft and weakly developed shoulder and constriction; body larger,

i up to 35 mm long .................................................................................. Mediomastus californiensis

!

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!

I 73
!

II

Mediomastus acutus Hartman, 1969 I[

Figure 4.10

!

Mediomastus acutus Hartman, 1969:385, figs. 1-3.--Warren et aL, 1994:231-233, figs. le, 3a-f, 4a-f, 5a-c,

Tables 1-2. i

Material Examined. California, Monterey Bay, Watsonville, Sta. R-5, Rep. 3, 19 Oct 1992, 20 m

(20).--Los Angeles outer harbor, Sta. 16.2, rep. 5, Aug 1992, 6 m (1).

Description. A small, threadlike species with abdomen usually coiled; reported 8-9 mm long, 0.1- l

0.8 mm wide across widest part of thorax, 60 or more setigers (Hartman, 1969). Thoracic region smooth,

expanded, consisting of achaetous peristomium and 10 setigers; thorax and abdomen distinctly separated m

between setigers 10 and 11. Pygidium bearing short caudal cirrus. Color in alcohol light tan to brown.

i

Prostomium long, pointed, bearing elongated, pointed palpode (Fig. 4.10A); single pair of eyes

present or absent; nuchal organs not apparent; proboscis eversible, with few small papillae. Peristomium 1

smooth, achaetous, merging completely with prostomium; II

Thorax with setigers 1-4(5) expanded, widest at setigers 2-3, lacking prominent annulafions (Fig.

4.10A); setigers 6-10 narrower, biannulate; setigers 1-4 with broadly winged capillaries in both noto- and In

neuropodia (Fig. 4.10B); about 5-6 setae in notopodia and 3-5 in neuropodia; setigers 5-10 with hooded l

hooks in neuropodia; setigers 5-7 with bidentate hooded hooks in notopodia (Fig. 4.10D); setigers 8-10 with

modified spinelike hooded notosetae (paddlelike setae according to Warren et al., 1994) (Fig. 4.10C). Noto- t

and neuropodia spaced equidistant from one another on setigers 1-7, then notopodia shifting dorsally and II

posteriorly, with both notopodia merging across mid-dorsum forming single setal fascicle.

I

Abdomen with anterior segments rounded in cross section, posterior segments bell-shaped, with l, _

ventral groove. Abdominal setae all bidentate hooded hooks resembling those of thorax; number of hooks 8 II

from setiger 11, decreasing to 5-6 per fascicle over next 10-15 segments, reduced to 1-2 per fascicle in far

posteriorMethyl segments, i Green Staining Pattern. Prostomium and anterior region through setiger 5 not retaining

stain. Setigers 6-9 and most of setiger 10 stain intensely. Thereafter, speckles of stain retained around and

behind parapodial tori and, in larger specimens, along midventral line. i

ml

Biology. This species is known only from central and southern California in f'me silty sands in

shallow subtidal habitats. Warren et al. (1994) report a gravid female collected in Aug 1984 with oocytes 60 i

gin in diameter. 1

I

Remarks. Mediomastus acutus, with its elongated, narrow prostomiurn, should not be mistaken for

related species. In addition, the form and arrangement of the notosetae of the posterior thoracic region are /

unique. These setae have been shown to be unidentate hooded hooks that have a hood that actually wraps Ii

around the shaft forming an expanded paddlelike shape to the seta (Warren et al., 1994). Readers are referred

to that paper for elegant SEMs of the setae and depiction of the unusual manner in which the notopodia of In

the right and left sides of setigers 8-10 shift dorsally and merge along the mid-dorsal line. 1

Distribution. California, Monterey Bay to San Diego, 6-20 m.

i

i'

l
!

!

Ii

Figure 4.10. Mediomastus acutus. A, anterior end, lateral view; B, winged thoracic capillary seta; C, modified

thoracic notoseta; D, bidentate hooded hook. (A, D, after Hartman, 1969; B, C, after Warren et at.,

I 1994).

I 75
I 

Mediomastus ambiseta (Hartman, 1947)

Figure 4.11 U

Capitita ambiseta Hartman, 1947:409-410, pl. 45, figs. 1-4; 1969:369-370, figs. 1-4.--Blake and Walton, t

1977:316. II

Mediomastus ambiseta: Hartmann-Schrrder, 1962:143, fig. 18.--Warren et al., 1994:234-236, figs. 2a, 5df, 6a-f, 12, Tables 1-2.

Mediomastus sp. A: Lissner et al., 1986: Appendix D. m

Material Examined. California, Santa Maria Basin, off Point Estero, Sta. 03,291 m, (25+, SBMNH); i

off Point San Luis, Sta. R-2, (50+, USNM); off Purisima Point, Sta. 42, 100 m, 1 specimen (SBMNH); off II

Point Arguetlo, Sta. 74, 201 m, (25+, USNM).

Description. A small, threadlike species, reported up to 14 mm long, 0.1 mm wide across thorax, !

for about 70 setigers; Santa Maria Basin specimens smaller, up to 8 mm long, for about 50 setigers. Thoracic 111

region smooth, consisting of achaetous peristomium and 10 crowded setigers (Fig. 4.11A), wider than long,

last thoracic setiger may be smaller than others, transitional with first abdominal setiger; abdominal region n

with elongate, lumpy looking segments, often with ventral groove; pygidium bearing caudal cirrus (Fig. II

11B). Color in alcohol light tan to brown.

Prostomium small, conical, often drawn out to pointed palpode (Fig. 4.11A); eyes absent; nuchal l

organs not apparent. Peristomium smooth, slightly longer than setiger 1; everted proboscis inflated, with g

small, indistinct papillae (Fig. 4.11A). Thorax smooth, lacking areolations, last 3-4 setigers distinctly

biannulate. Setigers 1-4 with broadly winged capillaries in both noto- and neuropodia; about 2-6 setae in i

notopodia and 3-8 in neuropodia; setigers 5-10 with hooded hooks. Thoracic hooks tong, with weakly g

developed shoulder and no constriction. Abdominal neuropodia with hooded hooks, 5-10 per fascicle

anteriorly, 2-6 posteriorly; anterior abdominal notopodia with 3-5 hooded hooks; middle abdominal notopodial B

hooks may be long, spinelike, accompanied by single, long, unwinged capillary seta; far posterior notopodia m

with 1-2 simple capillaries lacking wings; hooks absent. Neuropodial hooks short, with prominent node and

constriction, with large main fang surmounted by 3 minute apical teeth (Fig. 4.11C-D); hood voluminous; n

spinelike hooks elongated, thinner than thoracic hooks, with minute fang. II

Methyl Green Staining Pattern. Methyl green densely staining thoracic region from setiger 2 to 9,

with setiger 9 staining most intensely; prostomium, peristomium, and setiger 1 not staining; abdominal I

segments weakly stained, usually limited to posterior border. Thoracic staining pattern with numerous speckles II

and densely staining bands on most segments, but number of dense bands variable between specimens. Jm

Biology. Available information on the mode of life and life history of Mediomastus ambiseta has R

been summarized by Grassle and Grassle (1985). This species forms thin, fragile mucous tubes that are

often covered with fecal pellets. The worm lives head-down in the tube with the caudal filament beating the

terminal anus protruding into the water. The posterior end is flexible and fecal pellets are deposited in n

w

mounds around the tube. The worms feed on the sediment near the surface and when they have ingested all

of the sediment in the immediate vicinity of the tube, they move to another location. Jim

Mediomastus ambiseta is one of the most common capitellids in benthic communities of nearshore I

waters and continental shelf and also occurs in the intertidal. This species is one of the dominant polychaetes

in the Santa Maria Basin. I

Egg diameters of at least 50 pm have been recorded (Warren et at. 1994). See Biology section in i

this chapter for additional information about the biology of M. ambiseta.

I
!

m .,....

_...,..,L_

!

| J C D _

m cirrus _:

, | __

II i

i Figure 4.11. Mediomastus ambiseta. A, anterior end, lateral view; B, posterior end, lateral view; C,D. abdominal

hooded hooks. (after Hartman, 1947).

I

I 7_
!

II

Remarks. Mediomastus ambiseta is readily recognized by its elongate, pointed prostomium, presence

of long spinelike hooks and simple capillaries that occur in the abdominal notopodia. The species is

considerably smaller than its close relative, M. californiensis.

Distribution. East, West, and Gulf coasts of North America, intertidal to shelf depths.

Mediomastus californiensis Hartman, 1944 i

Figure 4.12

t

Mediomastus californiensis Hartman, 1944:264-265, pl. 26, figs. 64-65; 1947:408, pl. 46, figs. 3-4; 1969:387- I

388, figs. 1-4.--Ewing, 1984a:14-14 to 14-16, figs. 14-9, 14-10a-c.--Warrenetal., 1994:239-241,

figs. 2b, 7c-d, 9a-d, 12, tables 1-2. i

W

Material Examined. California, Morro Bay, intertidal, 3 July 1962, Sta. Y-2, coll. K.H. Woodwick

(10). I Description. A moderately-sized species, reported up to 35 mm long, 0.5 mm wide across thorax,

with 100 or more setigers; Morro Bay specimens slender, longer, up to 55 mm long, 0.5 mrn wide, for about

130 setigers. Body with thoracic segments appearing firm, robust; abdominal segments usually coiled (Fig.

12A), more fragile in appearance. Boundary between thorax and abdomen abrupt. Pygidium with midventral I

caudal appendage. Color in alcohol light tan to colorless.

Prostomium small, with short, rounded palpode (Fig. 4.12A, B); 2 small eyes present mid-dorsally; n

d

nuchal organs not apparent; eversible proboscis with short papillae, sometimes appearing dense. Peristomium U

as long as or longer than wide; longer than anterior setigers. Thorax consisting of 10 setigers, with segments

biannulate; setigers 1-5 shorter than 6-10. Nephridiopores usually present between setigers 5-6, 7-8, 8-9, I

and 9-10, but sometimes not apparent in all locations. Setigers 1-4 with narrow winged capillaries, setigers IB

5-10 with hooded hooks. Number of setae per fascicle variable, with capillaries ranging from 5-15 in notopodia

and 5-11 in neuropodia and hooks ranging from 4-8 in notopodia and 5-10 in neuropodia. Thoracic hooks

with long shaft and no shoulder or constriction, fang short, surmounted by 3 or more teeth in multiple rows, I

hood long.

Abdominal segments multiannulated, with parapodial ridge on posterior half. Abdominal setigers !

with hooded hooks only, anteriorly 5-7 hooks per fascicle in notopodia, 8-10 in neuropodia; posterior segments

w

with number of hooks reduced to 2 in notopodia and 4 in neuropodia. Abdominal hooks similar to those of

thorax, with slight shoulder and slight constriction, but not consistent, with 3-12 apical teeth, arranged in 1- •

3 rows (Fig. 4.12C, D) (small apical teeth difficult to observe); hood shorter than in thoracic parapodia. U

Methyl Green Staining Pattern. No stain uptake on prostomium, peristomium and setigers 1-4;

setigers 5-10 staining intensely, but variable, sometimes only 5-9 staining, or sometimes 9-10 staining; |

abdomen with large speckles on some anterior setigers and dark in intersegmental areas.

Biology. The species occurs in tidal fiats at the low intertidal and in shelf depths. Warren et al. li

(1994) record a gravid female with egg diameters of about 60 p_m. I!

Remarks. Mediomastus caIiforniensis is readily distinguished from M. ambiseta, with which it

may occur, by its larger size and more robust appearance, by having a similar structure to the thoracic and

abdominal hooks instead of long thoracic hooks with weak shoulder and no constriction and short abdominal il

hooks with prominent node and constriction, and the absence of abdominal capillaries and spinelike hooks.

Distribution. Widespread in North America, intertidal to shelf depths; S.E. Alaska to southern I_

California; New England to Chesapeake Bay; Gulf of Mexico. It

I
!I| 79
!

I

Genus Notomastus Saxs, 1851 •

Type Species. Notomastus latericeus Sars, 1851 by monotypy.

I

Diagnosis. Prostomium short conical lobe, usually bearing palpode, sometimes recessed into I

peristomium; eyes present or absent. Thorax with 12 segments, first achaetous; 11 setigerous, with capillaries

only (or with hooks in last 1-3 setigers: Ewing, 1982); setiger 1 complete or lacking neurosetae. Abdominal I_

neurosetae setae hooded hooks; notosetae hooded hooks or hooks restricted in distribution or entirely absent

(Ewing, 1982). Branchiae present or absent. Pygidium without appendages.

Remarks. The definition of Notomastus as used here follows that of Ewing (1982) and thus differs t

from that of Hartman (1947) in that some species axe now known to have hooks in some posterior thoracic II

setigers and some species lack hooks in middle and posterior abdominal setigers.

Notomastus is a common genus with several interesting and important benthic species present in !

the eastern North Pacific. Hartman (1947) divided Notomastus into two subgenera: Notomastus sensu stricto m

and Notomastus (Clistomastus). These subgenera were differentiated on the distribution of nephridial apertures i

and the shape and form of the hood surrounding the tip of the hooded hooks. Eight species of Notomastus I

are recorded from California by Hamnan (1969). Three species are described here, six are included in the key. ,1D

Key to Species of Notomastus I

1A. Nephridia absent in abdominal segments, present or absent in thoracic segments; hood of abdominal I

hooks inflated, but not voluminous ................................................................................................. 4

lB. Nephridia large, conspicuous middorsally on abdominal segments; absent in thoracic segments; •

hood of abdominal hooks greatly enlarged, voluminous ................................................................ 2 II

2A. Branchiae multiple, branched, in tufts .............................................................. Notomastus magnus I

S

2B. Branchiae single, long, filamentous ................................................................................................ 3

3A. Abdominal notosetae absent ............................................................................. Notomastus lobatus s I

3B. Abdominal notosetae present ....................................................................... Notomastus latericeus 9

I

4A. Setiger 1 uniramous, with notosetae only ....................................................................................... 5

4B. Setiger 1 biramous .......................................................................................... Notomastus linea_ 1° j

|

!

SNotomastus lobatus Hartman has been reported from southern California and western Mexico (Hartman, 1947, 1969).

The species was not recorded in the present study, but has been subsequently recorded from North Carolina l

(Day, 1973) and the Gulf of Mexico 0Ewing, 1984). |

9Notomastus latericeus Sars, is the type species of the genus. The species has been reported from worldwide localities, but

has not been well documented from California. The species was not encountered during the present study but

has been identified from monitoring programs in southern California (SCAMIT, 1994). Descriptions of the •

species may be found in Hartmann-Schr6der (t971) from northern Europe and Ewing (1984) from the Gulf of | Mexico.

toNotomastus tineatus (Clapar_de) has been reported from worldwide localities. Records from North America include

Hartman (1947, 1969) from California and Ewing (1984) from the Gulf of Mexico. I

|
I 5A Anterior thoracic segments gradually decrease in width from anterior to posterior, not changing

abruptly in size; prostomium with paired groups of numerous eyespots; methyl green staining body

more or less uniformly, without striking patterns ................................................ Notomastus tenuis

I 5B. First 4-5 thoracic setigers expanded, followed abruptly by narrower with

segments; only a

single

pair of minute eyespots; methyl green producing distinct patterns especially in abdominal segments

where a pair of longitudinal midventral bands extend to posterior end ....... Notomastus hemipodus

I

Notom_ms hemipodus _ma, 1945

I Figure 4.13

Notomasms (Clistomastus) hemipodus Hartman, 1945:38; 1947:424, pl. 48, figs. 1-5; 1951:103, pl. 24, figs.

i 1-3; 1969:393, figs. 1-5.

Notomastus hemipodus: Day, 1973:100.--Ewing, 1984a:14-28 to 14-31, fig. 14-24.

I Notomastus tenuis: Lissner et al., 1986: Appendix D.

Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. R-5,154 m (4, USNM);

off Point Conception, Sta. 84, 393 m (2, SBMNH).--Western Santa Barbara Channel, Sta. 86, 197 m (3,

I USNM).

Description. A relatively small species, reported up to 30 mm long, 1.5 mm wide for about 50

i segments. Thorax generally Santa Mafi smooth, a Basin weaklyspeareolated cimens smthrough aller, upfastto4-520 setigers mm long, and1those .0 mmsegments wide, forslightly about to50moderately segments.

expanded (Fig. 4.13A), appearing lighter in color than rest of thorax. Change from thorax to abdomen

I marked mostly smooth by changeexcept from for all capillaries parapodialtoriall dgeshooded meetinghooksacross inbotdorsum. h noto- andColor neuropodia. in alcoholAbdlight omitannal tosegmen brown,ts

with anterior expanded thoracic segments lighter in color than rest of body.

I with 2 minute Prostomi darkumeyespots elongate,andtriangular, pair of nuchal somewha slitst dalong epressed, postebea riortingmargin small(Fig. apica4.13B), l palpodthese e (Fig. difficult 4.13B)to;

see unless peristomial fold pushed back. Everted proboscis densely papillated proximally, smooth distally

i (Fig. everted4.13A). Peristomium achaetous, about one-third length of first setiger, hidden by folds of proboscis if .

Setiger 1 with notosetae only; setigers 2-11 biramous, with widely separated podia; with lateral

I organs between podia, but slightly closer to notopodia. Setigers 1-11 with simple parapodia bearing capillary setae. Last 5-6 thoracic setigers with lateral grooves bearing nephridial apertures located in intersegmental

grooves only observed when grooves pushed apart.

i Abdominal with parapodia well separated (Fig. 4.13A, C), tori with hooded hooks in

segments

single rows; notopodia shifted to more dorsal position so that glandular ridges lie adjacent to one another.

Neuropodia with longer glandular ridges and setal toil; dorsalmost edge of neuropodia elevated into small

I papilla in anterior abdominal segments. Hooks with curved main fang surmounted by curved row of 5-6

apical teeth (Fig. 4.13D, E); shaft long, straight, with thick node near middle. Branchiae absent. Pygidium

simple, without appendages.

I Methyl Staining Body staining heavily and with distinctive patterns. Peristomium

Green Pattern.

and thoracic segments staining more or less uniformly, concentrated in numerous minute reticulations (?cells)

especially in setigers 6-11, anterior expanded region staining lighter than rest of thorax; with distinctive

I triangular or wedge-shaped dark stained patch on dorsum of anterior half of setiger 7. Change to abdomen

abrupt, with abdominal segments exhibiting distinct stained and non-stained areas and forming patterns;

dorsal stain very dark, bluish, extending laterally, surrounding neuropodium, but leaving clear areas around

I and between neuropodia and that of next segment; ventrum with pair of longitudinal stained bands running

across segments and extending length of body.

I 81
I

I

_°:_ o,o_ I

I

... B _":_ _(_ I

'"_ ""''7"' 17...... "'/'II- '''£'_I' ."i.-_'''I'";... !"{"_"''':'' ''''!''I" -£.-'"'j_'_ r[]//!i_://_/! ' .-.:_--(_._!...>.l _.....7 iii71i,

i,!U;

__._ (i,.I'i1!,!77,7>1 I

......... z--..,---., ,..--........ ::_::- i,,-'/i,.,',_.'i '.!

.-.:....._t::-i-_ ._ : :::.::::-'- IIk'i,LI I

...... ._- -I_iiiii :-_-__:-:_< -:!_':"I:L. I_' 'li_ I

'i--

C

I

A i

Figure 4.13. Notomastus hemipodus. A, anterior end, lateral view; B, prostomium and peristomium, dorsal I

view; C, middle abdominal segments, dorsal view; D, E, abdominal hooded hooks. (after Hartman,

1947).

!

I

i
I Biology. Known from a variety of sediment types including f'me sands and silts; intertidal to shelf

depths.

I from it byRemarks. the distinctNoexpansion tomastus hemipo of the fLrst dus 5-6is clthoracic osely relsegments ated to Ninstead . tenuisof, but a unmay iformbesize readitolythosedistinguished segments,

the presence of only a single pair of minute eyespots instead of two groups of numerous eyespots, and a very

I distinct 7 insteadmethyl of a relatively green staining uniformpattern stainininstead g pattern. thatVoucher includesspecimens a prominentofNwedge-shaped . tenuis from thedorsal Phasepatch II Monitoring on setiger

Program from Sta. R-5 proved to be N. hemipodus.

I Distribution. North Carolina; Gulf of Mexico; California.

i Notomastus magnus Figure 4.14Hartman, 1947

I Dasybranchus giganteus: Moore, 1909:279. Not Moore, 1906:227.

Notomastus giganteus: Berkeley and Berkeley, 1941:48-49.

I Notomastus magnus Hartman, 1947:412-415, pl. 50, figs. 1-6; 1961a:35; 1963:63; 1969:401-402, figs. 1- 6.--Reish, 1963:429.--Blake, 1975:226.

Material Examined. California, Tomales Bay, Sta. 1-59-7, coll. 29 Jun 1959, shallow subtidal in

i sand (1); Sta. 3-57-1, coll. 8 Jul 1957, in White Gulch, 9m, medium sand (1); same Jul 1964 (1); Lawsons

Flat near pier, 16 Aug 1974, intertidal in fine sands, coll. J.A. Blake (l).---Santa Maria Basin, off Point San

Luis, Sta. 23, 195 m (1, USNM).

I Description. A large, robust species, up to 30 cm long, 7 mm wide for 200+ setigers; specimen

from Lawsons landing, 17 cm long, 5.0 mm wide, with 127 setigers; specimen from Sta. 23 incomplete, but

i nearly 1cm wide across thorax. Body thickest in thoracic region, remaining thick through middle abdominal segments, narrowing slightly posteriorly. Thoracic region areolated with conspicuous lateral organs on all

segments (Fig. 4.14A). Nephridial pores absent on the thorax, but paired genital (nephridial) pores present

i from first abdominal segment continuing posteriorly (Fig. 4.14C). Color in life reddish brown with dark red branchiae; color in alcohol brown. Branchiae present from anterior abdominal segments (Fig. 4.14D);

retractile, hollow, arising posterior to notopodium, in tufts. Pygidium simple, without appendages.

I present near Prostomi posteurimorshort, margin. flattened, Evertedwith proboscis roundedwithanterior numerous palpodelarge, (Fig. smooth, 4.14B);saclike eyes alobes bsent;distally, nuchal with slits

numerous small papillae proximally (Fig. 4.14A). Thorax with achaetous peristomium and 11 setigers;

peristomium, 11 with about one-half length of setiger 1; each segment biannulate, with rings subequal. Setigers 1- simple parapodia bearing capillary setae. Abdominal segments with parapodia formed into short

notopodial lobe and long neuropodial ridge with setae extending ventrally down body in long brown line,

Ii curving to near mid-ventrum and nearly meeting parapodium of opposite side; setae long handled hooded hooks with unusual short, flattened fang encompassed by a curved ridge of numerous small teeth and

crenulations (Fig. 4.14 E, F); hood high, expanded, with broad aperture.

I grooves Methyl of areolations Green not Staistaining; ning Pattlast ern.thoracic Prostomium segment andwith thoracic bluishregion colorstai onnposterior ing uniformly half. green, Abdominal with

segments also staining green, but with light blue hue to parts of dorsum; paired nephridia staining intensely

I blue, appearing Biology.asNotomastus paired bluishmagnus moundsis common on dorsumin tidal of eachflatsabofdominal bays andsegment. estuaries in sediments containing

fine sands and coarse silts; also in shelf and upper slope depths, recorded to 900 m.

i
I

!

0r_ B I

orob0_ I

A I

_. iI

i '

!

pore I

Y,

genital pore ,si_' _-,..'_ '_ _ i

,.,,. ...... ...___{(,..i_!-'_..

E F _'-_ D n

C

I

Figure 4.14. Notomastus magnus. A, anterior end, lateral view; B, prostomium and peristomium, dorsal view; C,

anterior abdominal segments showing nephridial openings; D, far posterior segments, dorsolateral I 

view, showing branchiae; E, F. abdominal hooded hooks. (after Hartman, 1947).

!

Remarks. Notomastus magnus is one of the most readily recognized of the California species of the

genus. This large species is distinguished in having an areolated thoracic region with distinct lateral organs;

nephridial pores are apparent on paired mounds first present on the first abdominal segment. The hooded I

hooks are unusual in having a short, flattened main fang surmounted by a double caplike structure consisting

of small teeth and crenulations; the hood is high, and inflated.

Distribution. California. i

_4 |
!

I Notomastus tenuis Moore, 1909

I Figure 4.15

Notomastus tenuis Moore, 1909:277-278, pI. 9, fig. 55.

i Notomastus (Clistomastus) tenuis: Hartman, 1947:420-422, pl. 47, figs. 1-5; 1961a:35; 1966:405; 1969:397- 398, figs. 1-5.

Notomastus angulatus Chamberfin, 1919:16-17.--Berkeley, 1929:312.--McGinitie, 1935:694.

I Material Examined. California, Tomales Bay, Lawsons Flat, Sta. S-6, fine sand flats, intertidal, 19

June 1962, col/. K.H. Woodwick (3); same data, Sta. S-7 (10).--Santa Mafia Basin, off Purisima Point, 395

i m (l).--Off Descript Pointion.Concepti A longon, , slenSta. der84,spe394 cies,mwith (2).--Western specimens Santa usuallyBarba fragment ra Channel, ing uponSta. preservati 86, 197on;m (2). body

knobby posteriorly because of elevated notopodia; up to ca. 200 mm long, 0.8 ram wide, for 200 or more

I areolated setigers; Tthrough omales first Bay5specime setigers;n thorax similar,slightly but onlexpanded y 0.5 mmthrough wide. Thorax first 4-6 generally setigers,thensmoogradually th, onlytape wakly ring

(Fig. 4.15A). Color in alcohol fight tan to brown.

I each Prostomium depressed, conical, with paired groups of 15-20 small eyes on either side (Fig. 4.15B); side with shallow longitudinal groove, or groove not apparent. Everted proboscis with papillae

proximally, smooth distally. Peristornium achaetous, slightly longer than first setiger.

I Setiger 1 with notosetae only; setigers 2-11 biramous (Fig. 4.15A); with lateral organs between podia, but closer to notopodia. Setigers 1-11 with simple parapodia bearing capillary setae. Last 5-6 thoracic

setigers with lateral grooves.

I Abdominal with well with setal short in middle

segments parapodia separated; ridges segments,

becoming elongate in posterior segments (Fig. 4.15C). Setae hooded hooks, generally few in fascicle, 4-5 in

notopodia and 6-10 neuropodia of middle segments, reduced to 1-2 in notopodia and 4-5 in neuropodia of

I posterior Hooks with distal end hooked, with hood extending slightly beyond main fang; segments. only

main fang surmounted by curved row of 4-5 apical teeth in curved row (Fig. 4.15D, E). Branchiae absent.

Pygidium simple, without appendages.

I Methyl Green Staining Pattern. Prostomium staining intensely; peristomium and thoracic setigers

with patterns of speckles, most dense on setigers 5-11; abdominal segments staining more or less uniformly;

farthest posterior parapodia staining intensely.

I Biology. Notomastus tenuis is a common inhabitant of California intertidal flats in bays and estuaries.

Remarks. Notomastus tenuis is readily recognized by the paired groups of small eyespots on the

i pro distinguished stomium andfromthetherelatively sometimes smooth, co-occur non-are ringolaNte.dhemipodus appearancebyogradual f the anterior rather ththan oracicabrupt setigers. narrowing It can beof

the thorax after setiger 5 or 6.

I Distribution. Eastern Pacific, Washington to southern California; intertidal to shelf depths.

!

!

!
I

I

:_ .oer !

___, B

e$

! "'_:: ° !_

:_,_i_,_:_.._ I

:?:i!/j lii',,_J c .... ._ l

 :iiI Jli"'l l

rlr l_i\!l il

nephridial pore D '° E i

i

Figure 4.15. Notornastus tenuis. A, anterior end, lateral view; B, prostomium and anterior part of peristomium,

dorsal1947), view; C, posterior setigers, lateral view; D, E, abdominal hooded hooks. (after Hartman, I

I

_6 |
I Neoheteromastus Hartman, 1960

Type Species. Neoheteromastus lineus Hartman, 1960, by original designation.

I Diagnosis. Prostomium short conical lobe, usually bearing palpode, sometimes recessed into

peristomium; eyes present or absent. Thorax with 12 segments, first achaetous, 11 setigerous, first 7 with

I capillaries hooks only.only; Branchiae setigerabsent. 8 with capillaries Pygidium unknown. in notopodia and hooded hooks in neuropodia; setigers 8-11 with

Remarks. The genus Neoheteromastus was established by Hartman (1960) for a single species, N.

I lineus. The main characters of the genus reflect a different combination of capillaries and hooks in the 11 thoracic setigers than exhibited by Notomastus (all capillaries), Heteromastus (first 5 setigers with capillaries),

Mastobranchus (11 setigers with capillaries, 10-11 with mixed capillaries and hooks), and Barantolla (first

I 6 setigers with capillaries, following setigers with hooks only). Neoheteromastus differs from these genera in having the first 8 setigers with capillaries (setiger 7 has capillaries in the notopodium and hooks in the

neuropodium.

I Neoheteromastus iineus Hartman, 1960

.wI Figure 4.16

Neoheteromastus lineus Hartman, 1960:137; 1969:389-390, figs. 1-2.--Fauchald, 1972:243.

I Material Examined. California, Santa Mafia Basin, off Point Arguello, Sta. 56, 900 m (3, SBMNH);

Sta. 69, 927 m (1, USNM).---Gulf of California (Mexico), Guaymas Basin, Southern Trough, hydrothermal

mounds, Alvin Dive 1175, 18 Jan. 1982, 27°03.0'N, 111°23.0 ', 1997 m, box core, 2 specimens (JAB).

I Description. A small, slender species, up to 25 mm long, 1.2 mm wide, with perhaps up to 40

setigers (all specimens examined to date fragments). Color in alcohol white to light tan. Prostomium short,

i bro peristomium adly rounded and 11anteriorly; eyes absent; nuchal organs not observed. Thorax consisting of achaetous setigers; peristomium longer than following segments; everted proboscis with numerous

small papillae (Fig. 4.16). Peristomium and first setiger areolated. First setiger with notosetae and large

i lateral pore; neurosetae absent. Following 7 setigers with capillary noto- and neurosetae, with interramal pore. Setiger 8 with capillaries in notopodia and hooded hooks in neuropodia; setigers 9-11 with hooks in

both rami; interramal pores present throughout thorax. Abdominal segments with hooded hooks. Branchiae

not observed.

I Methyl Green Staining Pattern. Thoracic region staining uniformly green; setiger 5 with faint

band on anterior half, setigers 10-11 with dark speckles, abdominal segments with speckles encircling setal

i fascicles.Biology. A deep-water species, in muddy sediments.

Remarks. Neoheteromastus lineus is poorly known. In addition to the two specimens from the

I MMS collections, I examined two additional specimens from soft sediments in the Guaymas Basin collected from the DSRV Alvin as part of exploration of active hydrothermal vents. These two specimens are smaller

than those from the type locality inthe San Nicolas Basin off southern California and the two MMS specimens

I and reflect some aspects of the growth patterns exhibited by capitellids. In the original account by Hartman

(1960), fragments with 20 setigerous segments measured 12 mm in length. The thorax consisted of 11

setigerous segments, with setiger 1 having only notosetal capillaries, setigers 2-7 with capillaries only in

I both the noto- and neuropodia, and setiger 8 with capillaries in the notopodia and long handled hooded hooks in the neuropodia. From setiger 9, all remaining thoracic and abdominal setigers had hooks. Fauchald

observed this same sequence of setae in specimens from specimens from the Guaymas Basin and other

i dsetal eep-water arrangement areas oisff prese Western nt inMexico. the MMSHespecimens. did not, however, indicate the size of this specimens. This same
!

!

nephridial pore

3er

'

prob I

Figure 4.16. Neoheteromastus tineus. Anterior end,lateral view. (afterHartman, 1969). I

'.|

The Gnaymas Basin specimens are both anterior fragments that are considerably smaller than those

described by Hartman (1960; 1969) and found in the MMS collections and have a slightly different setal

pattern that reflects morphological development with growth. The smallest Guaymas Basin specimen has J

26 setigers and measures 6.7 mm long. The thoracic region consists of 11 setigers with the first six having

capillaries in both the noto- and neuropodia. Hooks begin in both rami in setiger 7. The second specimen is •

larger bodied, but there are fewer segments on the fragment. It has 18 setigers and measures 5.7 mm long. l

This fragment has notosetae only on setiger 1; capillaries in both rami of setigers 2-6; capillaries in the

notopodia of setiger 7 and hooks in the neuropodia; and hooks in both rami from setiger 8. Thus, the first •

specimen represents more of an approximation of the genera Barantolla and Neomediomastus in setal II

ontogeny, while the second specimen agrees with Neoheteromastus, but the segmental transition occurs in

setiger 7 instead of setiger 8.

There has been little effort to investigate the setal changes during growth of capitellids and this has I[

resulted in considerable confusion at the generic level. It is highly likely that many species have been placed

in the wrong genera because they had not fully developed their adult setal arrangement. II

Distribution. Central California to the Gulf of Calffomia in deep water, 900 to 2000 m.

Neomediomastus Hartman, 1969 I

Type Species. Neomediomastus glabrus (Hartman, 1960), by original designation.

I1

Diagnosis. Prostomium short, triangular, rounded anteriorly, without palpode; eyes absent. Thorax I

with 11 segments, first achaetous; setiger 1 with both noto- and neurosetae; setiger 1-6 with capillaries;

setigers 7-10 with hooks. Branchiae present in far posterior segments. Pygidium unknown, n

[] Remarks. Neomediomastus is presently known from a single species, N. glabrus (Hamnan) fl'om

deep-water off California and western Mexico. The genus was reconsidered as part of a revision of the

genus Mediomastus by Warren et al., (1994). These authors concluded thatthe setal arrangement and presence n

of "branchiae" warranted a separate genus. These authors also redescribed the species based upon the type

material. Five specimens of N. glabrus are present in the MMS Phase I collections.

Although the identity of Neomediomastus and its species, N. glabrus were retained by Warren et al. I

(1994) and are also used here, the observations reported earlier on overlap in generic characters with n

Neoheteromastus and BarantoIla (see above) suggest that Neomediomastus might eventually be found to be

nothing more than a juvenile phase of another genus and species. 1
!

i Neomediomastus glabrus (Hartman, 1960)

Figure 4.17

! Mediomastus glabrus Hartman, 1960:138.

i Neomediomastus glabrus: Hartman, 1969:391-392, fig. 1.--Fauchald, 1972:243-244.--Warren et al.,

1994:250-251.

Material Examined. California, Santa Mafia Basin, off Point Arguello, Sta. 56,900 m (5, SBMNH).

I Description. A small species, all known specimens incomplete, up to 5.5 mm long, 0.4 mm wide

across thorax, with 33 segments. Thoracic/abdominal boundary indistinct, denoted by change in segment

i shape. CProstomium olor in alcoholshort, opaque triangular, white towith lightrounded ta. tip, palpode absent; eyes absent; nuchal organs rtot

observed. Thorax consisting of achaetous peristomium and 10 setigers; setiger 1 biramous. Peristomium

I. about numerous samesmall widthpapillae as following (Fig. 4.17). setigers.Thoracic Peristomium setigersandallfirwith st setiger notosetae arolated; and neurosetae; everted prosbetigers oscis with 1-4

biannulate, about as wide as long, setigers 5-10 narrower, more elongate. Setigers 1-6 with capillaries,

! I setigers surmounted7-10bywith smallhoapical oded hoteeth; oks. hood Thoracic closelyhooks applielodng, . Abdwithominal moresegments or less straight with swoshaft; llen parapodial main fangridges. short,

Abdominal setae hooded hooks; shorter than thoracic hooks with distinct shoulder, but no constriction;

i main fangMethyl larger Green than inStaining thorax, surmPattern. ountedBody by atstaining least 2 uniformly rows of apical green;teeth; thoracic hoodsetigers closely applied. with turquoise

bands from setiger 4, developed to varying degrees through setiger 11; bands on anterior half of segments,

I those of setigers 10-11 darkest, encompassing most of segment; abdominal segments with lateral speckles. Biology. A slope species, in mud.

!

!

! p

Pr_o '

I 0

I

i Figure 4.17. Neomediomastus glabrus. Anterior end, lateral view. (after Hartman, 1969).

i

| 89
!

|

Remarks. All five of the specimens from Sta. 56 are incomplete. They are all similar in size and •

appearance and have the thorax intact and 3-4 abdominal segments. All have the first 6 thoracic segments

with capillaries. As noted in the discussion forNeoheteromastus lineus, specimens from the Guaymas Basin

exhibit overlap with Neomediomastus glabrus. Although the setal patterns in the material examined in this I!

study do not overlap in a sample, the methyl green staining patterns are nearly identical, including a distinct

band on the anterior half of setiger 5. I suspect, therefore, that the two species are the same. Final resolution •

must await study of the types of both species. |

Distribution. California, offshore basins, 900-I749 m.

!

Acknowledgments

Ii

The text of this chapter was carefully reviewed by Brigitte Hilbig and Gene Ruff. Gene provided B

numerous insights from his own observations on the Capiteltidae.

i

Literature Cited

II

Amaral, A.C.Z. 1980. Breve caracterizaga6 des g_neros da fanfflia Capitellidae Grube (Annelida, Polychaeta) g

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w

Augener, H. 1914. Polychaeta Sedentaria. Fauna Sfidwest-Australiens 5(1):1-170, 1 pl.

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Blake, J.A. 1975. Phylum Annelida: Class Polychaeta. In: Smith, R.I. and J.A. Carlton (eds.), Light's Manual,

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I Blake, J.A. and C.P. Walton. 1977. New species and records of Polychaeta from the Gulf of the Farallones,

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Foret, J.P. 1972. l_tude des effets _ long terme de quelques d&ergents (isses de la prtroleochimie) sur la

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Fredette, T.J. 1982. Evidence of ontogenetic setal changes in Heteromastus filiformis (Polychaeta:

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Grassle, J.F. and J.P. Grassle. 1974. Opportunistic life histories and genetic systems in marine benthic

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Grassle, J.P. 1980. Polychaete sibling species. In: R.O. Brinkhurst and D.G. Cook, (eds.), Aquatic Oligochaete

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Grassle, J.P., C.E. Gelfman, and S.W. Mills. 1987. Karyotypes of Capitella sibling species and of several

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Hartman O. 1944. Polychaetous annelids from California, including the descriptions of two new genera

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53, 10 pls.

I Hartman O. 1947. Polychaetous annelids Part IV. Capitellidae. Allan Hancock Pacific Expeditions 10

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I' HartmanMarine O. 1951. The littoral marine annelids of the Gulf of Mexico. Publications of the Institute of Science 2:3-58.

i HartmanPreliminary O. 1955. Quantitative survey of the benthos of the San Pedro Basin, southern California Part I. results. Allan Hancock Pacific Expeditions 19(1):1-185, pls. 1-7.

Hartman O. 1959. Capitellidae and Nereidae (Marine Armelids) from the Gulf side of Florida with a review

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Hartman O. 1960. Systematic account of some marine invertebrate animals from the deep basins of Southern

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34 pls.

I Hartman O. 1961b. A new monslrillid copepod parasitic in capitellid polychaetes in Southern California.

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I Hartman O. 1963. Submarine canyons of Southern California. Part III. Systematics: Polychaetes. Allan Hancock Pacific Expeditions 27(3):1-93, 4 figs.

,i HartmanFinal O. 1966. resultsQuantitative and conclusions. surveyAllan of thehancock benthosPacific of the Expeditions San Pedro Basin, 19(2):southern 187-456.California Part II.

Hartman, O. 1969. Atlas of the Sedentariate polychaetous annelids from California. Allan Hancock

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Hartmann-Schrfder, G. 1962. Zweiter Beitrag zur Polychaeten fauna yon Peru. Kieler Meeresforschungen

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I Hartmann-Schrrder, G. 197I. Annelida, Borstenwiirmer, Polychaeta. Die Tierwelt Deutschlands 58:1-594.

i Hutchings P.A. and S.F. Rainer. 1981. Designation of a neotype of Capitellafiliformis Claparbde, 1864, type species of the genus Heteromastus (Polychaeta: Capitellidae). Records of the Australian Museum

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I James C.J. sediments. and R. Gibson. Estuarine1980. andThe Coastal distribution Marine ofSediments the polychaete 10:671-683. Capitella capitata (Fabricius) in Dock

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Lissner, A., C. Phillips, D. Cadien, R. Smith, B. Bernstein, R. Cimberg, T. Kauwling and W. Anikouchine. •

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MacGinitie, G. 1935. Ecological aspects of a California marine estuary. American Midland Naturalist 16:629- •

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I Reish, D.J. 1974. The establishment of laboratory colonies of polychaetous annelids. Thalassia Jugoslavica

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I Reish, D.J. 1977. Effects of chromium on the life history of Capitella capitata (Annelida: Polychaeta). In:

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96 I
I 5. FAMILY AMPHARETIDAE MALMGREN, 1867

| by

Brigitte Hilbig _

I

I Introduction

Ampharetids are small to medium-sized, tubiculous worms that rarely exceed lengths of 30 mm

I (large specimens of Melinna have been reported to reach 70 ram). They are somewhat difficult to identify because almost all diagnostic characters are soft-body features; with a few exceptions, the only hard structures,

the setae, do not differ much even among genera. If the specimens are poorly preserved or incomplete,

i ide Methyl ntification Green may only be possible to generic level. To see details of the anterior segments, staining with has proven to be very helpful; staining patterns vary and are species-specific. Staining patterns

are therefore added to all descriptions in this chapter. The Ampharetidae superficially resemble the

I Terebellidae, but can be distinguished by the simple, rather than palmate or dendritically branched, branchiae arising in a straight row across the dorsum of a single segment (although there axe some exceptions, see

below), rather than in a segmental fashion; the tentacles originating from a buccal cavity into which they

I can be retracted, rather than emerging freely from the ventral side of the prostomium.

I Morphology

Most ampharetids have a wide thoracic region dearly separated from anarrowabdomen; the thorax

I is characterized by the presence of noto- and neuropodia, whereas the abdominal parapodia consist of neuropodia only, although some genera have rudimentary notopodia in all or some abdominal segments.

The thorax encompasses between 12 and 18 segments; the ventrum of all but the last few uncinigerous

I tho Because racic segments has a highly glandular body wall, and distinct glandular shields or scutes axe visible. of their very large size, the gland cells were termed "Bauchriesenzellen" (giant ventral ceils) by

Hessle (1917).

I The prostomium is usually spatulate, roughly pentagonal, and often divided into a mid-superior ' lobe and a posterior lobe surrounding the mid-superior one on three sides, so that the anterior prostomial

margin is trilobed. These lobes are not discernible when the tentacles axe fully extended. Eyes, either single

I or as rows or patches of several eyespots, are often present, but may fade in alcohol (Ampharete labrops, a

common Californian species, has a band of eyes on the ventral side of the upper lip). Nuchal organs are

sometimes located in the postectal comers of the mid-superior lobe, and some genera have raised glandular

I ridges in the posterior part of the prostomium. Underneath the prostomium is the buccal cavity, with the front of the prostomium forming the upper lip. Buccal tentacles arise from the roof of the buccal cavity; they

can be retracted into the cavity and may be smooth, grooved, or pinnate (Gnathampharete lacks tentacles

i and has jaw pieces in the buccal cavity). The buccal cavity may or may not be separated from the mouth; if present, the separating membrane is sometimes extensible.

I

I 1ENSR8. 9WaterStreet,WoodsHole,Massachusetts02543.

I
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The first two segments are reduced, often fused or telescoped, and ventrally form the lower lip •

which may be smooth or crenulate. The shape of the next four segments (3 to 6) is of great importance at the

generic level, and differences in their interpretation by different authors have caused much of the confusion •

regarding the systematics of this family. Segments 3 to 6 may be fused and form lateral raised wings that II

bear needlelike neurosetae (subfamily Melinninae), or they may be more or less free and bear a varying

number of notopodia, but never neurosetae; the first notopodia may be enlarged, pointed forward, and bear •

specialized setae, the paleae (subfamily Ampharetinae). Many authors, including Day (1964), Fauchald |

(1977), and Holthe (1986), do not count the paleal segment as the first setiger, while others do (Hessle,

1917; Hartmann-Schrtder, 1971; Gibbs and Probert, 1973). Consequently, genus and species diagnoses are •

often not consistent, especially if the term "paleae" is used only for clearly enlarged setae. To avoid any of |

these confusions and to provide consistent diagnoses and character sets for the key, all setae emerging from

segment 3 are called paleae, and they are included in the count of thoracic setigers. In most cases, the i

number of thoracic uncinigers is a more useful character than the number of thoracic setigers because the i

anterior thorax is subject to reduction and fusion, the degree of which may vary even among congeneric

i

species.

Branchiae are associated with segments 3 to maximally 6, but usually pushed forward so that they i

arise in a more or less straight transverse row on segment 3. The original segmental arrangement can still be

seen in larval stages (Zottoli, 1974). Mexamage and Phyllampharete retain this arrangement as adults; in •

many other genera, the branchial bases can be seen as ridges on the dorsal body wall, extending backwards II

to the parapodium above which they arose originally. Two, three, or four pairs of branchiae may be present

(the number of branchiae for Uschakovius Laubier, 1973 is unknown, and may range from none to two). In •

the Melinninae, a pair of dorsal recurved hooks may arise from segment 3, and segment 6 is characterized |

by a dorsal membrane. In some genera, one or two papillae can be seen between the left and right branchiae.

These papillae are the openings of the first pair of nephridia. Figure 5.1 shows configurations of the anterior am

ends of genera and species treated in this chapter, indicating distributions of branchiae, noto- and neuropodia, |

and specialized setae among the first seven segments.

Ampharetids have two to five pairs of segmentally arranged nephridia, the anteriormost of which •

usually originates, and often also opens to the outside, in segment 5. This segment is the last of the anterior I

thorax; it is separated from segment 6, the first segment of the posterior thorax, by a diaphragm, the only

inner intersegmental wall (dissepiment) left in the thorax of ampharetids. Hessle (1917) studied the •

arrangement of the nephridia in great detail and based his keys and generic definitions mostly on this l

character. Because it is impractical for taxonomists to identify specimens using a character only to be seen

after tedious dissection or in series of thin sections, Hessle's work has been largely ignored; however, his •

introduction to the terebellomorph polychaetes is one of the most insightful written to date and should be |

consulted by any taxonomist new to ampharetids or any of the other terebellomorph families.

The first fully developed, biramous parapodia are always found in segment 7; notopodia are generally •

simple and cylindrical, but may have small pre- or postsetal lobes and ventral cirri. Neuropodia are more or l

less elongate pinnules with single rows of often very numerous uncini; sometimes dorsal cirri are present.

The number of uncinigerous thoracic segments is one of the most important diagnostic characters for •

ampharetid genera, and ranges from 9 (genus Mugga) to 14. The notopodia on one of the posterior segments l

may be raised, bear modified setae, or form a valvelike structure across the dorsum.

Aside from modified notosetae that usually have broadened, hirsute tips (for example, Anobothrus, •

Mugga, and Sosane) and the paleae, all ampharetid setae look basically alike; they are limbate in all biramous

parapodia and sometimes simple capillaries in the anterior thorax. The uncini, in contrast, can have different

shapes, both along the same animal and among species, although they are not often used for species •

discrimination. An elaborate review of the morphology of ampharetid and other terebellomorph uncini can

be found in Holthe (1986a). In somewhat simpler terms, each uncinus consists of several teeth of various

sizes that can be arranged in one, two parallel, or more, usually irregular rows; an additional small tooth i

98 l
1

1

Amage nn n n Paramage nn n n

I Mexamage

n n

I _,I,I•]_1_]_

Ampharete acutifror_

I labrops Ampharete finmarchica _nnn

l __' "_"°°,11".['_ I J F ,,_'I_t'l'_

Anobothrus gracilZs Anobo_hruzpaleazus

I ' i '7i

Lys_Te, Paralysippe

Pseudampharete, Sosane $osanopsis

|

I '

Amphicreis Amph_samytha

ildtlnii ; /!11/In

I _ I I l'l'l'} _, ) i ] I't'I'I',

Asabellides colifornica

l cornuta Asabellides iineata

t Sll _ i'1", I I I _ I_l,l", i II• J I,

1

Eclys_pe, Mugga Glyphanostomwn, Samytha

_lnna na_

l ) ![ l_l_l,I,_) ,iI I_,I_I,"'],

I Melinna heterodonta Melinna oculata lAn /IA/l# i ,A ,, ,

i I ! I

, ..,_-._.-,, , _,_., _.,

I Figure 5.1. Schematic lateral views of anterior ends of genera and species treated in this chapter, indicating

distribution of branchiae, noto- and neuropodia, and specialized setae among segments 1 through 7.

I
!

II

close to the base of the uncinus, called rostral point, that serves as an attachment point for a ligament; and a •

rounded, often quite large structure at the base of the uncinus, the basal prow. An additional ligament is

attached to the posterior "comer" of the uncinus; in large, spreading abdominal neuropodia, these ligaments •

can be seen through the body wall, resembling fascicles of embedded setae. II

The abdomen may consist of a few (about 10 to 15) to numerous (up to 70) segments; a long

abdomen is characteristic for the Melinninae. The neuropodial pinnules are usually narrower and longer I

than the thoracic ones, and the last pair may project beyond the pygidium. The anus is terminal, often

surrounded by two lateral cirri or flaps and rounded lobes or papillae.

The tubes are often considerably longer than the animals, and they usually consist of a fragile inner I

mucus wall and an outer layer of sediment particles which can be very thick, especially in very fine sediments.

The mucus for the tubes is produced in the large glands constituting the ventral shields.

!

Taxonomic History

! The first ampharetids were described by Sars in 1835 (Amphicteis gunneri and Sabellides octocirrata),

and a few additional species were described by Sars, Mtiller, and Grube before Malmgren established the mm

family Ampharetidae in 1866. Most of the shallow-water species were known by the early 1900's, the most

prolific authors being Grube, McIntosh, Moore, Chamberlin, and Caullery. Since about 1950, a wealth of

species and genera has been discovered mostly in Arctic and Antarctic waters and the deep sea, mainly by

Hartman and Fauchald.

The systematics of this family are still unclear; many genera are monotypic, little is known about

variability of characters used for the diagnosis of species and genera, and almost all characters are meristic.

While such characters are easy to use for the taxonomist, it is difficult to recognize the systematically II

meaningful ones as long as the variability is not known. If generic diagnoses are too narrow, potentially

every specimen with a slight difference in one character becomes a new genus. Day (1964) was one of the =.

first to address this problem, and his work is a stepping stone for establishing a more natural system of the

Ampharetidae. Two very comprehensive reviews of the ampharetids, together with the other terebellomorph

families, were provided by Hessle (1917) and Holthe (1986a), but the evolution of the terebellomorphs in

general and the ampharetids in particular is still not well understood. Both authors recognized groups of

more closely related genera within the family, and Holthe (1986a) proposed several tribes, but was only

able to group some of the many genera into those tribes. A comprehensive faunistic account of the ampharetids

of the northeastern United States was compiled by Zottoli in the late 1980's, but unfortunately was never

published. However, the manuscript has been used by taxonomists, and it is cited here as a reference for

widespread species.

,!

Distribution and Biological Notes

!

Many ampharetids have a very wide distribution geographically as well as bathymetrically, probably

because they adapt easily to new environments; for example, the most widespread hydrothermal vent m

polychaete known to date is an ampharetid, Amphisamytha galapagensis Zottoli (McHugh and Tunnicliffe,

1994), and species such as Ampharete finmarchica, Glyphanostomura pallescens, Mugga wahrbergi, and

Anobothrus gracilis are found in large areas both in the Atlantic and Pacific Oceans. Ampharetids are

euryhaline; one species, Hobsonia florida (Hartman) is known from estuarine waters along the U.S. east

coast, and two species have been described from fresh water in the Caspian Sea, Hypaniola kowalewskii J

Annenkova andHypania invalida (Grube). In the deep sea, ampharetids are sometimes among the numerically

!

|
!

I dominant species and constitute a major part of the deep-burrowing animals reaching several centimeters

down into the sediment. In shallower water, some species can reach densities high enough to form tube mats

I that matscanarecause apparently high sedim established entationnotratbyeslarval (Fauchald recruitment, and Jumars, but by1979). migration In disturbed of adultsenvir intoonamdepauperate ents, these area. tube

Generally, ampharetids are surface deposit feeders, feeding by stretching a short anterior part of

I their aboutbodies twice out theirofbody the tubes length.andParticles extendingarethetransported palps across alongthe thesediment, tentaclescovering toward anthearea mouth, withand a diameter certain

particle sizes may be selected during that transport process (Fauchald and Jumars, 1979). The few analyses

I of unselectively. gut contents One revealed knowndetritus, exceptionivertebrate is one of larvae, the freshwater and unicellular species, algae, Hypaniola whichkowalewskii are most likely Annenkova, taken in

that actively avoids ingestion of blue-green algae.

I A very interesting account of the feeding behavior of Amphicteis scaphobranchiata, a spectacular species that is treated in this chapter, was provided by Nowell et al. (1983). These investigators found that

the species uses its characteristic, scoop-shaped branchiae not only for respiration, but also in the feeding

I process. do, but Amphicteis scaphobranchiata does not maintain a bacterial "garden" as many other deposit feeders relies on a sufficiently high sedimentation rate in its habitat to provide it with food. The worm will

construct a burrow and mucous tube and then scoop out a depression around the tube opening, using its

I enlarged small sector pairatofabranchiae; time. To avoid sediment feedingcaptured on its inownthatfecal depression pellets, will the animal then beejects ingested, themoften out ofbythebrowsing range ofa

its tentacles with the aid of a mucus sling that has mechanical properties similar to rubber; after the fecal

I pellet bnd backwards, is pushed toward and throw the tubetheopening, pellet outtheofworm the depression. will hold theThemucus resulting strandmound betweenor the ringenlarged of pelletsbranchiae, forming

over time will, in turn, contribute to a slowing of the near-bottom currents and consequently enhance the

I settling of particles in the feeding depression. Other species move to a different location when the surrounding sediment is exploited, either by

leaving the old tube and reburrowing, or simply by lengthening the tube. This latter behavior may explain

I why ampharetid tubes usually are many times longer than the animal itself. The reproduction and larval development of ampharetids is not well known. Sexes are generally

separate, and the larvae are lecithotrophic, with a short, transitional pelagic phase. Two species of Melinna

I have been studied in detail, M. cristata by Nyholm (1950) and Hutchings (1973a-b), and M. paImata by Grehan (1991). Spawning in M. cristata takes place during a short time, during the last week of December

and the first week in January in Northumberland, United Kingdom (Hutchings, 1973a-b) and during the

I second half of January in the Gullmar Fjord, Sweden (Nyholm, 1950). Oocytes and sperm are shed through the posterior two pairs of nephridia which have more heavily ciliated walls and a wider lumen during the

spawning season than at the remaining time of the year (Hutchings, 1973b). Fertilization takes place while

I the gametes are still in the water; larvae emerge after 3 days at a length of 0.3 mm, provided with an apical ciliary tuft, a well-developed prototroch, and a weak telotroch. The pelagic phase is short, usually lasting 3

to 5 days, prolonged to maximally 14 days when no suitable substratum for settling can be found (Nyholm,

I 1950). Settling and metamorphosing larvae have 3 to 5 segments with larval setae and are about 3 mm long; they immediately burrow into the sediment, build a thin mucus tube, and then start feeding.

The adults start producing the next batch of gametes right after spawning; the gonads enter their

t proliferative phase. For about eight months, oocytes and spermatogonia (immature gametes) are released into the coelom where they mature just before spawning. During the last four months before spawning, the

gonads virtually shut down and remain in this non-proliferative phase until after the spawning season. Most

I individuals can spawn several times and live to be about three years old.

!

i I 101
!

Melinna palmata spawns between May and July (Grehan, 1991), for the first time when the animals I

are almost two years old, and probably more than once in its life time. The cycles of the gonads seem to

resemble those of M. cristata.

Brooding by the female has been described forHypaniola kowalewskii by Marinescu (1964) and for ll

Hobsoniaflorida (Hartrnan) by Zottoli (1974). Hobsoniaflorida releases eggs through nephridiopores into

the tube, where they are fertilized by sperm captured by ciliary currents. Larvae stay in the maternal tube at •

least until they are two to three setigers long (about two days), after which they settle directly to the bottom,

although juveniles up to 18 setigers long were found in the maternal tubes. The first emerging setae are

single spatulate notosetae; they are soon accompanied and replaced by capillary setae. Single crested larval •

uncini are present in all but the first (paleal) setiger in juveniles up to 13 segments long, but are shed

subsequently, and by the time the juvenile has 16 or 17 segments, the first uncinus is in setiger 4 (segment

7) as typical for adults. Tentacles are developed from the 6-setiger stage on, with a single tentacle appearing •

first; it is used for feeding from the 8-setiger stage on, replacing initial feeding by muscular contraction of I

the buccal mass and ciliary motion. Branchiae are first apparent when the animals are 11 setigers long, with

the first pair arising from segment 3 and the other three pairs emerging in segmental fashion, the last pair •

appearing when the worms are already more than 18 segments long. II

The larval development of Ampharete labrops, a species common in California, was followed to a

5-setiger crawling larval stage (J.A. Blake, unpublished). The early development resembles that of H. •

kowalevskii; gametes are shed into the seawater, and late trochophores are present after one to two days.

After three days, the larvae have grown three segments, and buds of the first branchial pair and a single

feeding tentacle are present. By the time the larvae are six days old, they have five segments and start •

feeding. After ten days, they settle and crawl on the sediment. The first pair of gills, arising from segment 3,

is fully grown. The parapodia bear single capillary and spatulate notosetae and, starting on segment 4

(setiger 2), single uncini with three teeth. 1

A recent investigation of the reproductive biology of the hydrothermal vent species Amphisamytha

galapagensis (McHugh and Tunnicliffe, 1994) showed continuous reproduction, with two spawning peaks

about two months apart; larvae are lecithotrophic and demersal. I

Key to the Ampharetidae iI

1A. Three pairs of gills (check scars if gills are broken off) ................................................................. 2

pairs of gills ............................................................................................................................ 5 '1

lB. Four

2A. Nine thoracic uncinigers; last notopodia elevated (Fig. 5.18A) ........................... Mugga wahrbergi i

2B. At least 11 thoracic uncinigers ........................................................................................................ 3

3A. Eleven thoracic uncinigers; paleae absent, first notosetae in segment 4; body slender and linear, I

prostomium scoop-shaped (Fig. 5.14B) .............................................. Glyphanostomum pallescens

3B. At least 12 thoracic uncinigers ........................................................................................................ 4 • !

i

4A. Twelve thoracic uncinigers; paleae present; first neuropodia slightly displaced dorsally, twice as i

tong as subsequent neuropodia (Fig. 5.13E) ....................................................... Eclysippe trilobata 1Ill

4B. Fourteen thoracic uncinigers; paleae absent, first notosetae on segment 4 ......... Samytha californiensis

I

102 i
!

I 5A. Segments 3 to 6 fused, forming lateral collar, bearing 3 or 4 fascicles of needlelike neumsetae;

segment 3 with dorsal nuchal hooks, segment 6 with postbranchial membrane across dorsum,

I abdomen with 40-50 segments: Genus Melinna (Fig. 5.25A-C) .................................................... 6 5B. Segments 3 to 6 free or partially fused, not forming collar; neurosetae absent before segment 7;

postbranchial membrane and nuchaI hooks absent; abdomen usually with 10-15 segments ......... 7

!

6A. Segments 4 and 5 with neurosetae only, segment 6 with neurosetae and fine notosetae; postbranchial

membrane with 13 or 14 teeth, the median 4 largest; branchiae long, unpigmented; eyes absent ...

I .......................................................................................................................... Melinna heterodonta

6B. Segments 4 and 5 with neurosetae only, segment 6 with notosetae only; postbranchial membrane

I with on prostomium about 5 coarse ......................................................................................................... teeth; branchiae short, with dark pigment bands; 2 transverse rows Melinnaof eyespots oculata

I Eleven thoracic uncinigers; paleae absent, abdominal notopodial present (Fig. 5.2D) ....... 8

7A. mdirnents

7B. At least 12 thoracic uncinigers ...................................................................................................... 10

I 8A. Branchiae clearly arising from successive segments; vestigial notopodia in segments 3 to 5, last one

with few notosetae ........................................................................................... Mexamage corrugata

I only partly arising segments; more or notopodia on segments

8B. Branchiae from successive less reduced

4and5 ............................................................................................................................................. 9

I 9A. Notopodia in segments 4 and 5 vestigial, last one with few reduced notosetae; prebranchial segments

with auricular lateral lobes ................................................................................... Paramage scutata

I 9B. Notopodia in segments 4 and 5 reduced in size, but bearing setae; body short and tumid, with relatively large prostomium and prebranchial segments, not drawn out into auricular lobes ..........

...................................................................................................................................... Amage anops

I

10A. Twelve thoracic uncinigers ........................................................................................................... 11

I 10B. At least 13 thoracic uncinigers ...................................................................................................... 21

11A. Paleae absent ................................................................................................................................. 12

I 1lB. Paleae present ................................................................................................................................ 15

I 12A. Third-to-last thoracic notopodia elevated, enlarged, with large fascicles of modified setae

.............................................................................................................................. Sosanopsis wireni

i 12B. Elevated notopodia absent; anterior abdominal neumpodia with dorsal cirri: genus Asabellides ...... 13

13A. Fourteen abdominal segments; abdominal dorsal cirri much longer than neuropodial lobes, filiform;

I median gap between branchiae narrow ............................................................... Asabellides lineata

13B. Ten or 11 abdominal segments; abdominal dorsal cirri rounded, no longer than neuropodial lobes;

I median gap between branchiae wide ............................................................................................. 14

I 103
1

I

14A. Eleven abdominal segments ......................................................................... Asabellides californica •

14B. Ten abdominal segments; comers of prostomium and fused segments 1t2 drawn out to acute, hornlike

tips ..................................................................................................................... Asabellides cornuta m

t 5A. Buccal tentacles smooth; 2 or 3 notopodia on segments 4-6; if 2, modified thoracic unciniger present m

(Fig. 5.10A, 5.24A) ....................................................................................................................... 16 l

15B. Buccal tentacles pinnate; 2 notopodia on segments 4-6; modified thoracic unciniger absent: Genus

Ampharete ..................................................................................................................................... 19 mm

16A. Posterior thoracic unciniger modified; notopodia elevated, with modified setae, and/or connected

by dorsal glandular ridge ............................................................................................................... 17 mm

16B. Modified thoracic tmciniger absent; paleae small; notopodia on segments 4, 5, and 6; 2 pairs of

branchiae smooth, other 2 pairs appearing ridged because of ciliary bands; lower lip crenulated...

................................................................................................................ Pseudampharete mexicana m

17A. Elevated notopodia on fifth-to-last or fourth-to-last thoracic unciniger, connected by glandular ridge: m

Genus Anobothrus ......................................................................................................................... 18 m

17B. Elevated, enlarged notopodia on third-to-last thoracic unciniger ...................... Sosane occidentalis !

18A. Modified notopodia and glandular ridge on unciniger 8; paleae slender, tapering gradually; notopodia

on segments 4, 5 and 6 ...................................................................................... Anobothrus gracilis m

18B. Modified notopodia and glandular ridge on unciniger 9; paleae wide, iridescent, tapering abruptly;

notopodia on segments 4/5 (fused) and 6 ......................................................... Anobothrus paleatus

I

19A. Ventral side of upper lip with band of eyespots (Fig. 5.6C); paleae short, brassy, indistinctly limbate;

segments 1 and 2 separate ................................................................................... Ampharete labrops •

19B. Lower lip without eyespots; paleae otherwise; segments 1 and 2 fused ventrally to form lower lip l

..................................................................................................................................................... ,. 20

I' 20A. Paleae slender, tapering gradually; branchiae in adults separated by distinct median gap; no dorsal

methyl green stain behind branchiae; pygidium with circlet of about 15 equally long anal cirri .... m

........................................................................................................................ Ampharete acutifrons 1

2013. Paleae wide, tapering abruptly; median gap between branchiae absent in adults; dorsum staining

with methyl green on few anterior segments; pygidium with 2 lateral anal cirri and about 6-8 low, •

rounded lobes (not cirri) .............................................................................. Ampharetefinmarchica l

21A. Thirteen thoracic uncinigers; lower lip large, scoop-shaped, crenulated ................. Lysippe labiata m

2lB. Fourteen thoracic uncinigers ......................................................................................................... 22

22A. Paleae present ................................................................................................................................ 23 n

22B. Paleae absent; prostomium without glandular ridges, mid-superior lobe quadrangular; thoracic

notopodia without ventral cirri; uncini with 2-3 rows of teeth ................. Amphisamytha biocuIata m

,04 I
!

I 23A. Prostomium with conspicuous glandular ridges and T-shaped mid-superior lobe (Fig. 5.7A); thoracic

notopodia with foliaceous ventral cirri; uncini with single row of teeth: Genus Amphicteis ....... 24

I 23B. Prostomium trilobed, mid-superior lobe simple; ventral cirri absent; uncini with more than 1row of teeth; lower lip deeply furrowed and crenulated ........................................... Paralysippe annectens

i 24A. Paleae with mucro (rounded if mucro is worn-off); lower lip crenulated, staining most deeply with

methyl green in posterior half; all branchiae cylindrical .............................. Amphicteis mucronata

l 24B. Paleae tapering to fine but continuous tip; lower lip smooth, staining most deeply with methyl green in anterior half; inner anterior pair of branchiae foliaceous, with scalloped anterior margin, others

cylindrical ........................................................................................... Amphicteis scaphobranchiata

I Description of Species

I The ampharetid fauna of California is diverse; in the Santa Maria Basin and Western Santa Barbara

Channel, 25 species assigned to 15 genera have been found. One genus and three species axe new to science.

I The following species are treated in this chapter: Amage anops (Johnson, 1901) Glyphanostomum pallescens (Threl, 1879)

Ampharete acutifrons (Grube, 1860) Lysippe labiata Malmgren, 1866

I Ampharetefinmarchica (Sars, 1864) Melinna heterodonta Moore, 1923

Ampharete labrops Hartrnan, 1961 Melinna ocuIata Hartman, 1969

I Amphicteis mucronata Moore, 1923 Mexamage corrugata Fauchald, 1972

Amphicteis scaphobranchiata Moore, 1906 Mugga wahrbergi Eliason, t955

i Amphisamytha Anobothrus gracilis bioculata (Malmgren, (Moore, 1866) 1906) Paralysippe Paramage scutata annectens (Moore, (Moore, 1923)1923)

Anobothruspaleatus Hilbig, new species Pseudampharete mexicana (Fauchald,

I AsabeUides californica ltilbig, new species 1972) new genus, new combination

AsabeUides cornuta I-Iilbig, new species Samytha californiensis Hartman, 1969

_I Asabellid1943) es Iineata (Berkeley and Berkeley, Sosanopse occide is wireni ntalisHess (Hartman, le, 19171969)

Eclysippe trilobata (Hartman, 1969)

I

i

!

I 105
!

Subfamily Ampharetinae !

Genus Amage Malmgren, 1866 i

Type Species. Amage auricuta Malmgren, 1866.

Diagnosis. Four pairs of smooth branchiae on 3 visible segments, arranged nearly segmentally with i

2 pairs on segment 3 and 1 pair each on segments 4 and 5 (2+I+1). Prostomium with nuchal organs (often

referred to as glandular ridges); buccal tentacles smooth. Paleae absent; 14thoracic setigers, 11 uncinigerous,

and 14-15 abdominal uncinigers; abdominal notopodial rudiments present, i

W Remarks. Species of Amage are usually characterized by a very short, tumid body with a wide and

square thorax and a much smaller, tapering abdomen. In comparison with the closely related generaParamage

• !

and Mexamage, this genus appears to be the most advanced in terms of cephalization in that the prebranchial

segments are the least distinct. The genus Amage consists of 11 valid species with a mostly eastern Pacific

distribution. Hartman (1969) listed five species in the Atlas of Polychaetous Annelids, of which two have i

been transferred to other genera. Only one species was present in the MMS Phase I and Phase 11material, i

Amage anops (Johnson, 1901) II

[] Figure 5.2

Sabellides anops Johnson, 1901:424-425, pl. 15, figs. 157-161, pl. 16, figs. 162-163. i

ill

Amage anops: Hartman, 1969:521-522, figs. 1-4.--Banse, 1979:1543-154, fig. la.--Lissneretal., 1986:D17.--Steinhauer and Imamura, 1990:F-1. II

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. R-2 (1); off Purisima _l

Point, Sta. 48 (1); Western Santa Barbara Channel, off Point Conception, Sta. 80 (1).

Description. Length to 27 mm, width to 5 mm; 14 thoracic setigers, 11 uncinigerous, and 14-15 i

abdominal segments. Body short, tumid, with wide thorax; abdomen quickly tapering to small pygidium an

Fig. 5.2A). Color in alcohol pale.

Prostomium with rounded mid-superior lobe bearing lateral extensions and U-shaped inferior lobe H

surrounding superior lobe and central crescentic area with nuchal organs. Segments 1 and 2 fused dorsally Jl

and laterally, separate ventrally, both forming lower lip. Branchiae on segments 3, 4, and 5, large, about as

long as 8 setigers, cylindrical, with filiform tips; posterior pair closest together, separated by median gap [I

about as wide as branchial bases (Fig. 5.2A, B). 10

Segments 3 and 4 fused laterally, bearing fast notopodia; reduced, displaced ventrally by outer i

anterior gill, bearing few very fine, short setae, easily overlooked. Subsequent notopodia gradually 1

lengthening, conical throughout, with short, triangular postsetal lobe (Fig. 5.2C); setae becoming longer

m

and thicker, reaching full size by setiger 4 (segment 7). Neuropodia with uncini first present on setiger 4 i

(segment 7), paddle-shaped; thoracic neuropodia about twice as long as abdominal ones, gradually diminishing i

in size toward pygidium, very crowded in last 5 segments or so. Notopodial rudiments large, bluntly digitiform i

and curved inward (Fig. 5.2D). Setae fully developed from setiger 4 (5 or 6 in juveniles), amber colored,

stiff, limbate, covered with numerous very regular rows of minute surficial spinules (Fig. 5.2E); thoracic i

uncini with 6 teeth in 1 row over rounded basal prow (Fig. 5.2F); abdominal uncini similar, slightly smaller

and often with 1 less tooth.

Pygidium small, about one-fifth as wide as thorax, with 2 short, clavate lateral anal cirri, i

Methyl Green Staining Pattern. Branchiae not staining except for single scattered cells, tips

completely white; lower lip and ventrum of segments 2 through 6; ventral scutes on all thoracic uncinigers; i

superior and inferior prostomial lobes and dorsum in front of branchiae. |

i
I Figure 5.2. Amage anops: A, entire animal, dorsolateral view, first notopodium concealed by branchiae; B,

anteriorend,dorsal view,schematic;C, notopodium, anteriorview;D, lastthoracicandfirst abdominal

segments, lateral view; E, detail of notoseta; F-G, thoracic uncini, lateral and frontal views. (lateral

I view from Johnson).

I Remarks. This species is easily recognized by its short, tumid body with large prostomium and

prebranchial segments, large branchiae, and the conspicuous notopodial rudiments.

I Type Locality and Type Specimens. Puget Sound (MCZ 1892)

Habitat. The examined specimens were found in sand mixed with silt.

I Distribution. Western Canada to southern California, 117 to 400 m.

i 107
!

Genus Ampharete Malmgren, 1866 •

Type Species. Amphicteis acutifrons Grube, 1860.

n

Diagnosis. Four pairs of smooth branchiae on 2 visible segments, arranged in an anterior row of 3 i

pairs, with fourth slightly behind (3+1), or in 2 rows of 2 pairs (2+2). Prostomium without glandular ridges;

buccal tentacles papillose. Paleae present; 15 thoracic set•gets including paleae, 12 uncinigerous (p+2), 12- •

17 abdominal segments; abdominal notopodial rudiments present or absent.

Remarks. The notopodial rudiments, often used for generic diagnoses, are variable within the

genus Ampharete. The consistency of a character in some genera and variability in others appears to be a i

phenomenon within the ampharetids that is not limited to the notopodial rudiments, but was also noted for

branchial configurations (e.g., Arnphisamytha, see below). The genus Ampharete contains 25 valid species

that are mostly from cold waters in northern Europe, the Arctic Ocean, and Antarctica. Hartman (1969) i

included four species in her Atlas of Polychaetous Annelids, three of which are treated in this chapter. I

Ampharete acutifrons (Grube, 1860) i

Figure 5.3

Amphicteis acutifrons Grube, 1860:109-110, pl. 5, fig. 6. n

Ampharete acutifrons: Hessle, 1917:96-97.--Hartman, 1969:537-538, figs. 1-4.--Hartmann-Schrrder, i

1971:459.--Banse, 1979:1544-1545,

fig. lb.--Zottoli, MS:15-17, figs. 3A-E. B

Material Examined. California: off Halfmoon Bay, Pioneer Canyon, Sta. C-3, 37°20.21'N,

123°08.84'W, 1400 m (1). B

i

Description. Length to 80 mm, width to t3 mm; 15 thoracic setigers, 12 uncinigerous (p+2), 13

abdominal segments. Body slender, widest anteriorly, gradually tapering toward pygidium. Color in fife I

Green, in alcohol pale. i

!

Prostomium trilobed anteriorly, with mid-superior lobe surrounded by inferior lobe; eyes, glandular

ridges, and nuchal organs absent. Segments 1 and 2 separate dorsally, very narrow, concealed by branchial •

bases; fused ventrally, produced into slightly crenulate lower tip (Fig. 5.3A, B). Segment 3 with 3 anterior l

pairs of gills and paleae; branchiae subulate, left and right group separated by narrow dorsomedian gap;

paleae long, brassy, tapering to gently curved, fine tips (Fig. 5.3C), arranged in nearly circular fascicles.

Segments 4 and 5 separate ventrally, fused laterally and dorsally, beating first notopodium and !

fourth pair of branchiae; notopodia small, setae shorter but not noticeably narrower than in subsequent

setigers. Branchial bases fused to dorsal body wall, branchiae emerging between inner and middle anterior •

pairs (Fig. 5.3A, B). Segment 6 with notopodium intermediate in size, setae as short as in preceding setiger. !1

First neuropodia on segment 7; long, narrow pinnules with dorsal extensions becoming cirri in abdominal

neuropodia. Notosetae stiff and transparent; thoracic uncini with 2 rows of 6-7 teeth above small rostral :•

point and basal prow, abdominal uncini with crest of several teeth (Fig. 5.3D, E). Anterior abdominal n

segments with tiny notopodial rudiments (Fig. 5.3F).

Pygidium short and wide, with terminal anus surrounded by circlet of about 15 equally long, slender •

cirri (Fig. 5.3G). l

Methyl Green Staining Pattern. Dorsal stain on prostomium, first 2 segments, and branchial

membrane in dorsomedian gap; ventral stain on prostomium and all segments through unciniger 10 (setiger i

12), with scutes on uncinigerous segments through unciniger 9; unciniger 10 with anterior narrow band, last

2 setigers not staining; notopodia and lateral body wall between abdominal neuropodia staining.

I

108 I
!

!

!

| 1;

4 _"

| : B

'

I| D E

, ti/ c

A .,!i;_!:....."ii_: " .."_: ::_. I

I

, G

I

Figure 5.3. Ampharete acutifrons: A, entire animal, lateral view; B, anterior end, dorsal view, schematic; C,

" I palea; D, thoracic tmcinus, lateral view; E, abdominal uncinus, frontal view; F, tast thoracic and first , abdominal segments, lateral view; G, posterior end, lateral view. (D, E, G after Zottoli).

I

Remarks. Ampharete acuU'frons is sometimes difficult to distinguish from the co-occurring A.

i I finmarchica staining pattern , particularly may be helpful if the characters; specimens the are paleae incomplete. ofA. acutifrons The shape areof the generally paleae more and the slnder methyl and green taper

more gradually than those ofA. finmarchica, andA. acutifrons does not show any dorsal methyl green stain

I behind the branchiae, whereas A. finmarchica has weak dorsal bands across 2 or 3 postbranchial segments.

!_! 109
I

|

Additionally, the branchiae of A. acutifrons are clearly separated by a median gap, whereas this gap is II

absent in A. finmarchica, at least in fully grown individuals. The Californian specimens do not reach the

size reported above; they are usually about half as large. Ill

Type Locality. Greenland. I

Habitat. In sand and mud.

Iml

Distribution. North Atlantic, Greenland to North Carolina; Mediterranean Sea; Bering Sea to Japan; 1

central to southern California; 0.5 to 1400 m.

Ampharetefinmarchica (Sars, 1864) I

Figure 5.4 I

Amphicteisfinmarchica Sars, 1864:6.

Ampharete finmarchica: Hartmann-Schrtider, 1971:458-459, fig. t 58a-h. II

Ampharete arctica: Malmgren, 1866:364, pl. 26, fig. 77.--Hessle, 1917:97-98, fig. 9.--Hartman, 1969:539- U

540, figs. 1-3.--Zottoli, MS:17-19, figs. 4A-I ( in part).--Lissner et al., 1986:D-17.--Steinhauer

I

and Imamura, 1990:F-1.

Ampharete acutifrons: Steinhauer and Imamura, 1990:F- 1.--Not Grube, 1860.

Material Examined. California: Santa Mafia Basin, off Point Sal, Sta. R-8 (1); off Purisima Point, i

Sta. R-5 (10), Sta. 42 (1), Sta. 46 (1). II

Description. Length to 45 ram, width to 7 mm; 14 thoracic setigers, 12 uncinigerous (p+2), 13

abdominal segments. Body widest in thorax behind branchiae, gradually tapering toward pygidium. Color •

in alcohol pale. ml

Prostomium roughly pentagonal, with large mid-superior lobe surrounded by inferior lobe, with Im

nuchal organs at postectal corners of mid-superior lobe (Fig. 5.4A). Segments 1 and 2 separate dorsally and l

ventrally, with segment 1 forming smooth lower lip; segment 2 very short but distinct, at least in large

specimens. Buccal tentacles slender, pinnate. Ill

Segment 3 with anterior 3 pairs of branchiae and paleae; branchiae usually not separated by 1

dorsomedian gap and bases of inner branchiae fused for short length (Fig. 5.4A, B). Paleae short, wide,

brassy, abruptly tapering to acute tips, in crowded semicircular fascicles (Fig. 5.4C). Gills short and stout, 1

subulate, barely surpassing anterior border of prostomium. Segment 4 fused to segment 3, bearing fourth |

pair of branchiae and 2 minute nephridial papillae just inside branchial bases. First 2 notopodia on segments

5 and 6, slightly raised dorsally, smaller than subsequent notopodia, notosetae of segment 5 shorter than Im

those in following segments. Setae stout and limbate, tapering to acute tip. Uncini first present on setiger 3

|

(segment 7), with double row of 5-6 teeth above rostral point and basal prow in thorax; abdominal uncini

smaller and sometimes with 3 rows of teeth, especially in posterior segments (Fig. 5.4D-F). Notopodia /

simple, cylindrical; neuropodia in thorax long and narrow, in abdomen only half as long but gradually II

becoming wider; last pair reaching edge of pygidium (Fig. 5.4G, H). Neuropodial cirri if present very short,

not projecting beyond anterior margin of neuropodial lobe, often present only as glandular upper edge of I

neuropodial lobes. Notopodial rudiments onfirst 4-8 abdominal segments, gradually decreasing and replaced 1

by low glandular ridges above neuropodial lobes.

i

Pygidium short, narrow, with 2 slender lateral cirri bearing basal pigment spot (may fade in alcohol) l

and circlet of low, rounded papillae between cirri (Fig. 5.4I). 1

!

110 !
I

i /

#

• i' rostral

,

,| r c

noRu

I G_

• .. __ aC

I ' '" .:".-- .-. -.-..L. :'. "- " ",_.. . pap

• i._.:: '....._:,, -

I H I ,,_

I Figure 5.4. Ampharetefinmarchica: A, entire animal, dorsolateral view; B, anterior end, dorsal view, schematic;

C, palea; D-F, thoracic, anterior abdominal, and posterior abdominal uncini, lateral and frontal views;

I G, pygidium. last thoracic and first abdominal segments, lateral view; H, posterior end, lateral view; I, detail of (D-F, H, I after Zottofi).

I Methyl Green Staining Pattern. Dorsal stain on prostomium and prebranchial segments; triangular

area between bases of inner anterior and posterior branchiae; transverse bands across 2 or 3 postbranchial

I segments. Ventral stain on lower lip and segment 2 except for narrow, distinct white band across middle of segment; scutes through unciniger 9, narrow anterior band across unciniger 10; no stain on last 2 thoracic

segments. All notopodia and lateral body wall between abdominal neuropodia staining.

i

i 111
!

Remarks. Among the examined material was a large specimen identified asA. acutifrons, probably I

based on the presence of well-developed papillae between the lateral anal cirri. However, these papillae are

clearly shorter than the lateral cirri, whereas the anal cirri ofA. acutifrons are all of the same length. Zottoti 1

(MS) may have described a mixture ofA.finmarchica and an undescribed species with an apparently northern !1

distribution (Ruff, pers. comm.). The undescribedAmpharete sp. has 12 rather than 13 abdominal segments,

a characteristic, scoop-shaped lower lip somewhat reminiscent of Lysippe labiata (see below), and a circlet I

of 10-15 crowded, filiform, short anal cirri in addition to the two long, lateral ones, and resembles somewhat II

A. baltica Eliason, 1955. Zottoli reported 12 abdominal segments for A. finmarchica (rather than 13) and

"10-13 smaller cirri" in addition to the lateral anal cirri, but showed six low, rounded lobes and two lateral lit

cirri in his figure 4. I

Aside from the differences in the pygidium, the two species can be distinguished as shown above

for A. acutifrons.

Type Locality. Spitsbergen.

Habitat. In silt and sand. gin

Distribution. Arctic Ocean; Greenland toDenmark, Labrador to New England; Alaska to northeastem 1

South America; Japan; intertidal to 2900 m.

I

Ampharete labrops Hartman, 1961 g

Figure 5.5 i

Ampharete labrops Hartman, 1961:127-128, pl. 34, figs. 1-4; 1969:543-544, 4 figs.--Blake, 1975:231, fig.

307.--Banse, 1979:1545.--Lissner et al., 1986:D- 17. I

I

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. 21 (1); off Point

Arguello, Sta. 64 (2); Tomales Bay, White Gulch, coll. J.A. Blake (19). 1

Description. Length to 25 mm, width to 3 mm; 15 thoracic setigers, 12 uncinigerous (p+2), 13 i

abdominal segments. Body stout, tumid, wide thorax with very prominent ventral scutes, tapering to about

half anterior width within last 2 or 3 setigers; abdomen slender, gradually tapering toward pygidium (Fig. 1

5.5A). Color in alcohol pale with brassy pateae and setae. i

Prostomium roughly rectangular, trilobed anteriorly, with mid-superior lobe surrounded by inferior

lobe (Fig. 5.5B); upper lip with row of eyespots along ventral margin, not visible in dorsal view (Fig. 5.5C). i

Segments 1 and 2 separate, not fused; dorsally very short, concealed by branchial bases on segment 3, often I!

one telescoped under the other; segment 1 ventrally produced into rounded lower lip, segment 2 visible as

very short ring with opaque white glandular band. Buccal tentacles numerous, very slender, pinnate; pinnae •

short and easily overlooked. Segment 3 with relatively few and short, indistinctly limbate, wide and stiff i

paleae, about 10 per fascicle (Fig. 5.5D), and 3 anterior pairs of branchiae. Segments 4 and 5 fused dorsally,

with separate ventral glandular shields, carrying small first pair of notopodia, paired dorsolateral nephridial i

papillae, and bases of posterior pair of branchiae fused to dorsal body wall, branchia emerging between il

inner and middle anterior pairs; dorsomedian gap very narrow. Segment 6 with second notopodia, intermediate

in size between first and third pairs. I

First neuropodia on segment 7, narrow pinnules, cirri absent. Notosetae f'me and short in first l

postpaleal setiger (segment 4t5), stiff and longer from segment 7, intermediate in segment 6, limbate with

hairlike tips throughout (Fig. 5.5E). Thoracic uncini with double row of 3 or 4 large teeth over basal prow

(Fig. 5.5F); abdominal uncini similar, largest ones about size of smallest thoracic uncini (Fig. 5.5G). I!

First 3 or 4 abdominal segments with tiny, papilliform notopodial rudiments. Pygidium small, with

2 long, slender dorsolateral cirri and several low, crowded papillae, i

112 I
!

!

| / ooo

! B

!

' i !il

! F

!

I Figure 5.$. Ampharete labrops: A, entire animal, lateral view; B-C, anteriorend,schematic, dorsal and ventral views; D, tip of palea; E, notoseta; F-G, dorsatmost and ventralmost thoracic uncini from same

fascicle; H, abdominaluneini, 3 different views.

!

I area between Methylbranchial Green bases Staining on segment Pattern.4, 5Dorsal and 6;stain ventrally on prostomi stainingum, through segments segment 1 and2 except 2, andfor triangu opaque lar

white band on segment 2; ventral scutes through unciniger 9, narrow anterior band on setiger 10; pampodia,

I branchiae,Remarks. and buccal A tentacles not staining. . labrops is easily distinguishable from its congeners by the band of eyespots on the

upper lip; it differs further from the co-occurring A. acutifrons and A. finmarchica by the short, brassy,

I indistinctly limbate paleae and the ventrally separate, rather than fused, first 2 segments.

i 113
!

II

Type Locality and Type Specimens. California, off Santa BarbaraPoint: holotype (LACM-AHF). n

Biology. The worms build in silty tubes incrusted with shell fragments; they live in sand and silt

with kelp. I Distribution. Central and southern California, 54 to 65 m.

Genus Amphicteis Grube, 1850 I

Type Species. Amphitrite gunneri Sars, 1835.

Ill

Diagnosis. Four pairs of smooth branchiae, usually cylindrical, occasionally flattened, on 2 visible 1

segments, arranged in 2 rows of 2 pairs (2+2), divided by median branchial lamella. Prostomium with

glandular ridges; buccal tentacles smooth. Paleae present; 18 thoracic setigers including paleae, 14

uncinigerous (p+3), 15 abdominal uncinigers; abdominal notopodial rudiments present. ii

Remarks. Species of Amphicteis are characterized by the elaborate, complex prostomium and a

high number of thoracic uncinigers; there are two other genera with 14 thoracic uncinigers (Samytha and

Amphisamytha) in the Californian fauna, but they lack paleae. Amphicteis is one of the largest ampharetid U

genera with 27 valid species that are widely distributed in the world's oceans. Hartman (1969) listed three

species, two of which were found in the MMS Phase I and II materials, n

Amphicteis mucronata Moore, 1923

Figure 5.6 I

Amphicteis mucronata Moore, 1923:203-206.--Hartman, 1969:547-548, fig. 1.--Fauchald, 1972:284-285, n

pl. 58, fig. a.--Lissner et al., 1986:D-17.---Steinhauer and Imamura, 1990:F-1. n

Amphicteis scaphobranchiata: Lissner et al., 1986:D-17 (in part). Not Moore, 1906.

Material Examined. California: Santa Maria Basin, offPoint Sal, Sta. R-8 (1), Sta. 40 (1); Western I

Santa Barbara Channel, Sta. 79 (1), Sta. 85 (2).

Description. Length to 48 mm, width to 4.5 mm; 17 thoracic setigers, t4 uncinigerous, and 15 n

abdominal uncinigers. Body long, slender, gently tapering toward pygidium (Fig. 5.6A).

m

Prostomium longer than wide, with mid-superior lobe divided into 2 L-shaped ridges joined by i/

thin, lamellar middle part; posterior prostomial lobe surrounding mid-superior lobe, bearing long, conspicuous II

glandular ridges on anterior margin; central part of prostomium bearing 2 lateral groups of small eyespots,

usually concealed by glandular ridges and not visible from above (Fig. 5.6B). Single (nephridial?) papilla in ll

center of prostomium between glandular ridges. Segments 1 and 2 fused, produced into crenulated lower lip I

ventrally. Branchiae long, with broad bases leaving fairly large scars when breaking off, tapered to filiform

tips; in 2 groups of 2, separated by narrow median gap occupied by notched branchial membrane or pad. i

Paleae well developed, about 10-15 per fascicle, coppery, tapering to slightly hooked, rounded tip 1

bearing a distinct mucron (Fig. 5.6C); older paleae lacking mucron, appearing distally rounded. First notopodia

on segment 4, reduced, with few fme setae; parapodia on segments 5 and 6 gradually increasing and with I

larger setae; uncini from segment 7 (setiger 4). Fully developed notosetae stiff, limbate, covered with minute 1

surficial spines (Fig. 5.6D); thoracic and abdominal uncini similar, with single row of 4-5 teeth above

inconspicuous rostral point and narrow basal prow (Fig. 5.6E, F). Notopodia with distinct, foliaceous ventral !1

cirri and conical presetal lobes (Fig. 5.6G); abdominal neuropodia with dorsal cirri (Fig. 5.6H). Abdominal |

notopodial rudiments present throughout, rounded, small; neuropodia progressingly elongate toward

pygidium, last pair projecting beyond anus, with small dorsal cirri. I

114 I
I Figure 5.6. Amphicteis mucronata: A, entire animal, dorsal view; B, anterior end, schematic, dorsal view; C,

palea; D, detail of notoseta; E-F, thoracic and abdominal uncini, laterai and frontal views; G,

notopodium, anterior view; H, abdominal neuropodium, anterior view; I, posterior end, dorsolateral

I view.

I 115
!

Pygidium small, with 2 long, filiform dorsal anal cirri and several low, rounded lateral and ventral •

lobes (Fig. 5.6I).

Methyl Green Staining Pattern. Prostomium staining except for median area around eyes, glandular

ridges dark green; branchial lamella and dorsal bands on first 5 setigers staining, remainder of dorsum II

remaining white; lower lip staining, with darkest cells anteriorly, weakly defined whitish band in the middle,

and lighter stain posteriorly; ventral scutes staining very dark throughout, fading in last 2 thoracic segments. •

Gills and buccal tentacles white. II

Remarks. Although the mucrons of the paleae are an unmistakable character ofA. mucronata, they

may be difficult to see and seem to wear off; however, the crenulated lower lip with its characteristic methyl

green staining pattern, as well as the distribution of segmental dorsal bands picking up the stain are diagnostic i

characters that can be readily observed in the dish under a dissecting microscope. The co-occurring species

A. scaphobranchiata is characterized by a smooth lower lip with a different staining pattern and dorsal stain II

throughout, and the branchial membrane has 2 anterior notches rather than a single notch. These characters

are sufficient for species discrimination even if the characteristic foliaceous branchiae are broken off, and

they were used to reassign some of the Phase I voucher specimens, i

i

Type Locality and Type Specimens. California, off Point La Jolla: holotype (USNM 17240) and

2 paratypes (ANSP 3109). Additional type material: Monterey Bay, off Point Pifios Lighthouse: 3 paratypes

(ANSP 3112, 3114), 2 paratypes (USNM 17257). i

Ill

Habitat. Moore (1923) reported the species from green mud and sand; the Santa Maria Basin

specimens were collected from sediments with 70 to 84% sand. i

Distribution. Western Canada to southern California, 110 to 1177 m. H

Amphicteis scaphobranchiata Moore, 1906 iw

Figure 5.7

Amphicteis scaphobranchiata Moore, 190@255-257, pl. 12,figs. 54-61; 1923:202-203.--Hartman, 1969:549- I

i

550, figs. 1-6.--Fauchald, 1972:286-287, pl. 58, fig. c.--Banse, 1979:1545.-- Uebelacker, 1984:51-

18 - 51-20.--Lissner et al., 1986:D- 17 (in part).---Steinhauer and Imamura, 1990:F- 1. 1

Ill

Material Examined. California: Santa Maria Basin, off Point Sal, Sta. PJ- 1 (1); off Purisima Point,

Sta. 42 (1). i

Description. Length to 54 mm, width to 3.5 mm; 17 thoracic setigers, 14 uncinigerous; 15 abdominal i

segments. Body with wide thorax tapering to narrower abdomen; color in alcohol uniformly tan.

Prostomium with mid-superior lobe divided into 2 straight, thick ridges; posterior lobe encircling i

mid-superior lobe on 3 sides, bearing long, L-shaped glandular ridges on anterior margin. Middle of W

prostomium beating lateral groups of 1 to many eyespots and paired nuchal organs median to eyes, often

concealed by glandular ridges. Segments 1and 2 fused, produced ventrally into smooth lower lip. Branchiae g

large, as long as 10 setigers, inner anterior pair foliaceous with scalloped anterior margin bearing a filiform

tip, others cylindrical and tapered; in 2 rows of 2, separated by simple branchial membrane (3-lobed in

juveniles) about half as wide as branchial bases (Fig. 5.7A). Anterior 3 notopodia small and conical, with •

triangular postsetal lobes; subsequent notopodia larger, with foliaceous ventral cirri in addition to postsetal 1

lobes (Fig. 5.7B, C). Both thoracic and abdominal neuropodia welt-developed, paddle-shaped, widest in

thoracic uncirliger 2, gradually narrowing to about half maximum width in last thoracic unciniger; abdominal 1

neuropodia longer and narrower than thoracic, gradually lengthening toward pygidium, with short dorsal I

cirri (Fig. 5.7B); last neuropodia projecting beyond pygidium. Abdominal notopodial rudiments small but

distinct, involute-digitiform to rounded. I

116 "1
!

I Figure 5.7. Amphicteis segments,latscaphobra eralviewnchiata ; C,poste :A,rior entirnotoeanima podiuml,dorsolat ,anterioreral viview ew; ;D,B,plale astathor ;E, athcicoracic andfiunrsteaini, bdominal lateral

andfrontalview; F, abdominaluncinus;G, posteriorend, lateral view. (A, E, F modified after

i Uebelacker).

i Paleae wide, straw-colored, tapering abruptly to hairlike tips (Fig. 5.7D), about 7-13 per fascicle. Setae fairly stout except in fast 2-3 setigers, limbate; thoracic uncini with 4-6 teeth in single row above

small, rounded basal prow; abdominal uncini with several rows of teeth above large basal prow (Fig. 5.7E, F).

tufts; noPygidium additional with pygidial 2 slender, lobes obclasverved ate dorsolateral (Fig. 5.7G).anal cirri covered with scattered papillae or ciliary

Methyl Green Staining Pattern. Prostomium staining except for area around eyes and nuchal

I organs, glandular ridges staining weakly; branchial membrane and 1 or 2 wide transverse dorsal bands directly behind branchiae, followed by lighter dorsal stain throughout length of thorax; lower lip with light

green anterior half, distinct white middle band, and very dark blue-green posterior part; ventral scutes

I distinct on all setigers; gills and buccal tentacles white or with scattered dark blue-green cells.

I 117
!

|

Remarks. If well-preserved and complete, A. scaphobranchiata is one of the most spectacular and •

unmistakable ampharetid species found in California; however, if the branchiae are missing, the species can

be misidentified as the co-occurring A. mucronata. The two species can best be told apart by the methyl •

green staining pattern, particularly that of the lower lip, and also by the shape of the lower lip; it is crenulated !1

in A. mucronata, but smooth in A. scaphobranchiata. Morphological details of the paleae are helpful for

species discrimination, but should be examined on dissected paleae under a compound microscope. Under i

low magnification and in situ, the mucrons of A. mucronata are difficult to distinguish from the abruptly II

tapering, but continuous tips ofA. scaphobranchiata.

Type Loeality and Type Specimens. British Columbia, Queen Charlotte Sound, Vancouver Island,

off Fort Rupert: holotype (USNM 5507). O

Habitat. Found in mud and fine sand; the Santa Mafia Basin specimen was collected in sediment

containing 69% silt. I

am Distribution. Washington to Baja California; Gulf of Mexico; 14 to 1980 m.

Genus Amphisamytha Hessle, 1917 I

d

Type Species. Samytha bioculata Moore, 1906.

Diagnosis. Four pairs of smooth branchiae on 2 or 3 visible segments, arranged 3+1, 2+2, or 2+1+1, i

median gap narrow. Prostomium without glandular ridges; buccal tentacles smooth. Paleae absent; 17 thoracic

setigers, 14uncinigerous; 13 or 14abdominal uncinigers, abdominal notopodiai rudiments present. Pygidium am

with 1pair of anal cirri, i

Remarks. The genusAmphisamytha was recently split by Williams (1987) based on the arrangement

of the branchiae and the development of the abdominal notopodial rudiments. Hessle (1917) described the Hi

branchial arrangement of the type species,A.japonica, as 3 anterior and 1 posterior pair, whereasA, bioculata |

has 2 anterior and 2 posterior pairs. Specimens of the third known species ofAmphisamytha, A. galapagensis,

were examined to determine whether the branchial arrangement is a valid generic level character. It was m

found that depending on the degree of relaxation at the time of death, the branchiae appeared to group into !i

2 anterior pairs on segment 3 and 2 posterior pairs on segment 4, or into an anterior pair on segment 3 and

2 single branchiae associated with segments 4 and 5. It seems therefore that branchial configurations are not m

a good character in this genus, even though it is very consistent in other genera. Another character listed by I!

Williams (1987) is the presence of"dorsal cirri" on the abdominal notopodial rudiments. In the type species,

the notopodial rudiments are "small papillae" (Moore, 1906), and inA. galapagensis, the notopodial rudiments i

are reduced to distinct but low ridges with a very small free distal part barely emerging from the dorsal body •

wall. Like the branchial arrangement, the development of the notopodial rudiments is apparently variable

i

among species. If Williams' (1987) generic def'mitions were to be followed through, Amphisamytha would nil

have to be split into three genera, which is neither justifiable nor desirable considering that so many monotypic

genera already exist in the family Ampharetidae. The genus Mooresamytha should, therefore, be considered

m

as a junior synonym of Amphisamytha.

Amphisamytha bioculata (Moore, 1906)

Figure 5.8 I

Samytha bioculata Moore, 1906:253-255, pl. 12, figs. 52-53. ID

Amphisamytha bioculata: Berkeley and Berkeley, 1952:73.--Hartman, 1969:551, figs. 1-5.--Fauchald and I

Hancock, 1981:40.--Lissner et al., 1986:D-17.

Mooresamytha bioculata: Williams, 1987:257, figs. 4A, B. I

118 1
I Material Examined. No specimens available, but species reported during Phase I reconnaissance.

Description. Length to 30 mm; 17 thoracic segments, 14 uncinigerous, 13 or 14 abdominal segments.

I Body tumid, with long and quadrate thorax and much thinner abdomen. Prostomium indistinctly trilobed (Fig. 5.8A), with large eyes at postectal comers of mid-superior

lobe. Segments 1 and 2 fused, ventrally forming large, areolate to crenulated lower lip (Fig. 5.8B). Segment

I 3 with first 2 pairs of branchiae; segment 4 with posterior 2 pairs of branchiae and minute first notopodia bearing few very fine setae. Branchiae slender, slightly flattened, about 3 times as long as body width.

Segments 5 and 6 with gradually increasing notopodia bearing gradually lengthening setae; fully developed,

i biramo abdominal us par parapodia apodia frowithm segment 7 through segment 20. Notopodia cylindrical, simple throughout; small neuropodial pinnules and larger, glandular notopodial rudiments (Fig.

5.8C). Notosetae narrowly limbate, uncmi similar throughout, with 4-5 teeth in a row over small basal prow

I (Fig. 5.8D-F). Pygidium with 2 ventrolateral anal cirri. Tubes about twice as long as animal, with thin mucous

wall covered with often thick layer of silt particles and incrusted with sponge spicules on upper end.

I Remarks. This species, transferred to anew monotypic genus, Mooresamytha, by Williams (1987),

is here moved back into Amphisamytha for the reasons listed above in the Remarks to the genus. A. bioculata

i is best recognized by the great number of thoracic uncinigers.

! B

| A .

!

!

i!

Figure 5.8. AmphisamNha bioculata:A, anteriorend,dorsal view; B,same,ventral view;C, abdominalsegments,

I lateral view; D-E, thoracicuncini;F, abdominaluncinus. (A-Cfrom Williams; D-Ffrom Hartman).

i 119
!

n

Type Locality and Type Specimens. Western Canada, British Columbia, Halibut Bank; holotype n

(USNM 5540).

Habitat. Moore (1906) reported sticky green mud and fine sand at the type locality. B

U Distribution. Canada to southern California, 57 to 774 m.

Genus Anobothrus Levinsen, 1883 N

Type species: Ampharete gracilis Malmgren, 1866.

Diagnosis. Four pairs of smooth branchiae on 2 visible segments, arranged in transverse row of 3 I_

pairs and single pair behind (3+1). Prostomium without glandular ridges; buccai tentacles smooth. Paleae

present; 15 or 16 thoracic setigers including paleae, 12 uncinigerous (p+2 or p+3); notopodia of unciniger 8 •

or 9 (f'Lfth-to-lastor fourth-to-last) elevated, connected by dorsal glandular ridge, but not structurally modified;

12-14 abdominal uncinigers; abdominal notopodial rudiments absent; pygidium without anal cirri, but

surrounded by papillae, n

Remarks. The genus Anobothrus has been subject to many erroneous definitions, mostly because U

of the different methods that were used in the past to count segments. The most reliable character at the

generic level, the modified thoracic segment with elevated notopodia and a well-defined connecting dorsal 1

ridge, has been described as setiger 13 (Hartman, 1969 and Fauchald, 1972 for A. gracilis), setiger 15 n

(Hartman, 1969 for A. occidentaIis), setiger 10 or 11 (Fauchald, 1977), and---correctly--as unciniger 8

(Day, 1964). Part of the confusion is due to the fact that the number of notopodia on segments 4 to 6 differs n

among species, and also the difference among authors in including or excluding the paleae from the notopodiai U

count. Day (1964) considered synonymizing Anobothrus and Sosane, a genus with elevated notopodia on

the third-to-last thoracic segment (unciniger 10); the suggestion was rejected by Williams (1987) and is also III

rejected here. U

There are, to date, five valid species of Anobothrus; of the three species in Hartman's (1969) Atlas

of Polychaetous Annelids, only one is still in that genus (A. gracilis), and it was also present in the MMS m

Phase I and II materials. Another species was found to be new to science. I

Anobothrus gracilis (Malmgren, 1866) n

Figure 5.9

I: Ampharete gracilis Malmgren, 1866:365, pl. 26, fig. 75.

Anobothrus gracilis: Levinsen, 1883:162.--Wolleb_ek, 1912:50.--Hessle, 1917:106-107 (synonymy). -- I

Hartman, 1969:553-554, figs. 1-2.--Zottoli, MS:27-28, figs. 10A-F.--Banse, 1979:1546. --Lissner i

et al., 1986:D-17.-'Steinhauer and Imamura, 1990:F-1.

Anobothrus bimaculatus: Lissner et al., 1986:D-17.

U

Material Examined. California: Santa Mafia Basin, off Point Sal, Sta. 104 (2); off Purisima Point,

Sta. R-5 (I); off Point Arguello, Sta. 66 (2). 1

Description. Length to 47 ram, width to 3 ram; 16 thoracic setigers, 12 uncinigerous (p+3), 14 n

abdominal setigers. Body slender, linear, gently tapering toward pygidium (Fig. 5.9A), with more or less

distinct constriction between second and third abdominal segment; color in alcohol uniformly tan. /

Prostomium divided into a median and 2 lateral lobes of equal size, sometimes with pair of eyespots. I

Peristomium and segment 2 fused laterally and ventrally, produced into smooth lower lip. Branchiae

cylindrical, slender, arising from segment 3 in a straight row of 3 and 1 behind on either side (Fig. 5.9B). m

120 I
I

I

I

I \!

I

J

I b

i /

I

I

I Figure 5.9. Anobothrus gracilis: A, entire animal, lateral view; arrows: dorsal ridges; B, anterior end, dorsal

view, schematic; C, tip of regular notoseta; D, tip of modified notoseta, unciniger 8; E-F, thoracic

I and abdominal uncini, lateral and frontal view; G, posterior end, lateral view.

I on Paleae slender, tapering gradually to hairlike tips, in fan-shaped fascicles of about 16. First notopodia segment 4, directly behind base of outermost gill, reduced to thin fascicle of short, very fine setae, easily

overlooked; notopodia 2 and 3 (segments 5 and 6) small but distinct, bearing larger, limbate setae (Fig.

i 5.9C). fifth-to-las Notopodia t setiger 6 (setiger (segment 11, 9, unciniger unciniger 3)8) connected slightly raised by oftendorsally, weaklyconnected defined dorsal by prominent, ridge; notopodia thickenedof

dorsal ridge, and bearing modified setae with finely pilose, more abruptly tapering tips (Fig. 5.9D). Uncini

I first present in setiger 4 (segment 7); thoracic and anterior abdominal uncini with double row of 4 teeth

i 121
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|

above rostral point and rounded basal prow, middle and posterior abdominal uncini with 3-4 rows of 3-7 i

teeth above rostra] point and basal prow (Fig. 5.9E, F). Notopodia simple, conical, without lobes or cirri;

thoracic neuropodia wide, paddle-shaped, abdominal neuropodia narrower and gradually lengthening toward i

pygidium; dorsal cirri absent. |

Pygidium with terminal anus surrounded by 2 large ventrolateral and 3 small dorsal papillae (Fig.

5.9G). Tubes with mucous inner lining and thin outer layer of fine mud particles. 1

M IIethyl Green Staining Pattern. Dorsum staining dark green through unciniger 3, with ridge

connecting notopodia appearing as thin white line; light green stain throughout posterior thorax, ridge on

setiger 11 white; lower lip white anteriorly, staining light green posteriorly; ventral scutes dark through 1

unciniger 7, then gradually fading.

i

Remarks. Hartman (1969) included some erroneous counts in her description of this species: the i

number of thoracic uncinigers is 12 rather than 13, and the elevated setae are on segment 11 rather than 13. •

Fauchald (1972) described Anobothrus bimaculatus, a species with 11 thoracic uncinigers, and may have iN

followed Hartman's erroneous segment counts for A. gracilis. It is therefore possible that A. bimaculatus

has to be moved to another genus. IIi

Type Locality. Western Sweden.

Habitat. In mixed and silty sediments, i

Distribution. Arctic and boreal Atlantic Ocean, New England, Bermuda; California; Sea of Japan; !1

10 to 2900 m.

I1

Anobothrus paleatus Hilbig, new species U

Figure 5.10 I

Anobothrus sp. A Lissner et aI., 1986:D-17. 1

Anobothrus sp. i SAIC, 1992:B-2. 1

Material Examined. California: Santa Maria Basin, off Point Sal, Sta. 36: holotype (USNM) and 4

paratypes (USNM); 5 paratypes (LACM-AHF Poly), 2 additional specimens (USNM), 3 additional specimens [I

(SBMNH); off Farallon Islands, north of Pioneer Canyon, Sta. 3-20, 37°25.04'N, 123°01.85'W, 610 m, coll.

J.A. Blake 22 September, 1991 (4).

Description. Holotype incomplete 5 mm long, 0.75 mm wide for 21 segments; other specimens: i

i

length to 5 mm, width less than 1 ram; 15 thoracic setigers, I2 uncinigerous (p+2), 12 or 13 abdominal U

segments. Body slender, with thorax accounting for at least two-thirds of body length; abdomen with

constriction after third segment, tapering slightly toward pygidium (Fig. 5.10A); color in alcohol pale. 1

m

Prostomium small, 3-lobed, concealed by paleae and branchial bases; eyes and nuchal organs not

observed. Segments 1and 2 free dorsally, hidden under branchial fold, fused and somewhat obscured laterally, im

and free ventrally, with segment 1 produced into an oval lower lip and segment 2 forming a short but I_

w

distinct ring (Fig. 5.10B). Branchiae about as long as fast 12 segments, filiform, arising in single row from

high dorsal fold; paleae prominent, iridescent, wide with curved sides, tapering to fine tip; arranged in

dense, fan-shaped, nearly circular fascicles (Fig. 5.10B-D). Segments 4 and 5 fused, bearing small notopodium 1

with well-developed notosetae; second notopodium on segment 6, also reduced in size but bearing wellm

developed setae. Fully developed, biramous parapodia from segment 7; notopodia simple, cylindrical

throughout, thoracic neuropodia very short pinnules, abdominal neuropodia longer but bearing fewer uncini. 1

Notopodia 5 (unciniger 3) connected by dorsal glandular band;notopodia of fourth-to-last setiger (unciniger

I

9) slightly elevated, connected by densely ciliated dorsal ridge, and bearing modified setae with rounded

I

122 l
I

I Figure 5.10. Anobothruspaleatus ltilbig, new species: A, entire animal, lateral view; arrows: dorsal ridges; B, anterior end, frontal view; C, same, lateral view; D, palea; E, modified seta; F, detail of tip; G,

normal seta (D, E, G drawn to scale); H, thoracic uncinus, anterolateral view.

!

I 123
!

II

tips and very thin mucron, freely piiose under high magnification (oil immersion) (Fig. 5.10E, F). Regular g

setae limbate, gradually tapering to capillary tip (Fig. 5.10G). Uncini similar throughout, with 3 rows of 4 to

5 teeth above rounded basal prow (Fig. 5.1 OH). III

Pygidium very small, ring-shaped, with 2 papilliform lateral anal cirri bearing basal pigment spots II

and about 4 very small dorsal and ventral papillae. Tubes delicate, with outer layer of fine sediment particles

and occasionally encrusted with very coarse sand grains. 1

II Methyl Green Staining Pattern. Ventrum staining strongly through unciniger 6, then fading, stain

absent after unciniger 9; lower lip white except for single dark glandular cells; no dorsal stain except for

prostomium and front of branchial fold. R

D

Etymology. The species is named for its most conspicuous character, the paleae.

Remarks. The striking paleae easily distinguish Anobothrus paIeatus from A. gracilis. In addition, i

the two species differ in the number of notopodia in segments 4 to 6 (3 in A. gracilis, 2 in A. paleatus), the l

position of the elevated notopodia (fifth-to-last setiger in A. gracilis, fourth-to-last setiger in A. paleatus),

the position of the abdominal constriction (between abdominal segments 2 and 3 in A. gracilis, between l1

segments 3 and 4 in A. paleatus), and the morphology of the modified setae. I

Habitat. In mixed sediments with about equal amounts of sand and silt.

Distribution. California, in 550 to 610 m. i

Genus Asabellides Annenkova, 1929 II

Type Species. Sabellides sibirica Wirtn, 1883. I_

Diagnosis. Four pairs of smooth branchiae on 2 visible segments, arranged in row of 3 pairs and the ill

fourth pair behind (3+1). Prostomium without glandular ridges; buccal tentacles pinnate. Paleae absent; 14 II

thoracic setigers, 12 uncinigerous, 10-14 abdominal uncinigers; abdominal notopodial rudiments present.

Remarks. Asabellides is a small genus with only four valid species that were all described from l

Arctic or subarctic waters. The only species known from California, A. lineata, was described by Hartman II

(1969) and was also collected during the MMS Phase I and II programs. In addition, two new species were

found. Asabellides is the only genus with four pairs of branchiae, 12 thoracic uncinigers, no paleae, and no II

modified thoracic setigers. I

am

Asabellides cahfornica Hilbig, new species 1

Figure 5.11

Asabellides sp. B Lissner et al., 1986:D-17. _l

Material Examined. California: Santa Maria Basin, off Purisima Point, Sta. BRA- 13, 34°42.57'N, in

120°47.90'W, 92 m: holotype (USNM); Western Santa Barbara Channel, Sta. BRC-1, 34°24.08'N, II

120°00.44'W, 74 m: 1 paratype (LACM-AHF Poly).

Description. Length to 8 mm, width to 1 mm; 14 thoracic setigers, 12 uncinigerous, 11 abdominal I

segments. Body linear in thoracic region, abdomen gradually tapering; thorax very long in relation to abdomen I!

(Fig. 5.1 IA). Color in alcohol pale.

Prostomium wider than long, trilobed, with all lobes of equal width; eyes or nuchal organs not I!

observed. Buccal tentacles few, wider than branchiae, distal part densely covered with cilia, with very long, I!

stiff pinnae about 3-4 times as long as tentacular width (Fig. 5.11A, B). Segments 1 and 2 fused dorsally,

obscured by branchial fold; free ventrally, with segment 1produced into small, smooth lower lip. Segment i

124 I
I

i

•...,_.x:- ..... ... :._.:._' .. .'_._

.....'_' - -'_Li_.'_'''_"?,_ i:-,l _:'"":'- ? ' _ _-

...,...-... .._.•.:=_,-_:..... , ._....:!_..'.._-_._- ..... .... _..

..,_......_:. :._-_iL_._ :. _ :=._ _._-......... - _ _'__

,..._-,::"i" .__-": _. .-:._..::,.._: : "....... 'I _F

_i_'_.:,;i; , "._I:__L..S'_ ,_ _/_ -'_ ' " 'J/_

I _'E_ _ F

I c

I Figure 5.11. Asabellidescalifornica Hilbig,new species:A, entireanimall,ateralview;B, detailofbuccaltentacle;

C-D, first and third abdominal neuropodium; E, thoracic uncinus, frontal view; F-G, anterior and

i posterior abdominaluncini, lateral views.

I 3 with closely set branchiae, 1pair slightly behind other 3,separated by wide median gap; 2 distinct nephridial papillae emerging between left and right branchiae. Segments 4 and 5 fused, bearing first notopodium;

second notopodium on segment 6.

I Biramous parapodia from segment 7; notopodia cylindrical, thoracic neuropodia short pinnules; abdominal neuropodia lengthening, with rounded dorsal cirri shorter than neuropodial pinnules (Fig. 5.11C,

D); last neuropodia reaching edge of pygidium. Notosetae in small fascicles, limbate; thoracic uncini with

I double row of 4 teeth over conical basal prow (Fig. 5.11E); abdominal uncini with 3 irregular rows of teeth, increase in number of teeth gradual (Fig. 5.11F, G).

Pygidium much narrower than last segments, with 2 long, filiform anal cirri, about as long as last 3

I to segments.

5

Methyl Green Staining Pattern. Dorsal stain very deep on prostomium, moderate on fused segments

I 1 and 2; 2 wide transverse bars across first 2 setigers; ventral stain on shields of all setigers except for last 4;

all notopodia staining, abdomen with short lines of staining ceils right above neuropodia; bases of anal cirri

staining; buccal tentacles and gills white.

!

i 125
!

I

Remarks. Asabellides californica is the only known species with 11 abdominal segments; it can []

further be distinguished from A. lineata (see below) by the short and round rather than long and filiform

dorsal cirri in the abdomen, and by differences in the methyl green staining pattern; A. lineata stains very

dark behind the branchiae and around the bases of the first notopodia, but does not stain in the abdomen, I

whereas A. californica exhibits two dorsal bands behind the branchiae and some stain on the abdomen.

Asabellides cornuta (see below) differs from A. californica in the presence of hornlike extensions on •

prostomium and fused segments 1 and 2, and also by the absence of any methyl green stain behind the I

branchiae.

Etymology. Named after its type locality. 1

ll Habitat. The species was collected at two hard-bottom stations; this may be the first record of a

species of Asabellides from hard substrate.

Distribution. Known only from California, 74 to 92 m. I

Asabellides cornuta Hilbig, new species I

U

Figure 5.12

Asabellides sp. A Lissner et al., 1986:D-17. I

Material Examined. California: Western Santa Barbara Channel, off Point Conception, Sta. 82,

34°18.71'N, 120°29.55'W, 394 m: holotype (USNM); Santa Maria Basin, off Point San Luis, Sta. 26, ll

35°04.38'N, 121°15.99'W, 590 m: 1 paratype (LACM-AHF Poly). II

Description. Length to 5 ram, width to 1 mm; 14 thoracic setigers, 12uncinigerous, longest specimen me

(holotype) with 3 abdominal segments (reported to possess 10 abdominal segments by Williams, unpublished |

data). Body wide anteriorly, tapering toward abdomen. Color in alcohol uniformly pale.

Prostomium indistinctly trilobed, roughly pentagonal, anterior corners drawn out to acute, hornlike •

tips; eyes, glandular ridges, and nuchal organs not observed (Fig. 5.12A). Buccal tentacles with 6 longitudinal I

rows of long, conspicuous pinnae on "dorsal" side, smooth on "ventral" side, with 2 rows of gland cells

(Fig. 5.12B). Segments 1 and2 fused laterally, separate dorsally and ventrally, drawn into acute anterolateral I

tips continuous with ventral anterior folds. Segment 3 also with ventral anterior fold, more conspicuous I

than that of segment 2; with slender branchiae arranged in 2 closely set rows of 2, separated by wide median

gap (Fig. 5.12A, C). Two nephridial papillae just inside branchial bases (Fig. 5.12A). First 2 notopodia on I

fused segments 415 and segment 6, large, well-developed, distally somewhat inflated, with full-sized fascicles |

of limbate notosetae.

Regular biramous parapodia from segment 7; notopodia large, slightly inflated to cylindrical, with []

stiff, long setae (Fig. 5.12D); thoracic neuropodia small pinnules, abdominal neuropodia longer, with rounded II

dorsal cirri shorter than pirmule (Fig. 5.12E). Notosetae narrowly limbate; thoracic and abdominal uncini

similar, with double row of 4-5 teeth above small rostral point and narrow basal prow; apical teeth smaller Q

than others (Fig. 5.12F-I). At least first 9 thoracic uncinigers with very pronounced ventral shields. I

Methyl Green Staining Pattern. Dorsal stain generally weak, on prostomium and fLrSt3 segments;

parapodia staining very deeply; ventral stain deep, encompassing all segments with ventral shields; buccal I

tentacles with 2 rows of deeply staining glandular cells opposite to rows of pinnae; branchiae not staining.

Remarks. AsabeUides cornuta is instantly recognizable by the strange, hornlike lateral extensions

of the prostomium, the fused first two segments, and the distinct ventral shields. Another unique feature is I

the short abdomen with only ten segments. Unfortunately, one of the specimens collected during Phase I, i

presumably the only complete one, is no longer available to confirm the number of abdominal segments.

I

126 i
!

I

il

i f C F

!

I igure 5 Hilbig, new specDies: A, anterior end, dorsal view,.b.,,, ottom of buccal cavity

I exnotopodium; tended; B, E,tipabdominal of buccal neuropodium; tentacle, 4 rowsF-G, of thoracic pinnae show and abdominal n; C, anteriuncini, or end,frontal ventralandview; lateral D,

views.

I Etymology. The species is named after the hornlike lateral extensions of the prostomium and the

fused first two segments.

I and is confined Habitat.toThe the supper peciesslope. seems to prefer very fine sediments with about 70% silt and up to 30% clay

Distribution. Known only from central California, 394 to 1000 m.

I

AsabeUides lineata (Berkeley and Berkeley, 1943)

I Figure 5.13

Pseudosabellides lineata Berkeley and Berkeley, 1943:131-132.

I Asabellides lineata: Berkeley and Berkeley, 1956:241.--Hartman, 1969:559-560, fig. 1.--Lissner et al.,

1986:D-17 (in part).--Steinhauer and Imamura, 1990:F-1.

I Sabellides octocirrata: Berkeley and Berkeley, 1942:201. Not (Sars, 1835).

Material Examined. Washington: Straits of Juan de Fuca, Dungeness Bay, MSMP Sta. 10,

48°10.50'N, 123°56.955V, 20 m, April 1989 (2).--California: Santa Maria Basin, off Purisima Point, Sta.

i R-5 (18); off Point Arguello, Sta. 65 (1).

I 127
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.. !

!

Figure 5.13. Asabeltides lineata: A, entire animal, dorsolateralview; B, tip of buccal tentacle; C, anterior end, 1

lateral view; D, last thoracic and ftrst abdominalsegments, lateral view; E, thoracic uncinus, frontal II

view;F-G, abdominal uncini, frontal and lateral views; H, posterior end, dorsal view. (Puget Sound

specimens). I

Description. Length to 15 ram, width to 0.6 mm; 14 thoracic setigers, 12 uncinigerous, 14 abdominal I_

segments. Body slender, linear, gently tapering toward pygidium (Fig. 5.13A). Color in alcohol pale.

Prostomium about as long as wide, trilobed anteriorly; mid-superior lobe largest, with acute tip; 2 []

large eyes at postectal comers of mid-superior lobe, connected by band of additional eyespots across posterior II

lobe. Segments 1 and 2 separate, segment 1 ventrally developed into smooth lower lip; buccal tentacles

slender, pinnate (Fig. 5.13B). Segments 3 and 4 fused; segment 3 bearing long, filiform branchiae, the 3 I

128 I
I anterior pairs in transverse row and fourth pair distinctly posterior to anterior pairs; middorsal gap small or

absent (Fig. 5.13C). Segments 5 and 6 bearing first notopodia, slightly smaller than subsequent ones and

I bearing Fully somewhdeveloped at reducedparapodia setae. present from segment 7; notopodia simple, cylindrical, with limbate

setae, neuropodia short, rounded pinnules with numerous crowded uncini. Abdominal neuropodia much

I longer with spreading than thoracic, row oflengthening uncini, withtoward conspicuous, posterior digitiform end, last pair dorsal reaching cirri longer edge ofthanpygidium; neuropodial broadlylobes rounded (Fig.

5.13D). Notosetae limbate; thoracic and abdominal uncini similar, with double row of 4-5 teeth over small

I• rostral (Fig. 5.13E-G). point and narrow basal prow; teeth in abdominal uncini in alternating rather than parallel positions

Pygidium very small, with terminal anus surrounded by several low, rounded or tapered lobes and

I 2 slender lateral anal cirri with brownish pigment spot or ring (Fig. 5.13H). Methyl Green Staining Pattern. Dorsal stain on prostomium except for areas around eyes, darkest

anteriorly; segments 1 and 2; narrow band right behind branchial bases; and very dark blue, broadly triangular

I band across middle of fused segments 4+5 (except for small areas around nephridial papillae) and large, dark rings surrounding white notopodial bases; ventral stain on all thoracic segments except for last 2.

i Remarks. This species is best recognized by its spectacular methyl green staining pattern. Diagnostic morphological details, such as the long dorsal cirri on abdominal neuropodia and the number of abdominal

segments, must be examined under a compound microscope because of the small size of this species.

i Ty Habi petat.LocInalitysandy . Northern silt, occupying Canada, fine Hudsomudn Btubes ay. resembling those of Myriochele heeri (Berkeley

and Berkeley, 1942).

I Distribution. Alaska to western Mexico, 15 to 200 m.

Genus Eclysippe Eliason, 1955

I Type Species. Lysippe vaneUi Fauvel, 1936.

I (occasionally Diagnos withis. 1Three pair slighpairs tly behind of smootherbranchiae 2), with oronwithout 1 visible mdiansegment, gap. Prostomium arranged inwithoutransverse glandular row

ridges; buccal tentacles smooth. Paleae present; 15 thoracic setigers including paleae, 12 uncinigerous (p+2),

I 12 abdominal Remarks. uncinigers; Ever abdominal notopodial rudiments absent. since Eliason (1955) described this genus, there has been a curious enigma in the

literature concerning one of the most basic diagnostic characters, the number of branchiae. Eliason based

I his genus, description on material of Eclysippe he collected , and himsubsequently self, ratherthethantransfer on Fauvel's of Fauvel's specimens (1936)which Lysippehe vanelli obviouslyto this didnewnot

examine. Eliason described three pairs of branchiae on his specimens, whereas Fauvel (1936) had reported

I four straight pairs, lineandof branchial he suspected basesthatto the sides. fourthOther "branchiae" authors were accepted indeedEliason's the paleal genenotopodia ric def'mition that ofcontinue Eclysippethe,

but often also acknowledged Fauvel's (1936) description of L. vanelli in form of a footnote or remark (see,

I for example, The Day, 1964 and Fauchald, 1977). genus contains two valid species. Hartman (1969) did not include Eclysippe in her Atlas of

Polychaetous Annelids, but did describe E. trilobata as Anobothrus trilobata. There are probably two or

I three 1987;yet 1992). undescribed deep-sea species collected both off the U.S. Atlantic and Pacific coasts (Blake et al.,

I

I 129
!

Eclysippe trilobata (Hartman, 1969) •

Figure 5.14

m

Anobothrus trilobatus Hartman, 1969:557, figs. 1-3. m

Eclysippe trilobata: Williams, 1987:252-253, fig. 1C.---Steinhauer and Imamura, 1990:F-1. 1

m

Material Examined. California: Santa Mafia Basin, off Point San Luis, Sta. 26 (7), Sta. 28 (1); off

Purisima Point, Sta. R-6 (2). m

Description. Length to 25 mm, width to 3 mm; 15 thoracic setigers, 12 uncinigerous (p+3), 12-14 m

abdominal segments. Body with wide and squafish thorax tapering quickly within last few setigers toward

narrower, linear abdomen (Fig. 5.14A); first 12 segments (through unciniger 6) considerably shorter and m

more rigid than last 6 thoracic uncinigers, separated ventrally by deep intersegmental furrow (Fig. 5.14C). l

Anterior thoracic notopodia in middorsal position and pointing upward because of expanded ventrum; in

middle and posterior thorax parapodia gradually shifting into lateral position and pointing to the sides, n

Color in alcohol pale. I

Prostomium pointing downward and surmounted by large branchial fold, anteriorly sometimes

divided into large median and 2 smaller lateral lobes, sometimes smoothly rounded depending on contraction im

of specimen; posterior part of prostomium forming a thick transverse ridge; central area between anterior II

and posterior part bearing paired nuchal organs. Segments 1 and 2 fused dorsally, concealed by branchial

fold, lengthening laterally, and ventrally produced into indistinctly crenulated lower tip (Fig. 5.14B, C). •

Branchiae cylindrical, fairly long and slender, arising from segment 3 in straight row, left and right group l

separated by narrow but distinct gap.

Paleae slightly wider and longer than subsequent setae, tapering to acute point, arising from large •

cylindrical lobes in narrow fascicles of about l0 (Fig. 5.14D). First notopodia on segment 4, simple, well

developed but very close to paleae and easily overlooked in small specimens, with setal fascicles similar to

those in following segments; setae stiff, limbate with short, acute tips. Last 5 notopodia with rounded dorsal •

and ventral lobes above and below setal lobe, indistinct in small specimens (Fig. 5.14F). Uncini first present 1

in setiger 4 (segment 7), first torus slightly displaced dorsally and about twice as long as all others (Fig.

5.14E); thoracic and abdominal uncini similar, with double row of 5 teeth above rostral point and rounded •

basal prow (Fig. 5.14G, H). Abdominal neuropodia short, narrow, inconspicuous; last pair not extending I

beyond pygidium.

Pygidium with terminal anus surrounded by 2 large, rounded lateral flaps and circlet of crowded, 1

narrow dorsal and ventral papillae. Tubes very thin and fragile, with outer layer of fine silt particles arranged

in regular, short, ring-shaped layers.

Methyl Green Staining Pattern. Prostomium with dark green frontal lobes and posterior ridge, II

middle part with nuchal organs unstained; first 2 segments visible as narrow green bands; front of branchial

membrane dark green. No additional dorsal stain. Buccal tentacles with 2 longitudinal rows of deeply staining

cells. All segments staining ventrally through setiger 10 (unciniger 7), after that staining ending abruptly; I

abdominal intersegmental furrows with ventral rows of glandular cells staining weakly; lateral pygidial

lobes staining dark green, papillae remaining white.

Remarks. In general, the examined specimens agree well with Williams' (1987) redescription, •

although the fairly wide median gap between the branchiae described from the holotype was not observed IB

on all present specimens. The width of the gap appears to be size dependent and was found to be largest m

(about 1.5 times branchial width) in small specimens, whereas the branchiae of the largest specimen were •

hardly separated at all. However, the other characters mentioned by Williams (1987) to separate E. trilobata Ig

from the very similar E. vanelIi, the large first notopodia and the methyl green staining pattern, were consistent

in allindividuals. The trilobed posterior notopodia are present in both species and may be a generic character, i

130 I
I Figure 5.14. Eclysippe trilobata: A, entire animal, dorsal view; B, anterior end, schematic, dorsal view; C, anterior

end, ventral view; D, palea; E, anterior end, lateral view; F, posterior notopodium; G-H, thoracic and

m abdominal uncini, 3 different views.

I 131
!

II

Type Locality and Type Specimens. California, east of Santa Catalina Island, 603 m: holotype •

(LACM-AHF Poly 0265) and paratypes (LACM-AHF Poty 0266).

Biology. Two of the largest Phase I voucher specimens were ovigerous; E. trilobata lives in silt or _l

silt mixed with sand.

Distribution. Central to southern California, in 400 to 691 m.

!

Genus Glyphanos¢omum Levinsen, 1884

Type Species. Samytha pallescens Threl, 1879. I

Diagnosis. Three pairs of smooth branchiae on 1 visible segment, arranged in nearly straight line

across dorsum, one pair slightly behind others (2+ 1). Prostomium without glandular ridges; buccal tentacles •

smooth. Paleae absent; 14 thoracic setigers, 11uncinigerous; 28 abdominal uncinigers, abdominal notopodial

rudiments absent.

Remarks. This genus, containing three valid species (Holthe, 1986a), seems to be typical for deep _l

water, although the type species G. pallescens occurs over a wide depth range from the upper shelf to the

continental rise. The species was also described by Hartman (1969).

I

Glyphanostomum paUeseens (Th_el, 1879)

Figure 5.15 I

Samytha pallescens Threl, 1879:61-62, pl. 4, figs. 60-62.

Sabellides pallescens: Wolleb_ek, 1912:67. I

Glyphanostomum pallescens: Hessle, 1917:105, fig. 11.--Hartman, 1969:561-562, figs. 1-3.--Zottoli, MS:32-

33, figs. 13A-C.--Holthe, 1986b:232.--Lissner et aI., 1986:D-17.

Material Examined. California: Santa Maria Basin, off Point Arguello, Sta. 69 (1). II

Description. Length to 32 mm, width to 1 mm; 14 thoracic setigers, 11 uncinigerous, and 28

abdominal segments. Body long, very slender, linear, tapering slightly toward pygidium (Fig. 5.15A). I_

Prostomium trilobed, scoop-shaped, dorsally and laterally surrounding large mouth opening, fused

with ftrst 2 segments forming lower lip. Buccal tentacles slender, very numerous (Fig. 5.15B). Branchiae 1

long, slender, cylindrical, 1 pair slightly behind other 2, arising from segment 3, separated by dorsomedian 1

gap.

Notopodia simple, cylindrical throughout; neuropodia small, with few uncini, slightly longer in ill

abdomen than in thorax. Notosetae slender and bilimbate; thoracic uncini with 5-6 large teeth in 1 or 2 rows II

over rounded basal prow (Fig. 5.15C); abdominal uncini smaller, with 2-3 rows of 2-4 crowded, very slender

teeth over basal prow (Fig. 5.15D). •

Pygidium with 2 short, slender lateral cirri and 2 rounded flaps inserted dorsally and ventrally. Tube I

thin, fragile, annulated, covered with f'me silt particles obscuring annulations and often encrusted with

sponge needles. I Remarks. The slender, linear body and the scoop-shaped prostomium cupped over the large mouth

give this species an external appearance that is unusual for ampharetids.

Type Locality. Arctic Ocean, Kara Sea, 75°40'N, 78°40'E, 45 m. !

u

Habitat. Muddy sand and silt mixed with sand.

Distribution. Arctic and Antarctic Oceans; western Canada to southern California; New England; Ill

Japan; 45 to 7100 m. i

132 I
Figure 5.15. Glyphano view,bucstcalomum pallescens: A, entireanimal,lateralview (after Zottoli);B, anteriorend,lateral tentaclesextended;C-D, thoracicandabdominaluncini,lateralandfrontalviews.

I Genus Lysippe Malmgren, 1866

Type Species. Lysippe Iabiata Malmgren, 1866.

I Diagnosis. Four pairs of smooth or areolate branchiae on 1 visible segment, arranged in 2 rows of

2 pairs (2+2). Prostomium without glandular ridges; buccal tentacles smooth. Paleae present, thin and short,

I not differing from other notosetae; 17 thoracic setigers including paleae, 13uncinigerous (p+3); 15abdominal setigers, abdominal notopodial rudiments present at least in anterior segments. Pygidium with 1 pair of anal

cirri.

I MalmgrenRemarks. There are only two genera with 13 thoracic uncinigers and four pairs of branchiae, Lysippe and Hypania Ostrooumov. They differ in the presence of glandular ridges on the prostomium

(absent in Lysippe); in addition, Hypania is a brackish and fresh water genus, whereas Lysippe is marine.

I Hartman (1969) described annectens in the Atlas of Polychaetous Armelids; that species

Lysippe

was moved to a new genus, Paralysippe, by Williams (1987), which differs from Lysippe in the number of

thoracic uncinigers (14 rather than 13).

!

I 133
Lysippe labiata Malmgren, 1866 •

Figure 5.16

Lysippe labiata Malmgren, 1866:367, pl. 26, fig. 78.--Hessle, t917:109- I 10.--Berkeley and Berkeley, i

1956:241.--Banse, 1979:1549.--Zottoli, MS.:34-36, figs. 15A-C.--Lissner et al., 1986:D-17; i

Steinhauer and Imamura, 1990:F-1. i

w

Lysippe sp. B: Steinhauer and Imamura, 1990:F-1.

Material Examined. California: Santa Maria Basin, off Point Sal, Sta. R-8 (1); off Point Arguello, 1

Sta. 69 (1); Western Santa Barbara Channel, Sta. 85 (1). am

Description. Length to 11 nun, width to 1 mm; 17 thoracic setigers including paleae, 13 uncinigerous I

(p+3), 11 to 15 abdominal segments. Body slender, widest in thorax, gradually tapering toward pygidium i

(Fig. 5.16A). Color in alcohol pale.

Prostomium trilobed anteriorly, with mid-superior lobe wider than lateral lobes; nuchal organs at •

postectal comers of mid-superior lobe (Fig. 5.16B), surrounded by large crescentic patches of golden brown l

pigment (may fade in alcohol). Buccal tentacles few, pinnate. Segments 1 and 2 fused, ventrally produced

into large, scoop-shaped, crenulate lower lip usually protruding beyond prostomium and visible in dorsal •

view. Segment 3 with small, thin paleae and anterior 2 pairs of branchiae. Posterior pairs of branchiae II

associated with segments 4 and 5, bases fused to dorsal body wall, free ends emerging between anterior

pairs; branchiae separated by wide dorsomedian gap. •

Segments 4, 5, and 6 with small notopodia, setae on segment 4 thin and needlelike, wider and II

limbate in all subsequent setigers. Notopodia from segment 7 fully developed; neuropodia large pinnules,

lengthening in posterior abdominal setigers. Thoracic uncini with 3 rows of 4-5 teeth above rostral point and •

basal prow (Fig. 5.16C); abdominal uncini crested, with about 4 rows of teeth above rostral point and basal

prow (Fig. 5.16D).

lobes. Pygidium with 2 thin lateral anal cirri, about as long as last segment, and several low, rounded i

Methyl Green Staining Pattern. Stain dorsal on prostomium, fused segments 1 and 2, front of

branchial membrane, and triangular patch on median branchial pad; scattered cells along entire body; ventrally: i

prostomium and all segments through unciniger 7 or 8, distinct ventral scutes from segment 6; last 5 thoracic i

setigers not staining except for scattered single cells, those continuing throughout abdomen. Notopodia

staining, also lateral band connecting neuropodial pinnules in both thorax and abdomen. H

Remarks. This species is easily recognized by the characteristic large, crenulated lower lip. It can

be distinguished from the very similar Paralysippe annectens (Moore) by the number of thoracic uncinigers i

(13 for L. labiata, 14 for P. annectens), the pinnate rather than smooth oral tentacles, and the scoop-shaped, i

posteriorly smooth rather than padlike, posteriorly deeply creased lower lip. The single Phase II specimen

originally identified as L. sp. B is identical to L labiata except for a very small size; the specimen is an

ovigerous female of only 6 mm length. It is therefore considered to belong to L. labiata. HI

Type Locality. Spitsbergen.

Habitat. Lysippe labiata is able to live in a variety of substrata; the Phase I voucher material is from •

locations with nearly 69% silt or about 72% sand. II

Distribution. Arctic, Greenland, Iceland, Denmark; Labrador to Massachusetts; Alaska to California;

Sea of Japan; 1 to 1030 m. •

!

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I

I i jj/

I

/

I c D

!

Figure 5.16. Lysippe labiata: A, entire animal, lateral view; B, anterior end, dorsal view; C-D, thoracic and

I abdominaluncirti,frontalview. (C, D afterZottoli).

• Genus Mexamage Fauchald, 1972

II Type Species. Mexamage corrugata Fanchald, 1972.

Diagnosis. Four pairs of wrinkled branchiae on 4 visible segments, arranged in longitudinal rows,

I wide interbranchial membrane. Prostomium with and lateral auricles; buccal separated by glandular ridges

tentacles smooth. Segments 1and 2 well separated, with lateral auricles. Segments 4, 5, and 6 with rudimentary

notopodia bearing few very thin notosetae, embedded or barely protruding; 11 clearly setigerous thoracic

I all uncinigerous; I 1 abdominal uncinigers, abdominal notopodial rudiments Pygidium segments, present.

with 0-2 papilliform anal cirri.

i Remarks. The genus Mexamage is one of the most interesting ampharetids because of the low

degree of cephalization; all prebranchial segments are well-developed and separate from each other (at least

ventrally). Fauchald (1972) considered this genus to be "structurally primitive". Mexamage is closely

related to Amage and Paramage; these genera differ from each other mainly in the development of the first

I six segments. To date, the genus consists of two valid species, both known from California.

I Mexamage eorrugata Fauchald, 1972

Figure 5.17

l Mexamage corrugata Fauchald, 1972:310-312, pl. 65, figs. a-c.

Amage arieticornuta: Lissner et al., 1986:F-17.--Not Moore, 1923.

I Material Examined. California: Santa Mafia Basin, offPoint Sal, Sta. 36 (1); Western Santa Barbara

Channel, off Point Conception, Sta. 89 (9).---Western Mexico, Guaymas Basin: holotype (LACM-AHF

i Poly 978).

I 135
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Description. Length to 37 mm, width to 5 mm; 14 podous thoracic segments, 11 uncinigerous, 11 II

abdominal segments. Body massive, with wide thorax about 3 times length of abdomen; body tapering

gradually toward pygidium. Color in alcohol pale. i

Prostomium appearing 3-lobed, with short, foliaceous mid-superior lobe surrounded by thick posterior

lobe; 2 nuchal organs at postectal corners of mid-superior lobe. Segments 1 and 2 fused dorsally, separate

ventrally, with segment 1 produced into deeply crenulated lower lip extending laterally into auricular lobes 1

(Fig. 5.17A, B), and segment 2 forming an apodous ring together with the fused dorsum of segments 1 and

2. Segment 3 bearing first pair of branchiae, ventrum extending laterally into auricular lobes; segments 4, 5,

and 6 with second, third, and fourth pairs of branchiae and rudimentary notopodia with f'me embedded •

setae, emerging in segment 6. Dorsum of segments 3, 4, and 5 with wide, crenulated branchial pad separating

left and right branchiae (Fig. 5.17A).

Segment 7 and subsequent segments with regular, biramous parapodia; notopodia simple, cylindrical 1

throughout, neuropodia fairly long pinnules with very crowded and numerous uncini, often more than 100

per fascicle. Abdominal neuropodia much narrower but longer than thoracic, with fewer uncini; last pair

projecting beyond pygidium (Fig. 5.17C). Notosetae stiff, coppery, limbate, covered with minute surficial 1

spines; thoracic and abdominal uncini similar, with row of 3 single teeth over 1 double tooth, 1 lower single

tooth, and small rostral point over rounded basal prow (Fig. 5.17D-F).

Pygidium short, inconspicuous, with wide anus surrounded by 8-15 low, rounded lobes and 2 short, 1

i

digitiform lateral anal cirri (Fig. 5.17C) Ill

Methyl Green Staining Pattern. Dorsal part of prostomium staining except for area around nuchal 1

organs; auricular lobes of segments 1 and 3, branchial pad; no dorsal stain behind branchiae. Ventral stain: i

lower lip and all subsequent thoracic segments; distinct ventral scutes from segment 5. All parapodial bases

i

with stain. m

Remarks. The examined specimens were originally identified as Amage arieticornuta, but the i

segmentally arranged gills and the rudimentary notopodia are a clear indication that they are Mexamage.

Aside from the exceptionally long gills in A. longibranchiata (Hartman, 1969) that surpass the body in []

length, M. corrugata can be distinguished from its very similar congener by proportions of the body; M. 1

corrugata tapers gradually toward the posterior end and has a wide pygidium, whereas M. Iongibranchiata

has an abruptly narrower abdomen and a small pygidium; the notopodia on segment 5 are larger in M. l1

corrugata than in M. longibranchiata; and the regular notopodia are much longer in relation to the segmental |

width in M. corrugata. Williams (1987) also mentioned differences in the methyl Green Staining Pattern,

but unfortunately she did not specify those differences. Fauchald (1972), in his description ofM. corrugata, II

reported simple capillary notosetae with "no trace of bilimbation;" examination of the large holotype revealed i

that the setae are narrowly limbate, with the wings more difficult to see than in the smaller Californian

specimens because of the thickness and intense coloration of the setae in the holotype. I

This record is the first for M. corrugata from California, representing a northward range extension I

from Mexico.

Type Locality and Type Specimens. Gulf of California, Guaymas Basin, ca. 1600 m: holotype _l

and 3 paratypes (LACM-AHF).

Habitat. The examined specimens were found in sand and silt.

Distribution. Western Mexico to central California, 525 to 1600 m. i

I

[]

I

136 I
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I A

I

I

Figure 5.17. Mexamage corrugata: A, entire animal, dorsal view; B, anterior end, lateral view; C, posterior end,

i ventrolateral view; D-F, uncini, 3 different views.

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Genus Mugga Eliason, 1955 •

Type Species. Mugga wahrbergi Eliason, 1955.

Diagnosis. Three pairs of smooth branchiae on 1 visible segment, arranged in transverse row or i

2+1. Prostomium without glandular ridges; buccal tentacles smooth. Paleae present; 13 thoracic setigers

including paleae, 9 uncinigerous (p+3), last notopodia elevated with modified setae crossing over dorsum; •

11 or 12 abdominal uncinigers, abdominal notopodiaI rudiments absent.

Remarks. Hotthe (1986b) described a second species ofMugga (M. bathyalis) from deep water off

southern Norway, but was obviously not convinced that this was indeed a new species because he could not

identify any good characters to discriminate the new species from M. wahrbergi. The genus Mugga may

therefore still be monotypic.

I

Mugga wahrbergi Efi_on, 1955

Figure 5.18 I

Mugga wahrbergi Eliason, 1955:7-8, fig. 2. mR

Mugga nr. wahrbergi: SAIC, 1991 :C-l.--Blake et al., 1992:A-1. i

Mugga sp. A Steinhauer and Imamura, 1990:F-1.

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. R-3 (1); off Point Sal, I

Sta. R-9 (2); off Purisima Point, Sta. R-6 (7).

Description. Length to 5.5 mm, width less than 1 mm; 13 thoracic setigers including paleae, 6 •

uncinigerous, 11 abdominal segments. Body slender, gracile, with relatively wide thorax and markedly II

thinner, nearly linear, beaded abdomen; color in alcohol pale.

Prostomium small, trilobed, often concealed by large peristomium; eyes or nuchal organs not

observed. Segments 1and 2 fused, forming large, collarlike structure, ventrally produced into smooth lower

lip (Fig. 5.18A). Segment 3 with numerous but very fine paleae and closely set, slender, long branchiae, left

and right group separated by wide median gap. Segments 4-6 with notopodia, those of segment 4 very close i

to paleae and easily overlooked.

Segments 7-14 with fully developed, normal biramous parapodia; notopodia slender, cylindrical,

long in relation to body width; thoracic neuropodia short, inconspicuous pinnules, abdominal ones wider,

with fewer uncini; parapodial cirri and notopodial rudiments absent. Notosetae limbate, slightly bent (Fig. I

5.18B); uncini crested, with 3 or 4 rows of 3-4 minute denticles above rounded basal prow (Fig. 5.18D-F).

Last notopodia (segment 15, unciniger 9) dorsally displaced, with ciliated, rounded basal flaps and modified I

notosetae crossing above dorsum; setae slightly wider than regular setae, roughly oar-shaped, distally hirsute I

(Fig. 5.18C); sometimes (when worn?) appearing distally truncate or excavate.

Pygidium short, wide, with terminal anus and 2 filiform lateral anal cirri about as long as last 2 I

segments (Fig. 5.18A). II

Methyl Green Staining Pattern. Dorsally prostomium and segments I and 2 staining, also membrane I

spanning median gap between branchiae; ventrally through unciniger 5, delineating distinct ventral scutes; 1

fading in subsequent segments, absent from last 2 setigers.

Remarks. Although this is a very small species, it is usually very easy to recognize even under tow II

magnification because of the manner in which the setae in the last thoracic segment cross over the dorsum. II

The genus Mugga is further characterized by the lowest known number of thoracic uncinigers among all

ampharetids. Despite these unique characters on the generic level, it seems to be difficult to find any species- I

l

.8 II
I

I

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I

I

I

I

I

I

I Figure 5.18. Mugga wahrbergi: A, entire animal, lateral view; B, regular notoseta; C, modified notoseta; D-F, uncird, different views.

I level characters (see Remarks on the genus above). The number of abdominal segments may be an additional

character not noted by Holthe (1986b) in his description and justification of M. abyssalis. According to

i Holthe, M. abyssalis possesses 12 abdominal segments, whereas the specimens examined here all have 11 abdominal segments. Eliason (1955) did not have any complete specimens, so that this issue could only be

resolved if additional material from the type locality were examined. Deep-sea specimens from the U.S.

i Atlantic slope have 12 abdominal segments and may be M. bathyaIis (personal observation), whereas two

specimens from the slope off the Farallon Islands, tentatively identified as M. hr. wahrbergi, were found to

have only 11 abdominal segments.

I Type BiologyLocality and Type Specimens. Skagerrak (Norway): holotype and paratypes (not located).

. Eliason (1955) reported ovigerous females occurring in July and August; in the present

material, two sexually mature females and one male were found in May and September. The eggs are

I nuthatmerous Muggaandwahrbergi flu the thomay racicbebody a deposit cavity.feeder. The inteThestinesspecies of some seemsspecimens to preferwere silt full or sandy of sedime silt.nt, indicating

Distribution. Widely distributed; Scandinavia, boreal Atlantic Ocean; California, 20 to 2400 m.

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Genus Paralysippe Williams, 1987 •

Type Species. Lysippe annectens Moore, 1923.

i

Diagnosis. Four pairs of smooth or areolate branchiae on 2 visible segments, arranged 2+2, median 1

gap as wide as branchia. Prostomium without glandular ridges, but with crescentic nuchal organs; buccal

tentacles smooth. Paleae present, small; 18 thoracic setigers including paleae, 14 uncinigerous (p+3); 14 l

abdominal segments; abdominal notopodial rudiments present. Pygidium with 1 pair of anal cirri. U

Remarks. Superficially, Paralysippe annectens closely resembles Lysippe labiata because it has a

similar crenulated lower lip, but it clearly does not belong to Lysippe because of the different number of I

thoracic uncinigers.

Paralysippe annectens (Moore, 1923) I

Figure 5.19

Lysippe annectens Moore, 1923:201-202, pl. 17,figs. 11-13.--Hartman, 1969:563, figs. 1-6.---SAIC, 1991:C-1. U

Paralysippe annectens: Williams, 1987:255-256, figs. 3A, B. 1

Material Examined. California: off Halfmoon Bay, south of Pioneer Canyon, Sta. EPA-1,

37°24.07'N, 123°14.89'W, 1263 m (1). n

Description. Length to 27 turn, width to 2 mm; 18thoracic setigers including paleae, 14 uncinigerous, 1

and 11 abdominal segments. Body widest at branchial bases, gradually tapering toward pygidium. Color in

alcohol pale with large, crescent-shaped brownish spots on prostomium.

Prostomium broadly trilobate, with large, crescentic nuchal organs at postectal margin of mid- 1

superior lobe, surrounded by conspicuous aggregation of golden brown pigment (Fig. 5.19A). Segments 1

and 2 very narrow, fused dorsally, separate ventrally, with segment 1 produced into deeply creased and •

crenutated lower lip; buccal tentacles numerous, slender, smooth. Segment 3 with small fascicles of fine, I

short paleae and closely set, slender branchiae, arising from high branchial fold, 2 anterior and 2 posterior

pairs, separated by median membrane. Segments 4 through 6 short but distinct, with well-developed notopodia •

and long setae. 1

Biramous parapodia from segment 7. Notopodia slender, cylindrical; neuropodial pinnules very

small, almost papilliform throughout. Notosetae long, slender, limbate; thoracic and abdominal uncini similar, •

with 3 or 4 irregular rows of about 5-6 small teeth above rostral point and small, rounded basal prow (Fig. 1

5.19B-D).

Pygidium very small, with 2 digitiform ventrolateral anal cirri and several small, rounded lobes or 1

papillae (Fig. 5.19E).

Methyl Green Staining Pattern. Dorsal stain on prostomium except for area surrounding nuchal i

organs, fused segments 1and 2, and anterior side of branchial fold; basal part of posterior side of branchial 1

fold; scattered single cells along entire body, particularly over last few abdominal segments. Ventral stain: 1

very deep on all segments through unciniger 7, weak bands on unciniger 8; scattered single cells to end of

body; notopodia and anal cirri staining, i

i

Remarks. This species was not collected during the Phase I and II programs, but is included here

because of its great similarity with the co-occurring Lysippe labiata Malmgren (see above). Aside from the i

additional thoracic segment, P. annectens can be distinguished from L. Iabiata by its lower lip that is not n

only anteriorly crenulated, but deeply creased posteriorly and padlike, whereas that of L. labiata is delicate,

w

posteriorly smooth, and scoop-shaped. Furthermore, the buccal tentacles ofP. annectens are smooth, whereas n

those of L. labiata are pinnate. •

140 I
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, i

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i I

I

B D

!

I Figure 5.19. Paralysippe annectens: A, anterior end, ventrolateral view (from Williams); B-C, thoracic uncini, frontal and lateral views; D, abdominal uncinus; E, posterior end, ventral view (from Hattman).

!

Type Locality and Type Specimens. Southern California, off Santa Rosa Island, San Miguel Island,

and San Diego: type (USNM 17326) and 2 syntypes (USNM 17327 and ANSP 3107).

I Habitat. In silty sediments.

Distribution. California, 70 to 2798 n_

I Genus Paramage Caullery, 1944

i Type Species. Paramage padurensis CauUery, 1944.

Diagnosis. Four pairs of smooth branchiae on 3 visible segments, arranged 2+1+1. Prostomium

without glandular ridges; buccal tentacles smooth. Paleae absent; 14 thoracic notopodia, 1 or 2 asetigerous

I vestigial, or setigerous, uncinigerous; or 11 uncinigers; abdominal notopodial

and 12 13 11 10 abdominal

rudiments present. Pygidium with 1 pair of papillae or flaps.

i below. ItRisemvery arksclosely . This gerelated nus currently toAmagecontains and differs onlyfromtwo spe theclatter ies, thein type the presence species anofdvestigial P. scutatanotopodia , describedin

the anterior thorax.

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Paramage scutata (Moore, 1923) •

Figure 5.20

I

Amage scutata Moore, 1923:210-212,pl. 17, figs. 19-24.--Lissner et al., 1986:D-17.--Steinhauer and

Imamura, 1990:F-1. Not Fauchald, 1972. Fide Williams, 1987. II

Paramage scutata: Williams, 1987:254-255, fig. 2B. B

Material Examined. California: Santa Maria Basin, off Point San Luis, Sta. R-1 (I); off Purisima

Point, Sta. 42 (1); Western Santa Barbara Channel, off Point Conception, Sta. 85 (1). 1

Description. Length to 25 mm, width to 4 ram; 17 thoracic segments with parapodia, 11 uncinigerous,

11-14 abdominal segments. Body short, tumid, quickly tapering in abdomen toward small pygidium (Fig.

5.20A). Color in alcohol pale. I

Ul Prostomium much wider than long, indistinctly trilobed, with nuchal organs at postectal corners of

mid-superior lobe. Segments I and 2 separate dorsally, fused laterally and ventrally, produced into lower lip i

and lateral auricular folds. Segment 3 also produced into lateral auricular folds, bearing fin-st2 pairs of gills. 1

Segments 4 and 5 with 1 pair of gills each and vestigial notopodia, those of segment 5 setigerous; notosetae

n

very fine, barely emerging from parapodium (Fig. 5.20B). Branchiae long, slender, subulate (broken in all

examined specimens), separated by roughly triangular branchial membrane; anterior pairs farthest apart, l

Segment 6 with fully developed notopodia; following 11 segments with regular, biramous parapodia. i

Notopodia cylindrical, simple throughout; neuropodia long, narrow pinnules. Abdominal neuropodia II

shorter, with less uncini, but more prominent; notopodial rudiments conspicuous, distally inflated, coiled 1

inwards (Fig. 5.20C). Notosetae in dense fascicles, stiff and bristly, narrowly limbate; in juveniles superior

thoracic uncini with small apical crest of a few small teeth over double row of 4 large teeth and rounded 1

basal prow; inferior thoracic and all abdominal uncini with crest of several teeth of varying size over basal i

i

prow; adults with thoracic uncini bearing single row of 5 teeth, sometimes slightly offset, and crested

abdominal uncini similar to those of juveniles (Fig. 5.20D-F).

Pygidium very small, with 2 lateral flaps about as large as last neuropodia, i

Methyl Green Staining Pattern. Dorsal stain on anterior margins of prostomium and first 2

segments; branchial membrane; transverse bands across segments 3 (in front of branchiae), 4, and 5; all •

notopodial bases including abdominal rudiments. Ventral stain: anterior margin of lower lip, segment 3, and 1

distinct scutes on all remaining setigers.

Remarks. The species, originally described as Amage, was reassigned by Williams (1987) to 1

Paramage because of the presence of rudimentary notopodia in the anterior thorax. With regard to

cephalization, Paramage is intermediate between Mexamage (all branchiae on successive segments, 3 pairs

of rudimentary thoracic notopodia, segments 3 through 6 separate) and Amage (branchiae arranged as in ll

Paramage, but rudimentary thoracic notopodia absent, segments 3 and 4 fused).

There appears to be some confusion as to how many abdominal segments this species has. Moore

(1923), in his original description, reported 10 segments, and Hartman (1969) repeated that statement in her i

description, but had apparently planned to position the species in her key as the counterpart ofA. arieticornuta,

which is characterized by 11 abdominal segments (there is only one entry in her key (p. 522) for the last

couplet). Williams (1987) reported 11 abdominal segments on the type specimens, but identified one of the II

examined specimens (Phase II) as P. scutata that has 10 abdominal segments. If indeed the number of []

abdominal segments is variable in P. scutata, then it is not a useful character for species discrimination in

this genus. It seems, at this point, unjustifiable to erect a new species of Paramage based solely on the 1

number of abdominal segments.

Type Locality and Type Specimens. California, Monterey Bay, off Point Pifios Lighthouse: holotype

(USNM 17209) and 2 paratypes (ANSP 3021). I

142 I
I

Figure 5.20. Paramagescutata: A, entire animal,dorsal view; B, anteriorsegmentys,lateralview; C,lastthoracic

andfirst abdominalsegments,lateral view;D, superiorthoracic uncinus,frontal view; E-F, abdominal

I uncird,2 different views.

I Habitat. Two of the three examined specimens were found in sediments consisting of more than

70% sand; the third one was from a station with 80% silt. Both Moore (1923) and Hartman (1969) report

I muddy sediments as this species' habitat. Distribution. California, 73 to 1229 m.

I Genus Pseudampharete Hilbig, new genus

Type Species. Lysippe mexicana Fauchald, 1972.

I Diagnosis. Four pairs of branchiae, arising from 1 visible segment in 2 closely set rows of 2 pairs

(2+2), outer pair of each row smooth, inner pairs ridged because of encircling ciliary bands. Prostomium

I without uncinigerous glandu(p+3), lar ridges 11 or; buccal 12 abdominal tentaclessegments; smooth. Pabdominal aleae presenotopodial nt; 16 thoracic rudiments setigersabsent. including paleae, 12

Remarks. Pseudampharete belongs to a group of some six genera that have four pairs of branchiae,

I modifications 12 thoracic uncinigers, in one orandmorepaleae; thoracicit issetigers, most closely but differs relatedfromto Athese mpharete generaandinGna having thampha smooth rete rather in lacking than

pinnate buccal tentacles (the buccal tentacles may be entirely absent in Gnathampharete) and in the number

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i

of notopodia on segments 3 to 6 (4 in Pseudampharete, 3 in Ampharete and Gnathampharete). While •

attempts should be made to emend generic def-mitions to avoid an increase in monotypic genera within the

Ampharetidae, these two characters are so consistent in Ampharete that a new genus seems to be justified. _l

Etymology. The name reflects the morphological similarity of the new genus to Ampharete.

Pseudampharete mexicana (Fauchald, 1972), new combination i

Figure 5.21

ill

Lysippe mexicana Fauchald, 1972:299-301, pl. 61, figs. b-d. 1

Ampharetidae gen. Asp. A Lissner et al., 1986:D-17.---Steinhauer and Imamura, 1990:D-1.

Material Examined. California: Santa Maria Basin, off Point Buchon, Sta. 16 (1); off Point San l[

Luis, Sta. 28 (1); off Purisima Point, Sta. R-4 (2).--Western Mexico, vicinity of Cedros Island: holotype

(LACM-AHF Poly 1015). I

I1

Description. Length to 8 nma, width to 1 mm; 16 thoracic setigers including paleae, 12uncinigerous,

and 11 or 12 abdominal segments. Body slender, widest in anterior thorax, with very distinct ventral shields Hi

through unciniger 7 (Fig. 5.21A), gradually tapering toward pygidium. Color in alcohol pale with golden to II

brown pigment over nuchal organs and blackish pigment spots at bases of anal cirri.

m

Prostomium roughly pentagonal, trilobed, with mid-superior lobe sometimes incised, narrower than III

inferior lobe; with 2 eyes approximately at postectal comers of mid-superior lobe and darkly pigmented, I

crescent-shaped nuchal organs along posterior margin of inferior lobe (Fig. 5.21B). Segments 1and 2 fused,

dorsally concealed by branchial bases, ventrally produced into slightly inflated, distinctly crenulated lower am

lip, projecting beyond prostomium if mouth opened, visible in dorsal view (Fig. 5.21A, B). Buccal tentacles l_

smooth, wide. Segment 3 with small fascicles of long, but very fine paleae and branchiae arranged in 2

closely set rows; branchiae slender, long, 2 on each side smooth and 2 slightly wider and appearing ridged 1

because of encircling ciliary bands (Fig. 5.21A, C, D), especially when contracted. Segments 4-6 with small ll

parapodia bearing short, thin setae.

I

Fully developed biramous parapodia from segment 7; notopodia long, cylindrical, neuropodia II

inconspicuous, simple, short pinnules; setae broadly limbate, uncini with 5 rows of 3-4 minute teeth above I

basal prow, similar in thorax and abdomen (Fig. 5.21E, F). Abdominal neuropodia gradually lengthening,

crowded in last 4 or so segments, j

Pygidium very small, with 2 filiform lateral anal cirri about as long as last 2 segments, bearing 1

black basal pigment spot (Fig. 5.21G).

Methyl Green Staining Pattern. Dorsal stain on small band just in front of branchial bases; ventral 1

stain along anterior edge of lower lip (weak); very distinct shields from segment 3 through 11 (unciniger 7),

narrow band on segment 6.

Remarks. A fh-st examination of specimens labeled "Ampharetidae gen. Asp. A" from the Phase I 1

and 1I collections suggested that these specimens did indeed represent a new genus and species, although I1

the new species appeared to be very similar to Lysippe mexicana Fauchald, 1972; the only differences,

according to Fauchald's (1972) description, were the crenulated rather than smooth lip and 11-12 rather I

than 13 abdominal segments in Ampharetidae gen. A sp. A. Subsequent examination of the holotype of L.

mexicana, however, revealed that Ampharetidae gen. Asp. A is the same asL. mexicana; it was found that

the lower lip is crenulated, and the number of abdominal segments is 11. Establishment of a new genus was

still warranted because L. mexicana has 12 rather than 13 thoracic uncinigers. U

l

,44 |
I

Figure 5.21. Pseudampharete mexicana: A, anterior end, ventral view; B, same, dorsal view; C-D, tips of smooth

i and annulated branchiae; E-F, uncini, anteror and lateral views; G, posterior end, lateral view.

I

i 145
!

I

Habitat. The specimens were found in different types of mixed sediment with equal parts of sand •

and silt-clay to more than 98% silt and clay. Two specimens were collected on the continental shelf (less

than 100 m), but the species seems to be more abundant on the upper slope (about 600-2500 m); all of []

Fauchald's (1972) records are from that depth range. I

Distribution. Western Mexico and California, 91 to 2500 m.

Genus Samytha Malmgren, 1866 i

Type Species. Sabellides sexcirrata Sars, 1856. I

l

Diagnosis. Three pairs of smooth branchiae on 1 visible segment, with 1 pair slightly behind the

other 2 (2+1). Prostomium without glandular ridges; buccal tentacles smooth. Paleae absent; 17 thoracic •

setigers, 14 uncinigerous; up to 19 abdominal segments, abdominal notopodial rudiments present or absent. l

Remarks. This genus consists of seven valid species that are scattered over the world's oceans.

Samytha californiensis is the only Californian species; it is included in Hartman's (1969) Arias of Polychaetous •

Annelids, and it was collected in low abundances during the Phase I and II programs.

m

Samytha californiensis Hartman, 1969 iu

Figure 5.22

m

Samytha californiensis Hartman, 1969:573, figs. 1-3.--Banse, 1979:1549-1550.--Lissner et al., 1986:D- n

17.---Steinhauer and Imamura, 1990:F-1.

Samytha hr. californiensis Lissner et al., 1986:D-17. m1

Material Examined. California: Santa Maria Basin, off Pt. San Luis, Sta. 22 (1), Sta. R-1 (1)

Description. Length to 43 mm, width to 7 mm; 17 thoracic setigers, 14 uncinigerous, 16-19 abdominal 1

segments. Body slender, with thorax much longer than abdomen, gradually tapering toward pygidium (Fig. U

5.22A). Color in alcohol pale.

n

large, trilobed, without eyes, nuchal organs, or glandular ridges. Segments I and 2 i

Prostomium

fused, concealed by branchial bases dorsally, forming large, somewhat inflated, finely crenulated lower lip

um

ventrally. Segment 3 with slender branchiae in nearly straight row, 1 pair slightly behind remaining 2, i

separated by membrane about as wide as branchial bases; segment 4 with minute first notopodia bearing n

few very fine, short setae. Segments 5 and 6 with gradually increasing notopodia and notosetae.

n

First fully developed biramous parapodia in segment 7; notopodia simple, cylindrical throughout, i

neuropodia short, narrow pinnules barely emerging from lateral body wall; abdominal neuropodia somewhat |

longer, with tiny notopodial rudiments, at least in anterior segments, very crowded. Notosetae limbate;

thoracic uncini with single row of 5 or 6 teeth over basal prow (Fig. 5.22B); abdominal uncini crested, with n

2 or 3 rows of 5 small teeth over basal prow (Fig. 5.22C, D). n

Pygidium with 2 rounded lateral flaps and several small papillae (Fig. 5.22E).

Methyl Green Staining Pattern. Stain generally weak; no dorsal stain except for prostomium; i

ventrally lower lip and all segments through unciniger 8, stain fading after that.

Remarks. The specimen originally identified as Samytha. nr. californiensis was thought to differ

from the stem species in the number of abdominal segments, but there seems to be some variability in this I

character. Hartman (1969) reported "about 20" for S. californiensis, whereas Banse (1979) found only 19 I

during a reexamination of the type; the Phase II specimen has 17, andS. nr. californiensis has 16. Additionally,

the anal papillae and crenulations thought to be diagnostic for S. californiensis (one of the few features 1

146 I
!

I

I

I

I

I

I

I

i

Figure 5.2.2. Samytha californiensis: A, entire animal, lateral view; B, thoracic unciniger, lateral view; C-D,

I abdominal Hartman). uncinigers, frontal and lateral views; E, posterior end, ventrolateral view. (B, E after

i Hartman (1969) illustrated in the original description) are not discernible in the specimen named S. nr.

californiensis. However, the specimen is damaged, with the lower lip appearing cushion-like because the

i layers ofTyp integument were tom apart, and details like small papillae are likely to be obscured. e Locality and Type Specimens. California, San Pedro Channel: holotype (LACM-AHF).

Habitat. In silt and muddy clay.

i Distribution. Central and southern California, 110 to 370 m.

i Genus Sosane Malmgren, 1866.

Type Species. Sosane sulcata Malmgren, 1866

' _ Diagnosis. Four pairs of smooth branchiae on 1 visible segment, arranged in transverse row.

Ill Prostomium without glandular ridges; buccal tentacles smooth. Paleae present; 16 thoracic setigers including

i paleae, 12 uncinigerous (p+3), third-to-last (unciniger 10) elevated, structurally modified, with hirsute setae;

i 12 or 13 abdominal segments, abdominal notopodial rudiments present.

Remarks. Sosane is a small genus with five valid species; the distribution tends to be concentrated

in warm, subtropical to tropical waters. Only the type species was collected in the boreal North Atlantic.

I Although specimens of Sosane are rare, at least in samples from California, they are easily spotted because of the large, elevated fascicles of notosetae in the modified setiger. The only genus it could be confused with

is Sosanopsis (see below); the two genera differ in the presence of paleae in Sosane and their absence in

i Sosanopsis.
!

1

Sosane occidentalis (Hartman, 1969) g

Figure 5.23 i

Anobothrus occidentalis Hartman, 1969:555, figs. 1-3.

Sosane occidentalis: Williams, 1987:251-252, fig. 1B.--Steinhauer and Imamura, 1990:F-1. i

IB

Material Examined. California: Santa Maria Basin, off Point Sal, Sta. PJ-17 (1).

Description. Length to 19 mm, width to 2 mm; 16 thoracic setigers, 12 uncinigerous, and 12 1

abdominal segments. Body gracile, with relatively wide, square thorax and slender, tapering abdomen (Fig.

5.23A). Color in alcohol pale with pigmented anal cirri.

Prostomium trilobed, nearly quadrangular, with 2 rows of eyespots across posterior half and along 1

anterior margin. Segments 1 and 2 concealed by branchial bases dorsally, separate ventrally, with segment

I forming smooth lower lip and segment 2 narrow, simple ring. Segment 3 with very small fascicles of fine,

short paleae and branchiae, arranged in nearly continuous transverse row with practically no median gap; i

each branchia short and subulate. Segments 4, 5, and 6 with small, gradually increasing notopodia bearing

free notosetae.

Segments 7 through 15 with regular biramous parapodia; notopodia papilliform, neuropodia short, 1

inconspicuous pinnules. Notosetae stiff, limbate (Fig. 5.23B); thoracic and abdominal uncini similar, crested,

1B

with 4 rows of 4-5 teeth above inconspicuous rostral point and small basal prow (Fig. 5.23C-E). Segment 16

(third-to-last setiger) with dorsally displaced, enlarged notopodia bearing dense fascicles of modified, i

flattened, distally hirsute setae reminiscent of modified setae of Mugga (see above) (Fig. 5.23F). Notopodia II

connected by indistinct dorsal ridge (Fig. 5.23A, G). Segments 17 and 18 with regular noto- and neuropodia.

Abdominal neuropodia gradually lengthening toward posterior end, last pair reaching margin of pygidium, i

III

Pygidium small, with 2 relatively large, digitiform lateral anal cirri, each with large aggregation of

pigment granules. l

Methyl Green Staining Pattern. Dorsal stain on prostomium, segments 1 and 2, and front of 1

branchial membrane; ventral stain on entire thorax, with distinct ventral scutes through setiger 10, last 2

setigers with fading stain; branchiae unstained. •

Remarks. Sosane occidentalis is a small species that tends to occur in low abundance, and it may !

be difficult to spot in a dish under low magnification. The species is best recognized by the dense fascicles

of modified notosetae; it can be distinguished from juvenile Anobothrus by the position of the modified /I

setiger (third-to-last in Sosane, fourth- or fifth-to-last in Anobothrus) and the development of the dorsal II

ridge connecting the modified parapodia (barely visible in Sosane, distinct in Anobothrus). Hartman (1969)

erroneously stated in her original description ofS. occidentalis (asAnobothrus occidentalis) that 11 thoracic •

uncinigers are present; the error was corrected by Williams (1987). See also remarks onAnobothrus gracilis. 1

Type Locality and Type Specimens. California: off Santa Catalina Island: holotype (LACM-AHF

Poly 0263) and 1 paratype (LACM-AHF Poly 0264). i

i Habitat. Occurs in silty sand.

Distribution. Central and southern California, 76 to 257 m. _,

I

!

148 I
I Figure 5.23. Sosaneoccidentalis:A, entireanimal, ventrolateralview; B,regular notoseta,tip; C,thoracic uncinus,

frontal view; D-E, abdominaluncini, 2 different views;F, modified seta,tip; G, setiger 13, dorsum,

i anteriorview.

I Genus Sosanopsis Hessle, 1917. Type Species. Sosanopsis wireni Hessle, 1917

I segment Diagnosis. 5 (3+1). Prostomium Four pairs without of branchiae, glandular3 pairs ridges; arising buccalfromtentacles segment smooth. 3 andPalea fourthe absent; pair arising 15 thoracic from

setigers, 12 uncinigerous; third-to-last setiger (unciniger 10) with elevated, enlarged notopodia bearing

i modified setae; occasionally dorsal ridges present on posterior setiger(s); 11-14 abdominal segments. Remarks. As the name indicates, Sosanopsis is closely related to Sosane, but differs from the latter

in lacking, rather than having paleae. The arrangement of the gills is very unusual in that an abranchiate

I segment divides the anterior and posterior pairs.

There are only three valid species in this genus (Holthe, 1986a), none of which are included in

Hartman's (1969) Atlas of Polychaetous Annelids. This record may be the first of Sosanopsis wireni for

I California.

I 149
I

Sosanopsis wireni Hessle, 1917 •

Figure 5.24

i

Sosanopsis wireni Hessle, 1917:112-113, fig. 17a-d, pl. 1, fig. 6.--Holthe, 1986b:228.--Maciolek et al.,

Ampharetidae 1987:D-1. genus B Ispecies A Lissner et al., 1986:D-17.

Material Examined. California: Santa Maria Basin, off Purisima Point, Sta. 56 (1). I

Description. Length to 8 mm, width less than 1 mm; 15 thoracic setigers, 12 uncinigerous, 11 I

abdominal segments. Body slender, widest in anterior thorax, gradually tapering toward posterior end.

Color in alcohol pale.

Prostomium trilobed, pentagonal, without eyes or nuchal organs. Segments 1 and 2 fused dorsally, i

free ventrally, with segment 1 forming scalloped to coarsely crenulated lower lip; buccal tentacles smooth.

Segment 3 with 3 anterior pairs of branchiae in straight row, separated by wide median gap; segment 4 fused U

to segment 3, bearing small notopodia with reduced setal fascicle; segment 5 with last pair of branchiae l

above notopodia (Fig. 5.24A).

Regular biramous parapodia on segments 7 through 15 (uncinigers 1 through 9); notopodia simple, n

cylindrical, neuropodia short pinnules. Third-to-last thoracic segment (unciniger 10) with dorsally displaced, I

enlarged notopodia and dense fascicles of slightly modified setae; last 2 setigers normal. Abdominal

neuropodia much longer than thoracic, markedly increasing toward posterior end. Regular notosetae limbate, I

modified setae simple, weakly hirsute (Fig. 5.24B, C). Both thoracic and abdominal uncini crested, abdominal I

uncini slightly smaller than thoracic (Fig. 5.24D, E).

I

o I

I

A .........-:::i"::":@:-i.-.: :_ '_7 ,_-,.,-_,_,. ." :':i"iii-_."_i;_}i'_::,?

C B

'

I

Figure 5.24. Sosanopsiswireni: A, anteriorend,lateral view;arrow:insertionof posteriormostbranchia;B, regular

unpublC,ished modiseta; fiedsketcR.E.D-E' hSeta;byRuff).th°rac and iacbdominal uncini, fi-ontaland lateral views. (after an I

I
!

I Pygidium short, produced into 2 rounded lateral lobes, each bearing short, digitiform anal cirrus,

easily breaking away.

I rounded Mpatches ethyl Green behind Staini outer nanterior g Patternand. posterior Dorsal stain branchial on prost bases; omiuventral m, fusedstainsegments on all segments 1 and 2,through and 2

setiger 9.

I Remarks. The examined specimen is in poor condition; none of the branchiae are still attached, but

the scars could be seen with methyl green stain.

Type Locality and Type Specimens. Swedish west coast, Kosterfjord: 3 types (Zoological Museum

Uppsala).

Habitat. In mixed sand and silt.

I Distribution. Northern Europe; U.S. Atlantic; California; 50 to 1220 m.

I Subfamily Melinninae

Genus Melinna Malmgren, 1866

I Type Species. Sabellides cristata Sars, 1851.

Diagnosis. Four pairs of usually smooth branchiae on 2 visible segments, arranged 2+2 or 3+1. One

I pair of nuchal hooks present; dorsal crest on segment 6 present. BuccaI tentacles all similar. Twenty thoracic segments, 17 or 18 setigerous, 14 uncinigerous. Abdominal segments numerous, varying between 50 and 70.

Remarks. Melinna is one of the larger ampharetid genera with 19 valid species, widely distributed

I in the world's oceans. Both species found in the Phase I and II materials were also listed by Hartman (1969).

I Melinna heterodonta Moore, 1923 Figure 5.25

I Melinna cristata heterodonta Moore, 1923:212-213, 17, 25. pL fig.

Melinnaheterodonta: Hartman, 1960:157; 1969:565-566, figs. 1-6.--Lissneretal., 1986:D-17.--Steinhauer

i and Imamura, 1990:F-1. Material Examined. California: Santa Mafia Basin, off Point Sol, Sta. R-9 (1), Sta. PJ-23 (1); off

Point Arguello, Sta. 76 (6).

I Description. Length to 90 mm, width to 3.5 ram; 18 thoracic segments, 15 setigerous, maximally

54 abdominal segments. Body long and slender, widest atlevel of postbranchial membrane, gradually tapering

toward pygidium (Fig. 5.25A). Color in alcohol pale.

I Prostomium slightly wider than long, with scalloped anterior margin, raised Ixansverse glandular

ridge posteriorly, and paired nuchal organs just anterior to ridge ("lateral pads" of Moore, 1923). Segments

I 1stout, and 2largest fused toonesprostomium about as wide dorsall asybranchiae., produced into lower lip ventrally. Buccal tentacles smooth, slender to

Segments 3 to 6 fused dorsally and laterally, forming lateral wings with 4 rows of needlelike

I neurosetae edge. Segments (Fig. 5.25 5 andB); 6segments separate, 3with and 4ill-de fusedfined ventrall glandular y, forminscutes. g deepSegment ventral fo3ldbearing with slightly dorsalcrerecurred nulated

hooks and branchiae, arranged I+3 or 2+2, fused basally, long and subulate; segment 6 with first notosetae,

I very membrane fine andwithshort, 11 toemerging 16 teeth,more the 4ormedian lss directly ones coarsest from body(Fig. wall, 5.25noC).visible notopodia; and postbranchial

i 151
152 I
I First unmodified parapodia on segment 7, with fully developed, cylindrical notopodia and neuropodial

pinnules bearing uncini. Notosetae stiff, brassy, broadly limbate; thoracic uncini with 1row of 3 teeth above

i large rostral point and rounded basal prow; abdominal uncini with 2 large teeth surmounted by crest of about 7 smaller ones, large rostral point, and rounded basal prow (Fig. 5.25D-H). Notopodial rudiments

present in anterior abdominal segments (Fig. 5.25A).

I scalloped.Pygidium very small, with terminal anus surrounded by 2 lateral pads; dorsal margin sometimes

Methyl Green Staining Pattern. Prostomium with anterior edge and raised glandular ridge staining

I dark band green; across dorsum segmentsof segments 7 and 8. Ventrum 1-5 stainingstaining weakly;solidgillsthrough and postbranchial segment 6,membrane pronouncedwhite; ventralbroadscutes greenin

segments 7-16 (uncinigers 1-9), then starting to fade; no stain or weak anterior band on f'u-stthoracic segment.

I Body wall staining just above and between notopodia. Remarks. Melinna heterodonta is very similar to M. cristata and was originally described as its

subspecies. The two species differ in the number of setigerous segments (15 in M. heterodonta, 16 in M.

I cristata) and the dentition of the postbranchial membrane (median teeth distinctly larger than others in M.

heterodonta, all teeth equal in M. cristata).

Type Locality and Type Specimens. California, off Point Pifios Lighthouse: holotype (USNM

I 17331) and 15 paratypes (USNM 16991); 11 paratypes (ANSP 3129).

Habitat. Moore found most of his specimens in Green mud; the present material is from localities

i with 74%Disilt. stribution. Central and southern California, 198 to 1528 m.

I Melinna oculata Hartman, 1969

Figure 5.26

I Melinna oculata Hartman, 1969:567-568, figs. 1-6.--Lissner et al., 1986:F- 17.---Steinhauer and Imamura,

1990:F-1.

I Material Examined. California: Santa Mafia Basin, off Purisima Point, Sta. R-4 (3); off Point Arguello, Sta. 73 (1). Western Santa Barbara Channel, off Point Conception, Sta. 79 (1).

i Description. Length to 40 mm, width to 2.5 mm; 18 thoracic segments, 16 setigerous, and 56 abdominal segments. Body slender, almost linear, gently tapering toward pygidium (Fig. 5.26A). Color in

life drabgreen, branchiae with browntransverse pigment bands; color in alcohol pale with striped branchiae,

pigment fading over time.

I Prostomium wider than long, with scalloped anterior margin; posterior edge developed into raised

transverse glandular ridge bearing 2 parallel rows of eyespots (Fig. 5.26A, B). Segments 1 and 2 fused to

prostomium dorsally, forming smooth, padlike lower lip ventrally. Buccal tentacles smooth, of 2 sizes, few

I large ones about the size of branchiae.

Segments 3-6 fused, laterally developed into wings with 3 rows of needlelike neurosetae. Segment

I 3+1, withbasally dorsal fused recurved(Fig.hooks 5.26C). andSegment branchiae,4 with ventrally neurosetae formingonly; collarlike segmentfold. 5 with Branchiae neurosetae short,andarrange smalld

fascicle of very f'me notosetae; segment 6 with small notosetae and short postbranchial membrane with

I about 10 coarse dentations (Fig. 5.26C).

!

i 153
I

I

I

, I

i

.... "'" I

I

•... .i:_":

_'_ i

I

_I__ _ i

,

Figure 5.26. Melinna oculata: A, entire animal, lateral view; B, anterior end, dorsal view, branchiae pulled back

to show prostomium; C, same, branchiae in natural position; D-E, thoracic uncini, lateral and frontal •

views; F, uncinus from middle of abdomen, frontal view; G-H, uncini from far posterior segment, U

lateral and frontal views; I, posterior end, ventrolateral view.

I

First unmodified parapodia in segment 7, with well-developed notopodia and neumpodial pinnules

bearing uncini. Thoracic neuropodia short and wide, with numerous very crowded uncini, gradually l

lengthening in last setigers, becoming narrower but more conspicuous in abdomen. Notosetae stiff, limbate; |

uncini in thorax and anterior abdomen with single row of 4 teeth over basal prow (Fig. 5.26D, E); in middle

of abdomen with crest of 6 small teeth above large main fang and basal prow (Fig. 5.26F); and in far m

posterior segments crested with 3 rows of 3 slender teeth (Fig. 5.26G, H). |

Pygidium small, with wide terminal anus surrounded by 2, sometimes pigmented, dorsolateral flanges

and additional ventral crenutations (Fig. 5.261). i

154 I
I Remarks. The pigment stripes on the gills, a very characteristic and easily recognizable diagnostic

feature for this species, may fade in alcohol over time; however, M. oculata is readily distinguished from its

I co-occurring congener, M. heterodonta by the much shorter postbranchiat membrane, the eyes on the prostomium, and the distribution of neuro- and notosetae in segments 3-6: M. oculata has neurosetae in

segments 3 and 4, both neuro- and notosetae in segment 5, and notosetae only in segment 6; whereas M.

i heterodonta has neurosetae in segments 3 and 4 and both neuro- and notosetae in segments 5 and 6 (the notosetae in segment 5 are not always easy to see).

Type Locality and Type Specimens. California: Palos Verdes shelf: holotype (AHF? write to

I Leslie) Habitat. Hartman (1969) reported mixed sediment as the habitat of M. oculata; the examined

specimens are from locations with mostly silt or silt and sand.

I Distribution. Central and southern California, 22 to 450 m.

I Literature Cited

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I Zoology 57: 1543-1552.

Berkeley, E. and C. Berkeley. 1942. North Pacific Polychaeta, chiefly from the west coast of Vancouver

I Island, Alaska, and Bering Sea. Canadian Journal of Research 20:183-208.

Berkeley, E. and C. Berkeley. 1943. Biological and oceanographical conditions in Hudson Bay. 11. Polychaeta

from Hudson Bay. Journal of the Fisheries Research Board of Canada 6: 129-132.

I Berkeley, E. and C. Berkeley. 1952. Polychaeta sedentaria. Canadian Pacific Fauna 9b(2): 1-139.

Berkeley, E. and C. Berkeley. 1956. On a collection of polychaetous annelids from Northern Banks Island,

I Beaufort Sea, and northwest Alaska; together some new

from the south from with recordsfrom the

east coast of Canada. Journal of the Fisheries Research Board of Canada 13: 233-246.

I Blake, J.A. Light's 1975Manual: . PhylumInter Annelida: tidal Invertebrates Class Polychaeta. of the Pp. Central 151-243 California In: R.I.Coast. SmithUniversity and T.J. Car ofltCalifornia on (eds.),

Press, Berkeley.

I J.H. 196I. The fauna of South Africa. Part 6. off Day, Polychaet [sic] Sedentary species dredged Cape

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I Day, J.H. 1964. A review of the family Ampharetidae (Polychaeta). Annals of the South African Museum

48: 97-120.

I Day, J.H. 1967. A monograph of the Polychaeta of Southern Africa. Part 2. Sedentaria. British Museum (NaturalHistory) London, Publication No. 656, xvii + pp. 459-878.

I Eliason, Vetenskaps- A. 1955. Neueochoder Vitterhets wenig bekannte Samhalles;schwedisc Handlingar, he Ampharetide Serien B, 6:n 1-17. (Polychaeta). Grteborgs Konglia

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.

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.

i 155
Fauchald, K. and D.R. Hancock. 1981. Deep-water polychaetes from a transect off Oregon. Alan Hancock •

Monographs in Marine Biology 11: 1-73.

Fauchald, K. and P.A. Jumars. 1979. The diet of worms: a study of polychaete feeding guilds. Oceanography D

and Marine Biology Annual Review 17: 193-284. i

Fauvel, P. 1897. Recherches sur les Amphar_tiens. Bulletin Scientifique de la France et de la Belgique 30: •

t-212, pls. XV-XXV. II

Gibbs, P.E. and K. Probert. 1973. Notes on Gyptis capensis and Sosane sulcata (Annelida: Polychaeta) from

the benthos off the south coast of Cornwall. Journal of the Marine Biological Association of the •

United Kingdom 53: 397-401. I

Grehan, A.J. 1991. Demography and reproductive biology ofMelinnapalmata (Ampharetidae: Polychaeta) •

in Inner Galway Bay on the west coast of Ireland. Marine Biology 109: 459-467. II

Grube, A.-E. 1860. Beschreibung neuer oder wenig bekannter Anneliden. Archiv fiJrNaturgeschichte Berlin i

26: 71-118, pls. m-V. i

Hartman, O. 1960. Systematic account of some marine invertebrate animals from the deep basins of southern

California. Allan Hancock Pacific Expeditions 22: 69-216. •

Hartman, O. 1961. Polychaetous annelids from California. Allan Hancock Pacific Expeditions 25: 1-226.

Hartman, O. 1969. Atlas of the Sedentariate Polychaetous Annelids from California. Allan Hancock, •

Foundation, Los Angeles. 812 pp. I

Hartmann-Schr6der, G. 1971. Anne•ida, Borstenwiirmer, Polychaeta. Die Tierwelt Deutschlands 58: 1-594. i

Hessle, C. 1917. Zur Kermtnis der terebellomorphen Polychaeten. Zoologiska Bidrag from Uppsala 5: 39- i

258, pls. I-V.

Holthe, T. 1986a. Evolution, systematics, and distribution of the Polychaeta Terebellomorpha, with a catalogue i

of the taxa and a bibliography. Gunneria 55: 1-236. m

Holthe, T. 1986b. Polychaeta Terebellomorpha from the northern Norwegian Sea and the Polar Sea, with •

descriptions of Mugga bathyalis sp.n. and Ymerana pteropoda gen. and sp.n. Sarsia 71: 227-234. I

Hutchings, P.A. 1973a. Gametogenesis in a Northumberland population of the polychaete Melinna cristata.

Marine Biology 18:199-211. i

Hutchings, P.A. 1973b. Age structure and spawning of a Northumberland population of Melinna cristata

(Polychaeta: Ampharetidae). Marine Biology 18: 218-227. I

Johnson, H.P. 1901. The Polychaeta of the Puget sound region. Proceedings of the Boston Society of Natural m

History 29(18): 381-437.

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s6rie D, 277: 2723-2725. •

Levinsen, G.M.R. 1883. Systematisk-geografisk Oversigt over de nordiske Armulata, Gephyrea, Chaetognathi

og Balanoglossi. 2. Videnskabelige Meddelser fr_ Dansk Naturhistorisk Forening I KObenhavn

1883: 92-350. I

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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 Mafia Basin and

i Western Minerals Santa Management Barbara Service, Channel Pacific - Phase OCS I. Report Region, prepared underforcontract the U.S. No.Department 14-12-0001-30032. of the Interior,

Maciolek, N., J.F. Gmssle, B. Hecker, B. Brown, J.A. Blake, P.D. Boehm, R. Petrecca, S. Duff-y,E. Baptiste,

I and R.E. Ruff. 1987. Study of Biological Processes on the U.S. North Atlantic Slope and Rise. Final Report prepared for the U.S. Department of the Interior, Minerals Management Service, Washington,

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I Malmgren, A.J. 1866. Nordiska Hafs-Annulater. 0fversigt af Kongliga Vetenskaps-Akademien Forhandlingar

22: 355-410.

I Mafinescu, Danube: V.P. Hypaniola 1964. La r6production et le d6veloppement des polych_tes reliques Ponto-Caspiens du kowalewskii (Grimm) et Manayunkia caspica. Revue Roumaine de Biologic,

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Transactions of the American Microscopical Society 93: 78-89.

Zottoli, R.A.75 pp.MS"Ampharetid polychaetes of the northeastem United States, shallow and deepwater species. I

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158 I
Appendix

Lists and Maps of Stations

Table A.I. 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°10.20"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°20.00'N 121°06.58"W 308

9 35°19.48'N 121°10.06"W 398

10 35°18.28'N 12t°18.65'W 591

11 35°17.80'N 121°22.13"W 690

12 35°15.03'N 120°57.31'W 98

13 35°14.54"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 t21°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

2 t 35006.1 I'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°56.8 I'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 121°15.37'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°00.81'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 120052.92"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 34045. ! I'N 120°52.85"W 196

49 34045.03 'N 120056.31 '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 120056.3 I'W 590

56 34°30.32'N 121°01.02"W 900

159
!

Table A.1 (Continued)

Station Latitude Longitude Depth l

(m) II

58 34°34.35_N 120°45. t8%V 99

B

59 34°33.65_N 120°47.18_/ 216

60 34°33.25'N 120°48.34_V 275

61 34°33.01'N 120°48.89_W 345

62 34°30.46"N 120°52.13"W 582 •

l

63 34°26.29'N 120°58.08"W 930

64 34°33. t5'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°54.2tYW 927 n

70 34°29.67'N 120°43.70'W 200 •

71 34°29.04'N 120°44.0 I'W 306 U

72 34°28.4 I'N 120°44.76'W 40 l

73 34°28.2 I'N 120°36.80'W 98 •

|

74 34°26.84'N 120°38.6 I'W 20 l

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 •

II 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 n

81 34°2t.26"N 120°28.83'W 294 •

82 34° 18.7 I'N 120°29.55'W 394 U

83 34° 17.20"N 120°30.20NV 444

84 34° 13.54"N 120°31. l 9'W 394 n

85 34%5.88'N 120°16.31'W 113 •

86 34°24.45 I 'N 120° 17.02"W 197

87 34°21.60"N 120° 17.11'W 299

88 34° 17.89"N 120° 16.86'W 393 •

|

89 34°13.79'N 120°16.56'W 471

90 34°09.44'N 120° 16.30'W 375

91 34° 11.73'5/ 120°07.43"W 540

92 34°08.70'N 120°07.5tYW 444 •

93 34°07.63'N 120°07.5 I'W 357 1

96 34°22.9 I'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 i

100 34°08.67"N 120°05.50'W 443 •

101 34°07.5 J t'N 120°05.65"W 357

102 34°59.7l'N 120°48.22'W 99

103 34°59.63"N 120°53.56'W 197 •

|

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 t21°15.08'W 573 •

l

108 34°58.21'N 121°17.88'W 492

Note: Sample labels from the Soft-substrate 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 = Q

Benthic Sediment Variance (subsamples); 01-09 = replicate numbers; TX = a taxonomy sample. Sample labels having the

designation BRA, represents a sample from recks taken as part of the hard bottom survey, m

160 I
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I : _ "_ mESTERO I I

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i ,+ . ,_ I+ _.g • 20 I;_ 12 ( l_ BUCHON

• 14"-.- -._

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it9 - ! o _ e'-" ]

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•

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i "'"-. • ". 52

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'" 55 " i:" 581 64 LPt'AFIGUE/LO

• ....... • ++.+- •

m ...... ":-- -- " + ' "Z J _ . +_

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i • 72 71.......... 41,74 • 85

......... e,so......... 200 . %....

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ii' KEy e _ 92 • 0100

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Figure A.1. Map showing location of soft-substrate stations from the Phase I Reconnaissance and Phase II

Monitoring Programs.

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Table A.2. Location of soft-substrate stations taken during the Phase 1] Monitoring Program. I

Station Latitude Longitude Depth B

(m)

mat

R- l 35_05.83'N 120_49. ]6'W 91 •

R-2 35°05.50'N 120°53.40"0/ 161

R-3 35°05.30'N 121°00.90"W 409

R-4 34°43.01'N 120°47.39'W 92 i

R-5 34°42.69'N 120°50.83"W 154 •

R-6 34°4 t.40rN 120°57.90"W 410 g

R-7 34°52.90'N 121°10.30tW 565

R-8 34°55.30'N 120°45.87"W 90 •

|

R-9 34°53.68"N 120o59.12'W 410

PJ- 1 34°55.79"N 120o49.9 I'W 145

PJ-2 34°55.32'N 120°49.59'W 142

PJ-3 34°56.26'N 120°49.58'W 138 •

PJ-4 34°56.26'N 120°50.24'W 150

PJ-5 34°55.32'N 120°50.24'W 152

PJ-6 34°54.7 I'N 120°49.9 I'W 148

PJ-7 34°55.79'N 120°48.60'W 123 •

PJ-8 34056.87'N 120049.9 I'W 142

PJ-9 34°55.79'N 120051.23'W 169

PJ-10 34°53.63'N 120°49.9t'W 147 I

|

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- 14 34°55.79"N 120°49.26'W 134 •

l

PI- t 5 34°55.79"N 120°50.57'W 155

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 35°01.23'N 120°51.15'W 143

PJ-22 34°55.25'N 120°49.93'W 143 •

PJ-23 34°56.33"N 120°49.90'W 143 It

!

Table A.3. Sampling dates of MMS Phase ITMonitoring Program. i

Cruise Date i

1-1 October 1986

1-2 January 1987

1-3 May 1987 •

2-1 July 1987

2-3 October 1987 |

2-4 January 1988

2-5 May 1988 m

3-I October 1988

3-4 May 1989 |

!

162 R
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°00.443"W 34°24.184'N 120°01.48(YW 73.5-78

2 A/B 34°11.377'N 120°29.318'W 34° 11.289"N 120°28.774'W 110-126

2 C/D 34° I0.984"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°40.189'W 168-237

6 A/B 34°30.246"N 120°35.555"W 54-63

6 C/D 34°30".42 I'N 120_34.315"W 54-63

13 A/B 34°42".570'N 120°47.899'W 34°42.107"N I20°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 CA) 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 120050.197"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°46.470'N 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°50.990'N 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°05.33 I'W 35°27.805'N 121°05.277'W 102-106.5

Table A_. MMS Phase I1 - Locations of hard-substrate photosurvey stations.

Station Latitude Longitude Depth

(m)

PH-E 34°30.26_N 120°42.76'_ 119

PH-F 34°30.8 I'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 1t7

PH-K 34°29.37'N 120°42.26"W 160

PH-N 34°29.21'N 120°42.05'W 166

PH-R 34°29.11'N 120°42.67"W 213

PH-U 34031.48'N 120°43.5 I'W ! 13

PH-W 34°31.52'N 120°45.86'W t95

163
I

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....._ '_:" 6

34o32,N ...._._. J LPt. ARGUELLO

PH-U "_-_ I • _

Pt. CONCEPTION

t ............ " . • 2

PH-W ",, "

/' • - .... .

PH-E • PH-I -_) / ...................... 200 ..............

•,o_o.N _-_ II " ,/ I

PH-.K _l i' /"' /

PH-R PH-N .,

34° 28'N I / 'I

120 °44'W

Figure A.2. Map showing location of hard-substrate stations from the Phase [ Reconnaissance and Phase II

Monitoring Programs. Phase II stations are indicated in the inset, i
I

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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 biological diversity;

i preserving the environmental and cultura! 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 thefr care.

The Department also has a major responsibilityfor American Indian reservation communities

and for people who live in islandterritories under U.S. administration.

The Minerals Management Service Mission

I __'_ As pr a bureau of the Department of the Interior, the Minerals Management Sarvice's (MMS) imary 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

lands, and dLqtribute those revenues.

I Moreover, in working to meet its responsib_litiss,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 reffi esources. cient, timeThely MMS and accurate Royltycollectio Management n andProgra disbursem meent esofitsrevenue responsibil fromitimineral es by ensuring leasing the and

production due to indian tribes and allottees, States and the U.S. Treasury.

I The respons MMSivestrives to the topubli fulfill c's itsconresponsibilities cerns and interethrough sts by the maintagenining eral guiding a dialogueprincwithiplesalof: l potentially (1) being

affected parties and (2) carrying out its programs with an emphasis on working to enhance the

quality of life for aft Americans by lending MMS assistance and expertise to economic

I development and environmental protection.

I