I

l OCS Study

i MMS 96-0041

i

I TAXONOMIC ATLAS OF THE BENTHIC FAUNA OF

THE SANTA MARIA BASIN AND

I WESTERN SANTA BARBARA CHANNEL

I

I FINAL REPORT Volume 14 of 14

l

Miscellaneous Taxa

l

i Edited by: James A. Blake

Paul H. Scott

I ndrew Lissner

Submitted by:

1 Science Applications International Corporation 10260 Campus Point Drive

San Diego, California 92121

I For:

U.S. Department of the Interior Minerals Management Service

Pacific OCS Region

770 Paseo Camarilto

Camarillo, California 93010

Under Contract No. 14-35-0001-30484

I

Minerals Management Service

Pacific OCS Region July 1996

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DISCLAIMER N

TService, his reportU.S. hasDepartment been revieweofdthebytheInterior PacificandOuapproved ter Continenfortapubl l Shel ication. f RegionThe, Minopinions, erals Manfindings, gement I

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

nnot ecess consitiut arily refl e endors ect theevi ment ew ofortherecommendation Minerals Managfemoreusnte.SeThis rvice.report Mentihas on onot f trabdeeennamedited es dofeors I

conformity with Minerals Management Service editorial standards.

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TAXONOMIC DISCLAIMER

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

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

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I PROJECT ORGANIZATION

I

PROGRAMMANAGER

I

DR. ANDREWL. LISSNER

t 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

89 Water Street

i Woods Hole, Massachusetts 02543

i PROJEC'r TECHNICAL OFFICER

I MR.FRANKJ. MANAOO

U.S. Department of the Interior

I Minerals Management Service, Pacific OCS Region

770 Paseo Camarillo

I Camarillo, California 93010

I CONTRIBUTING AUTHORS

MARY BERGEN

I DEBRAHART

GORDONHENDLER

I ANF.G.DREWHOCHBERG LISSFmR

GRETCHENLAMBERT

' PAULA JOHNS.F.WINCHELL Pa._ER

KErrH H. WOODW1CK

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ACCESS NUMBER: 30484.14

TECHNICAL SUMMARY i

STUDYTITLE: TaxonomicAtlas of the Santa Maria Basinand WesternSanta BarbaraChannel. i

REPORTTITLE: TaxonomicAtlasof the Santa Maria BasinandWesternSantaBarbaraChannel.Volume

t4. MiscellaneousTaxa.

IB

CONTRACT NUMBER: MMS ContractNo. 14-35-0001-30484. I

SPONSORINGOCS REGION: Pacific. i

i

APPLICABLEPLANNINGAREA: SouthernCalifornia.

FISCALYEARSOF PROJECT FUNDING: 1990; 1991;1992; 1993;1994; 1995;1996. I

COMPLETIONDATEOF REPORT: July1996.

COSTS: FY 1989,$229,013; FY 1990,$150,000; FY 1991$150,000; FY 1993$150,000;FY 1995$45,371. i

CUMULATIVEPROJECT COST: $724,384.

II PROJECTMANAGER: Andrew L. Lissner.

AFFILIATION: Science ApplicationsInternationalCorporation. i

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

PRINCIPALINVESTIGATORS""JamesA. Blake,AndrewL. Lisener. i

KEYWORDS: Marineinvertebrates,Brachipeda,Echiura,Sipuncula,EchinodermataC, rinoidea,Ophiumidea, i

Asteroidea,Echninoidea,Holothuroidea,Hemicordata, Enteropneusta,Cordata, Urocordata,Ascidacea, I

taxonomy,California,Santa MariaBasin,SantaBarbaraChannel,continentasl helf,

BACKGROUND: The TaxonomicAtlasof theSanta Maria Basinand westernSanta BarbaraChannelisan

extensionofthe benthicreconnaissance(PhaseI)and monitoringprograms(PhaseII)thatwere conducted

bytheMMS since1983. The organismsthatwerecollectedas partofthoseprogramsprovidethe material

onwhichthe Atlasisdeveloped, in ordertofullydocumentthefaunacollectedbythoseprograms,a series i

of 14 volumes will be preparedthat provide keys, descriptions, and illustrationsof the benthic fauna of the II

hardandsoft substrate environments. A team of40 experts on the fauna hasbeen assembledto carry out

this work and their contributions are distributed amongthe 14volumes.

OBJECTIVES: The objectives of Volume 14 are to provide keys, descriptions, and illustrations to the _

miscellaneoustaxa of the Santa Maria Basinand Western Santa BarbaraChannel. This volume treats the

brachiopods,echiurans,sipunculans,crinoids,ophiuroids,asteroids,holothuroids,enteropneusts,and IBm

ascideans.

DESCRIPTION: Volume14includesa treatmentofthe brachiopods,echiurans,sipunculans,echinoderms, I

enteropneusts(hemichordates),and ascideans(urochordates). The eleven chaptersare organizedinto I

sectionsthatincludethe morphology,taxonomichistory,biology,keysofspecies,anddescriptionsofgenera

andspecies. Eachspeciesisfullyillustratedwith relevantcharacteristicslabeledandidentified. !

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ACCESS NUMBER: 30484.14

I SIGNIFICANT CONCLUSIONS: A total of 97 species and 73 genera are includedas representativesof 6

phyla:Brachiopoda(7 genera,9species),Sipuncula(4 generaandspecies),Echiura(4 genera,3 species),

Crinoidea(1 genusandspecies),Asteroidea(8 generaandspecies),Ophiuroidea(14 genera,19species),

i Echinoidea and Holothuroidea 6 28 species), Enteropneusta(4 and

(4genera species), (1 genera, genera

species),and Ascidiacea(12 genera, 16species). Forthe Ophiuroidea,a lectotypewas designatedfor

Amphiuragranulataandthespecieswastransferredtothe genusAmphichondrius. Amhiopfushexacanthus

I wdeasscrib determedininaesdetparat o be eapap syneorn(Lamb ymof eDougafop rt,1993).tusamphacanthus. Three new speciesof ascidianswere

i STUDY RESULTS: The faunaofthe Santa Maria Basinand WesternSanta BarbaraChannel is evaluated fromsamplesthat were taken from depthsof about50 to 1,000 m. Organismswere collectedfromsoft

sedimentsusingboxcoresandfromrocksusingmanipulatorarmsofsubmersiblesand remotelyoperated

vehicles. The collectionsare organizedinto sets of vouchersfrom Phase I and II that havebeen made

availableto the teamof investigators.Additionalmaterialfromthe bulkcollectionsnowarchivedwiththe Los AngelesMuseumof NaturalHistoryisalsobeingexamined.Thetotalnumberofspeciestreatedintheentre

TaxonomicAtlas(14volumes)mayexceed1,000species.

tm

'1 Volume14representstheconclusionof the Atlas. Includedinthisvolumearerepresentativesof 6 different

coelomatephyla:Brachiopoda,Sipuncula,Echiura,Echinodermata,Hemichordata(Enteropneusta),and

i Urochordaa(Ascidiacea).FiveclassesoftheEchinodermatare a includedCdnoid : eaAsteroidea, , Ophiuroidea, Echinoidea,and Holothuroidea.A total of 97 speciesand 73 generaare includedamongthesevarious

coelomatephyladistributedasfollows:Brachiopoda(7 genera,9 species),Sipuncula(4 generaandspecies),

Echiura(4 genera, 3 species), Crinoidea(t genus and species),Asteroidea(8 genera and species),

I Ophiuroidea species),Ent(14eropngenera, eusta(419gspeci eneraeas), ndEchinoid species),eaand(4 Ascidi generaacandea(12spgeeciner es),a,Holot 16speci huroid es).eaEach (16 gener speciae,28sis

• fullydescribedandillustrated,withkeysproducedthatallowusersto identifythefauna. FortheOphiuroidea,

i II a lectotype was designated for Amphiura granulata and the species was transferred to the genus

II Amphichondrius. Amhiopfushexacanthuswas determinedto be a synonymof Dougafopfusamphacanthus.

Three new speciesof ascidians, Motgula napiformis, Boftenia pofyplacoderma,and Styefa tesseris were

I described inseparatepaper(Lambert, 1993:Proc.Calif.Acad. Sci.48).

STUDYPRODUCT: Blake,J.A.. P.H.Scott and A.L. Lissner (Editors).1996. TaxonomicAtlasofthe Benthic

Faunaof the Santa Maria Basin and Western Santa Barbara Channel. Volume 14. MiscellaneousTaxa. A

I fiMinerals nalrepoManag rtpreparement edby SSecirevice, nceAPacific pplicatioOCSnsInt Region, ernationa Camarillo, Col rporati CA.on,fOCS ortheStudy U.S. Dep MMSart96-0041. mentoftheContract Interior,

No. 14-35-0001-30484.

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I "P.l'saffiliationmaybedifferenthanthatlistedfor ProjectManager(s).

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ACCESS NUMBER: 30484.14

I_" i_ lip II?* I

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S0:_S2- _01

REPORTDOCUMENTATIONz. REPORTNO. MMS 96-0041 z. 3.hci_e._=A==,,m.N_

PAGE

4. Title alKI Ikurbtitle L Report Oete

TaxonomicA'dasoftheSantaMariaBasinandWestemSantaBarbaraChannel.Volume July 1996

14. MiscellaneousTaxa. 6.

7. Authoc(s) 8. PeRorrninEO_nizatlon Rept. No.

James A. Blake,PaulH. Scott andAnderw L. Lissner(editors)

9. Per_rmk_l OtllnizMion Nmme and Addms 10. Project/Tesk/Work Unit No.

ScienceApplicationsInternationalCorporation zz.c_=_c) _ Gr.,d(m,_.

10260CampusPointDrive

San Diego,California92121 _{c) 14-35-001-30484

' (G)

|2. SponsoringOqianizatlon N=me and Adclress 13. Type of Report & PeriOdCovered

Final14 of 14volumes,

MineralManagementService,U. S. Departmentof theInterior, 9/89to 7/96 PacificOCS Region,770 PaseoCamarillo

Camarillo,California93010 _4.

IS. Supplementary Notes

Contributingauthors: Mary Bergen, DebraHart, GordonHendler, F. G. Hochberg,Andrew Lissner,Gretchen Lambert,

John F. Pilger,PaulaS. Winchell, Keith H. Woodwick.

li, Abstract (Limit: 200 words)

Volume14representstheconclusionofthe Atlas. Includedin thisvolumearerepresentativesof 6differentcoelomate

phyla:Brachiopoda,Sipuncula,Echiura,EchinodermataHemichordata , (Enteropneusta)a,ndUrochordaa(Ascidiacea).

Fiveclassesofthe Echinodermataareincluded:Crinoidea,Asteroidea,Ophiuroidea,Echinoidea,andHolothuroidea.

Atotalof97speciesand73 generaareincludedamongthesevariouscoelomatephyladistributedasfollows:Brachiopoda

(7 genera,9 species),Sipuncuta(4 generaand species),Echiura(4 genera, 3 species),Crinoidea(1 genusand

species),Asteroidea(8 generaandspecies),Ophiuroidea(14 genera,19species),Echinoidea(4 generaandspecies),

Holothuroidea(16 genera,28 species), Enteropneusta(4 generaandspecies),Ascidiacea(12 genera,16species).

Each speciesis fullydescribedand illustrated,with keys producedthatallowusers to identifythefauna. Forthe

Ophiuroidea,a lectotypewas designatedfor Amphiura granu/ata and the specieswas transferredto the genus

Amphichondrius. Arnhiop/ushexacanthus wasdeterminedto be a synonymof Douga/op/usamphacanthus. Three

newspeciesofascidians,Mofgufanapiformis,Bofteniapofypfacoderma,and Styelatesseriswere describedinseparate

paper(Lambert,1993:Proc. Calif.Acad.Sci.48).

iT. Document k_tys_s L O_c_=tem

Marineinvertebrates,Brachipoda,Echiura,Sipuncula,Echinodermata,Crinoidea,OphiuroideaAsteroi , dea,Echninoidea,

Holothuroidea,Hemicordata,Enteropneusta,Cordata,Urocordata,Ascidacea,taxonomy, California,Santa Maria Basin,

Santa BarbaraChannel, continental shelf.

Identlflers/Ol_m-F.nde¢tTerms

TaxonomicAtlas,Keysto species,faunalguide,marineecology,crustaceans,marinebenthos.

c. COSATI FleldlGmup

ill Availability _l_tement 19. Security Class (This Retx_t) 21. No. of PN_es

Unlimited Unclassified 302

20. Security Class (This P_e] 22. Price

_lmAN$1-Z39._-83 See Instructkms on Reverse OPTIONAL FORM 272 (4.71)

I (Formerly NTIS-351

De_memt of Commerce
I

/

Table of Contents 1

1. N BRACmOVODA(by F. G. Hochberg) I

Introduction ..................................................................................................................................... i /

Glossary ........................................................................................................................................... 7 II

List of Brachiopod Species Treated in this Chapter ........................................................................ 9

Key to Genera of West Coast Brachiopods ................................................................................... 10

Descriptions of Species ................................................................................................................. 10 |

Class Inarticulata Huxley, 1869 .................................................................................................... 10

Family Lingulidae Menke, 1828 ...................................................................................... 11 II

Glottidia albida (Hinds, 1844) ............................................................... 11 |

Suborder Craniidina Waagen, 1885 .............................................................................................. 13

Family Craniidae Menke, 1828 ........................................................................................ 13

Neocrania californica (Berry, 1921) ..................................................... 14 .I

Class Articulata Huxley, 1869 ...................................................................................................... 15

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Family Frieleiidae Cooper, 1959 ...................................................................................... 16 II

Frieleia halIi Dall, 1895 ......................................................................... 16

Suborder Terebratulidina Waagen, 1883 ...................................................................................... 18

Family Cancellothyrididae Muir-Wood, 1955 ................................................................. 18 /i

Key to Species of the Genus Terebratulina ........................................................ 18 II

TerebratuIina crossei Davidson, 1882 ................................................... 19

Terebratulina unguicula (Carpenter, 1864) ........................................... 21

Family Laqueidae Hatai, 1965 ......................................................................................... 23 1

Laqueus californianus (Koch, 1848) ..................................................... 23

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Suborder Terebratellidina Muir-Woods, 1950 .............................................................................. 25

Family Dallinidae Beecher, 1893 ..................................................................................... 25 n

Key to Species of Terebratalia ........................................................................... 26 Iff

Terebratalia occidentalis (Dall, 1871) .................................................. 27

Terebratalia transversa (Sowerby, 1846) .............................................. 29 l_

Family Platidiidae Thomson, 1927 .................................................................................. 32 n

Platidia hornii (Gabb, 1861) .................................................................. 32

Acknowledgments ......................................................................................................................... 34 1

Literature Cited ............................................................................................................................. 34 m

List of Specimens Examined ......................................................................................................... 42

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2. Pm'LtrM SWtrNCOLA(by Paula S. Winchell)

Inla'oduction ................................................................................................................................... 49

Taxonomic History ........................................................................................................................ 49 II

Systematics .................................................................................................................................... 50

Morphology ................................................................................................................................... 51 I

Biology .......................................................................................................................................... 53 I

Glossary ......................................................................................................................................... 53

List of Abbreviations on Figures ................................................................................................... 54 I

List of Species from the MMS Surveys ........................................................................................ 54 U

Key to the Sipuncula of the Santa Mafia Basin ............................................................................ 55

Descriptions of Species ................................................................................................................. 55 i
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I Nephasoma diaphanes (Gerould, 1913) ................................................. 55

Phascolion lutense Selenka, 1885 .......................................................... 57

i Siphonosoma (Siphonosoma) ingens (Fisher, 1947) .............................. 59 Thysanocardia nigra (lkeda, 1904) ....................................................... 60

Literature Cited ............................................................................................................................. 62

i 3, Ec_ (by John F. Pilger)

Introduction ................................................................................................................................... 67

Morphology ................................................................................................................................... 67 Taxonomic History ........................................................................................................................ 68

Biology .......................................................................................................................................... 68

n Glossary ......................................................................................................................................... 69 Abbreviations Used in the Figures ................................................................................................ 70

Key to the Orders, Families, Genera, and Species of Echiura from the Santa Mafia Basin and Western

Santa Barbara Channel .................................................................................................................. 71

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

of 72

Listriolobus pelodes Fisher, 1946 .......................................................... 72

Listriolobus hexamyotus Fisher, 1949 ................................................... 72

I Arhynchite californicus Fisher, .....................................................

1949 75

Nellobia eusoma Fisher, 1946 ................................................................ 77

Literature Cited ............................................................................................................................. 79

N 4, PnvI.trM EOnNOI)E_t_rA (by Andrew Lissner and Debra Hart)

Introduction ................................................................................................................................... 81

Sources of Material 82 Acknowledgments ......................................................................................................................... 83

Literature Cited ............................................................................................................................. 83 D

n 5. CLASSCm_on)_, (by Gordon Hendler)

Introduction ................................................................................................................................... 85

i History 85 Systematics .................................................................................................................................... 86

Morphology ................................................................................................................................... 86

I Ecology Preservationand Behavior ................................................................................................................................... ................................................................................................................... 8788

Glossary ...... ................................................................................................................................... 88

I Terminology Description and Conventions ...................................................................................................... 88 of Species ................................................................................................................... 89

Family Antedonidae ......................................................................................................... 89

I Subfamily Heliometrinae Florometra................................................................................................. serratissima (A.H. Clark, 1907) ......................................... 8989

Acknowledgements ....................................................................................................................... 92

i Literature Cited ............................................................................................................................. 93

6. CLASSASTEaO_r._ (by Andrew Lissner and Debra Hart)

I Introduction ................................................................................................................................... 97

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Taxonomic History of the Asteroids ............................................................................................. 98 M

Morphology of Asteroids .............................................................................................................. 98

Glossary ....................................................................................................................................... 100 I

Methods of Laboratory Examination for Asteroids .................................................................... 100 U

List of Species ............................................................................................................................. 101

Key to Orders and Species of Asteroidea Collected from the Santa Maria Basin and Western "ll

Santa Barbara Channel ................................................................................................................ 101 U

Descriptions of Species ............................................................................................................... 102

Family Luidiidae ............................................................................................................ 102 Ill

Luidia foliolata Grube, 1866 ................................................................ 102 n

Family Astropectinidae .................................................................................................. 104

Astropecten verrilli (Fisher, 1906) ....................................................... 104 m

Family Goniasteridae ..................................................................................................... 105 I

Hippasteria spinosa Verill, 1909 .......................................................... 105

Family Pterasteridae ....................................................................................................... 106 I

Pteraster tesselatus arcuatus (Ives, 1888) ........................................... 106 1

Family Poraniidae .......................................................................................................... 107

Poraniopsis inflata (Fisher, 1906) ....................................................... 107 l

Family Echinasteridae .................................................................................................... 108 ]1

Henricia leviuscula annectens (Fisher, 1910) ..................................... 108

Family Zoroasteridae ..................................................................................................... 109 i

Myxoderma platyacanthum H.L Clark, 1913 ....................................... 109 I!

Family Asteridae ............................................................................................................ 110

Rathbunaster californicus Fisher, 1906 ............................................... 110

Literature Cited ........................................................................................................................... 111 i

7. CLASSOt,muaomE_, (by Gordon Hendler)

Introduction ................................................................................................................................. 113 "1

I History ......................................................................................................................................... 114

Systematics .................................................................................................................................. 114

Morphology ................................................................................................................................. 115 l[

Ecology and Behavior ................................................................................................................. 116 U

Collection and Preservation ........................................................................................................ 119

Glossary ....................................................................................................................................... 119 i

Terminology and Conventions .................................................................................................... 123 III

Description of Species ................................................................................................................. 123

Family Asteronychidae .................................................................................................. 123 _l

Asteronyx Iongifissus D6dedein, 1927 ................................................ 123 II

Family Gorgonocephalidae ............................................................................................ 125

GorgonocephaIus eucnemis (Miiller and Tmschel, 1842) ................... 125 ]l

Family Amphiuridae ...................................................................................................... 128 II

Amphichondrius granulatus (Ltitken and Mortensen, 1899) new

combination .......................................................................................... 128 am

Amphiodia (Amphispina) digitata Nielsen, 1932 ................................. 131 I

Amphiodia (Amphispina) urtica (Lyman, 1860) .................................. 134

Amphioplus (Amphioplus) strongyloplax (H. L. Clark, 19I 1) ............. 137 I

Amphipholis pugetana (Lyman, 1860) ................................................. 140 il

Amphipholis squamata (Delle Chiaje, 1828) ....................................... 143

Amphiura arcystata H.L. Clark, 1911 .................................................. 145 i

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i Amphiura diomedeae Liitken and Mortensen, 1899 ............................ 147

DougalopIus amphacanthus (McClendon, 1909) ................................ 150

i Family Ophiacanthidae Amphiuridae .................................................................................................. sp. juvenile ..................................................................... 154 I55

Ophiacantha diplasia H. L. Clark, 1911 .............................................. 155

i Family OphiactidaeOphiopholis........................................................................................................ 157 bakeri McClendon, 1909 ................................................. 157

Family Ophiodermatidae ................................................................................................ 160

11 Ophiuroconis bispinosa Ziesenhenne, 1937 ........................................ 160

Ii Family Ophiotrichidae ................................................................................................... 162

Ophiothrix (Ophiothrix) spiculata Le Conte, 1851 ............................. 162

Family Ophiuridae .......................................................................................................... 164

I Ophiosphalmajolliense (McClendon, 1909) new combination .......... 164

Ophiura leptoctenia H. L. Clark, 1911 ................................................ 167

Ophiura luetkenii (Lyman, 1860) ........................................................ 169

I Acknowledgements .....................................................................................................................

172

Literature Cited ........................................................................................................................... 172

I 8. CLASSEcm_omra (by Andrew Lissner and Debra Hart) Introduction ................................................................................................................................. 181

Regular Urchins ........................................................................................................................... 181

I Irregular Echinoids 183

Taxonomic History of the Echinoids .......................................................................................... 185

Morphology of Echinoids ............................................................................................................ 185

I Glossary 185

• Methods of Laboratory Examination for Echinoids .................................................................... 185

List of Species ............................................................................................................................. 186

I Ke Channel y to the Species of Echinoidea Collected from the Santa Maria Basin and Western Santa Barbara ........................................................................................................................................ 186

Family Toxopneustidae .................................................................................................. 187

Lytechinus pictus (Verrill, 1867) ......................................................... 187 Family Strongylocentrotidae .......................................................................................... 188

Allocentrotusfragilis Jackson, 1912 .................................................... 188

I Family Schizasteridae Brisaster.................................................................................................... 189 latifrons (A. Agassiz, 1898) ................................................. 189

Family Brissidae ............................................................................................................. 191

I Literature Cited ........................................................................................................................... Brissopsis pacifica A. Aggasiz, 1898 .................................................. 193 191

i 9. CLASIntroduc SHOLOTHURO1DEA tion ................................................................................................................................. (by Mary Bergen) 195

Morphology ................................................................................................................................. 195

I Glossary Taxonomic....................................................................................................................................... History ...................................................................................................................... 200 199

Dissection and Slide Preparation ................................................................................................. 201

I Biology Key to Families, ........................................................................................................................................ Genera and Species from Shallow Water and Continental Shelf Depths of California 202

..................................................................................................................................................... 203

I Descriptions of Species ............................................................................................................... 206

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Family Synaptidae Burmeister, 1837 ............................................................................. 207 g

Leptosynapta spp.................................................................................. 207

Family Chiridotidae Ostergren, 1898 ............................................................................. 209 i

Chiridota spp ........................................................................................ 209 l

Family Molpadiidae Miiller, 1850 ................................................................................. 210

Molpadia intermedia (Ludwig, 1894) .................................................. 210 i

Family Caudinidae Heding, 1931 ................................................................................... 211 U

Caudina arenicola (Stimpson, 1857) ................................................... 211

Hedingia califomica (Ludwig, 1894) .................................................. 213 aim

Family Stichopodidae Haeckel, 1896 ............................................................................ 214 il

Parastichopus californicus (Stimpson, 1857) ...................................... 214

Parastichopusjohnsoni (The61, 1886) ................................................. 214 Ill

Parastichopus parvimensis (H.L. Clark, 1913) ................................... 216 |

Family Psolidae Perrier, 1902 ........................................................................................ 216

Psolus chitinoides H.L. Clark, 1901 .................................................... 217 n

Psolus squamatus (Koren, 1844) ......................................................... 218 i

Lissothuria nutriens (H.L. Clark, 1901) .............................................. 219

Lissothuria deichmannae Pawson, 1967 .............................................. 220 i

Family Cucumariidae Ludwig, 1894 .............................................................................. 220 l_

Subfamily Cucumariinae Panning, 1949 .......................................................... 220

Cucumaria curata Cowles, 1907 ......................................................... 221

Cucumaria miniata (Brandt, 1835) ...................................................... 222 1

Cucumaria piperata (Stimpson, 1864) ................................................ 223

Cucumaria pseudocurata Deichmann, 1938 ....................................... 225 I

Cucumaria saIma Yingst, 1972 ............................................................ 226 •

Pseudocnus lubricus (H.L. Clark, 1901) .............................................. 227

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Family Sclerodactylidae Panning, 1949 ......................................................................... 229

Eupentacta quinquesemita (Selenka, 1867) ......................................... 229 l

Pachythyone rubra (H.L. Clark, 1901) ................................................ 231

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Family Phyllophoridae Ostergren, 1907 ........................................................................ 231

Subfamily Thyoninae Panning, 1949 ............................................................................. 231 II

Havelockia benti (Deichmann, 1937) .................................................. 233

Havelockia charlottae (Deichmann, 1938) .......................................... 236

Pentamera lissoplaca (H.L. Clark, 1924) ............................................ 237 M

Pentamera populifera (Stimpson, 1857) .............................................. 239 u

Pentamera pseudocalcigera Deichmann, 1938 ................................... 241

Pentamera pseudopopulifera Deichmann, 1938 .................................. 243 i

Stolus trachyplaca (H.L. Clark, 1924) ................................................. 244 U

Thyone montereyensis (Deichmann, 1938) .......................................... 246

Literature Cited ........................................................................................................................... 247 1i

10. Hemichordata: Enteropneusta (by Keith H. Woodwick)

Introduction ................................................................................................................................. 251 i

Morphology ................................................................................................................................. 251

Taxonomic History ...................................................................................................................... 251

Biology ........................................................................................................................................ 252 i

Methods ....................................................................................................................................... 252

Systematic Account ..................................................................................................................... 253

Family Ptychoderidae Spengel, 1893 ............................................................................. 253 I

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U Family Spengeliidae Willey, 1899 ................................................................................. 254

Family Harrimaniidae Spengel, 1901 ............................................................................ 254

i Keys I. KeytoBaEnteropneusts sed on External................................................................................................................. Characteristics .................................................................................... 255 255

II. Key Based on Internal Characteristics .................................................................................... 255

i Literature Cited ........................................................................................................................... 258 11. PHYLUMCHORDATAI SUBPHYLUMUROCHORDATA, CLASSASCIDIACEA(by Gretchen Lambert)

i Introduction Morphology................................................................................................................................. 261 and General Biology ............................................................................................... 261

Taxonomy .................................................................................................................................... 265

Methods of Preservation and Dissection ..................................................................................... 266

I Key to Genera and Species of Ascidians listed in this Atlas ...................................................... 266

Description of Species ................................................................................................................. 269

Family Didemnidae ........................................................................................................ 269

I Didemnum carnulentum var. Iacteolum Ritter and Forsyth, 1917 ....... 269

Trididemnum opacum (Ritter, 1907) .................................................... 269

n Family Polycitoridae ...................................................................................................... 271

Distaplia occidentalis Bancroft, 1899 ................................................. 271 _F

Eudistoma carolinense (?) Van Name, 1945 ....................................... 271

Family Synoicidae .......................................................................................................... 273

I (Ritter and 191 273 Aplidium californicum Forsyth, 7)

Aplidium spauldingi (Ritter, 1907) ...................................................... 273

Euherdmania claviformis (Ritter, 1903) .............................................. 275

I Family Rhodosomatidae 277 Chelyosoma columbianum Huntsman, 1912 ........................................ 277

Family Molgulidae ......................................................................................................... 278

I Molgula regularis Ritter, 1907 278 Molgula napiformis Lambert, 1993 ..................................................... 279

Family Pyuridae ............................................................................................................. 281

I Boltenia polyplacoderma Lambert, 1993 281 HaIocynthia igaboja Oka, 1906 ........................................................... 283

Pyura haustor (Stimpson, 1864) .......................................................... 285

I Family Styelida .............................................................................................................. 286 Styela gibbsii (Stimpson, 1864) ........................................................... 286

Styela milleri Ritter, 1907 .................................................................... 287

I Literature Cited ........................................................................................................................... Styela tesseris Lambert, 1993 .............................................................. 28992

I _rE_ ..................................................................................................................................... 297

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List of Figures II

Figure 1.1. Diversity of shell types in the two classes of brachiopods - Articulata and Inarticulata. A. I

anterior end view; B, F, H, K. dorsal views; C, G, J. lateral views; D & E. interior view of II

dorsal valve; I. interior view of ventral valve; L. ventral view. A-C. Terebratella; D & E.

Magellania; F,& G. Lingula; H & I. Crania; J-L. Discinisca ........................................... 4 I

Figure 1.2. Diagramatic representations of brachiopod anatomy and morphology. A. principal organs,

lateral view; B. articulation and muscle system, median section; C & D. muscle scars, shell _i_

and brachidium terminology, interior views; C. ventral or pedicle valve; D. dorsal or brachial I

valve; E. mantle canal patterns, often seen through the shell, in a diversity of brachiopod

genera. [A & E, from Williams & Rowell, 1965; B, modified from Rudwick, i961; C & D, m

modified from Davidson, 1886] ......................................................................................... 5

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Figure 1.3. Diagramatic representations of brachiopod anatomy and morphology. A. lophophore loop

patterns; B. types of folding patterns in the anterior commissure of the shell, anterior end _l

views. Abbreviations: m, mouth; tb, tips of brachia. [A, from Rudwick, 1962; B, from II

Hatai, 1937] ........................................................................................................................ 6

Figure 1.4. A-C: Glottidia albida (Hinds, 1844); depth unknown, San Pedro Bay, California (SBMNH I

[SSB 155]); 24.4 mm TL. A. dorsal view; B. lateral view; C. end view, anterior margin. DE: Glottidia paImeri Dall 1871; depth unknown, San Fetipe, Baja California, Mexico ml

(SBMNH 46125); 42.2 mm TL. D. dorsal view; E. end view, anterior margin ............. 12 !

Figure 1.5. Neocrania californica (Berry, 1921). A-D: Holotype; 180 m, off Santa Monica, California

(SBMNH 34501); 13.0 mm TL. A. dorsal view; B. lateral view; C. end view, anterior i

margin; D. interior view of dorsal valve. E: 124 m, Cordell Bank, California (CAS 105948); II

20.0 mm TL. E. dorsal view ............................................................................................ 15

Figure 1.6. Frieleia halli Dall, 1895:1150-1170 m, off San Diego, California (SBMNH [SSB 3038]); e1

I7.7 mm TL. A. dorsal view; B. ventral view; C. lateral view; D. end view, anterior margin;

w

E. end view, posterior margin .......................................................................................... 17 lira

Figure 1.7. A-E: Terebratulina crossei Davidson, 1882; 88 m, off Santa Monica, California (SBMNH I

000000); 30.7 mm TL. A. dorsal view; B. ventral view; C. lateral view; D. end view,

anterior margin; E. end view, posterior margin. F: Terebratulina kiiensis Dall & Pilsbry, Ill

1891; 183 m, off La Jolla, California (SBMNH [SSB 3265]); 31.8 mm TL. F. dorsal view t

........ , .............................................................. , ......... ,,°, ........ ,,., ........ , ...................... , ......... 20

Figure 1.8. Terebratulina unguicula (Carpenter, 1865). A-E: 132 m, off Gaviota, California (SBMNH); I

18.3 mm TL. A. dorsal view; B. ventral view; C. lateral view; D. end view, anterior margin;

E. end view, posterior margin. F: depth unknown, off La Jolla, California (SBMNH [SSB

13507]); 17.0 mm TL. F. 2 specimens attached to solitary coral ................................... 22 1

Figure 1.9. Laqueus californianus (Koch, 1848). A-E: 90 m, off Santa Catalina Island, California,

(SBMNH); 45.7 mm TL. A. dorsal view; B. ventral view; C. lateral view; D. end view, ii

anterior margin; E, end view, posterior margin. F: 155 m, off Morro Bay, California I

(SBMNH [SSB 18432]); 42.5 mm TL. F. dorsal view ................................................... 26

Figure t.10. Terebratalia occidentalis (Dall, 1871): off Santa Catalina Island, California (SBMNH I

32400); 38.5 mm TL. A. dorsal view; B. ventral view; C. lateral view; D. end view, anterior I

margin; E. end view, posterior margin ............................................................................. 28

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I Figure 1.11. Terebratalia transversa (Sowerby, 1846). A-E: depth unknown, off Point Conception,

California (SBMNH 46074); 34.4 mm TL. A. dorsal view; B. ventral view; C. lateral view;

i DSan. endDiego, view,California anterior margin; (SBMNE./H); end22.5view,mmposte TL.riorF.margin. dorsal view; F & G:G.0.5endm,view, Missioante n Bay, rior

margin ............................................................................................................................... 31

I Figure 1.12. Platidia hornii (Gabb, 1861). A & B: 97 m, off Point La Jolla, California (SBMNH [SSB 30052]); 3.6 mm TL. A. dorsal view; B. ventral view. C-E: 45 In, off Monterey, California

(CASIZ 105953); 4.6 mm TL. C. dorsal view; D. ventral view; E. detail of punctae .... 33

i Figure 2.1. Morphology of a generalized sipunculan: A, external morphology; B, internal anatomy of

dissected specimen. (modified from Frank, 1983) ........................................................... 52

I Figure 2.2 Nephasoma diaphanes: A, internal, dissection of extreme anterior end; B, internal, dissection of posterior end showing origin of retractor muscles; C, portion of posterior end showing

papillae; D, hooks; E, reduced tentacles. (A-D after Cutler, 1973; E after Cutler et al.,

I 1984) ................................................................................................................................. 56

Figure 2.3 Phascolion Iutense: A, dissection showing internal anatomy; B, external view; C, hooks; D,

i glands with papillae; E, tentacular lobes (A-C after Murina, 1975b; D, after Selenka, 1885; E, after Curler and Cutler, 1985a) .................................................................................... 58

Figure 2.4 Siphonosoma ingens: A, internal, dissection of anterior portion of tnmk; B, internal, dissection

I C, of anterior end of paratype showing contracted retractor muscles, partly invaginated introvert; tentacles; D, internal, posterior trunk showing tubular, fusiform glandular bodies. (A-C

after Fisher, 1947; D after Cutler and Cutler, 1982) ........................................................ 59

I Figure 2.5 Thysanocardia nigra: A, internal view of dissected specimen; B, tentacles; C, papilla from

posterior of trunk. (A after Ikeda, 1924; B after Cutler and Cutler, 1981; C after Ikeda,

i Figure 3.1. Listriolobus 1904) ................................................................................................................................. pelodes: A, external anatomy, ventral view;(from Pilger, 1980). B, anterior 61

internal anatomy, dorsal view (after Fisher, 1946); C, posterior internal anatomy, dorsal

I view (after Pilger, 1980) .................................................................................................. 73

Figure 3.2. Listriolobus hexamyotus: A, external anatomy, ventral view; B, proboscis, detached, ventral

view; C, anterior internal anatomy, dorsal view .............................................................. 74

I Figure 3.3. Arhynchite californicus: A, external anatomy, ventral view; B, proboscis, ventral view; C,

anterior internal anatomy, dorsal view ............................................................................. 76

I Figure 3.4. Nellobia eusoma: A, external anatomy, ventral view; B, proboscis, ventral view; C, anterior

internal anatomy, dorsal view .......................................................................................... 78

I Figure 5.1. FIorometra mm) ................................................................................................................................. serratissima (A.H. Clark): Lateral view (LACM 41-176.3; arm length = 160 .91

Figure 6.1. Common external features of many seastars (Astropecten is depicted). Source: Word and

I Charwat, ..................................................................................................................

1975 99

Figure 6.2. Aboral view of arm from Luidiafoliolata (Station 022-BRA-02-TX), emphasizing lack of

i prominent supramarginal plates, but distinct arm spines ............................................... 103

Figure 6.3. Aboral view of arm fromAstropecten verrilli (Station 001-BRA-02-TX), showing prominent,

I spinous, supramarginal plates and relatively small paxillae .......................................... 104

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Figure 6.4. A. Aboral view of arm from Hippasteria spinosa (Station 020-BRA-03-TX), indicating U

extensive spination on surface and marginal spines. B. Oral view of arm from Hippasteria

spinosa (Station 020-BRA-03-TX), depicting lack of prominent surface spines and paxillae

surrounding clam-like pedicellariae ............................................................................... 105 I1

Figure 6.5. A. Aboral view of Pteraster tesselatus arcuatus (Station 021-BRA-02-TX), emphasizing

stubby body form, smooth surface, and central osculum. B. Oral view ofPteraster tesselatus •

arcuatus (Station 021-BRA-02-TXr), showing spinous ambulacral grooves and Prominent I

mouth plates ................................................................................................................... 106 l

Figure 6.6. A. Aboral view of Poraniopsis inflata (Station 021-BRA-02-TX), indicating body form I

and relatively sparse and evenly distributed surface spines. B. Oral view of Poraniopsis

inflata (Station 021-BRA-02-TX), emphasizing numerous, small podia per ambulacral row /

........................................................................................................................................ 107 l

Figure 6.7. Aboral view of arm fromHenricia leviuscula annectens (Station 01-BRA-03-TX), depicting

"pock-marked" appearance of papulae surrounded by pseudopaxillae ......................... 108 I

Figure 6.8. Aboral view of Myxoderma platyacanthum (Station R-7, Pep. 2), showing body form and

reticulate pattern of surface plates ................................................................................. 109 •

Ii Figure 6.9. A. Aboral view ofRathbunaster caIifornicus (Station 026-BRA-02-TX), indicating extensive,

stalked pedicellariae on surface. B. Oral view of Rathbunaster californicus (Station 026-

BRA-02-TX), emphasizing rows of numerous podia and large width of ambulacrum li

compared to overall arm width ...................................................................................... 110 lib

Figure 7.1. Major anatomical features of art ophiuroid: A, the dorsal surface of the disk; B, the dorsal Ill

surface of an arm; C, the ventral surface of an arm; D, the ventral surface of the disk; E, a U

ventral interradius and the jaw structures that project into the mouth. (After Hendler et al.,

1995) ............................................................................................................................... 117 /

I

Figure 7.2. Asteronyx longifissus Dtdedein: A, dorsal view; B, disk, ventral view; C, distal portion of

an arm, ventral view, showing elongate, club-tipped ventral-most spine and hooked dorsal

arm spines (LACM 77-204.1; dd = 25.0 ram) ............................................................... 124 i

m

Figure 7.3. Gorgonocephalus eucnemis (Miiller and Troschel): A, dorsal view (LACM 42-39.4; dd =

35.0 ram); B, ventral view (LACM 65-331.1; dd = 56.0 ram) ...................................... 127

II Figure 7.4. Amphichondrius granulatus (Liitken and Mortensen): A, disk, ventral view; B, disk, dorsal

view; C, ventral interradius, showing granule covering (USNM 19539, lectotype,Amphiura

granulata; dd = 6.4 mm); D, disk, ventral view; E, disk, dorsal view; F, ventral interradius, i

showing granule covering (LACM 38-122.4; dd = 6.1 mm) ......................................... 130 I

Figure 7.5. Amphiodia (Amphispina) digitata Nielsen: A, disk, dorsal view; B, disk, ventral view III

(syntype, ZM; dd = 4.9 ram); C, disk, dorsal view (LACM 39-82.7; dd = 4.9 mm); D, disk II

ventral view (LACM 39-109.11; dd = 7.4 ram) ............................................................. 133

Figure 7.6. Amphiodia (Amphispina) urtica (Lyman): A, disk, dorsal view; B, disk, ventral view; C, i

disk edge, showing spinose scales on disk margin (LACM 56-66.7; dd = 4.9 mm) ..... I36 II

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I Figure 7.7. Amphioplus (Amphioplus) strongyloplax (H.L. Clark): A, disk, dorsal view; B, disk, ventral

view (USNM 25615, holotype, Amphiodia strongyloplax; dd = 4.9 mm); C, disk, dorsal

I view; ventralDview , disk, ventral view (LACM 64-242.1; dd = 7.2 mm); E, disk, dorsal view; F, disk,

(LACM 55-110.1; dd = 9.3 mm); G, disk, dorsal view; H, disk, ventral view

(LACM 75-409.4; dd = 6.6 mm). Note variability in disk scalation; all disks are regenerating

i ........................................................................................................................................ 138 Figure 7.8. Amphioplus (Amphioplus) macraspis (H.L. Clark): A, disk, dorsal view (USNM 25584,

syntype,Amphiodia macraspis; dd = 11.5 ram); B, disk, ventral view; C, oral frame, showing

I broad oral shields (USNM 25584, syntype, Amphiodia macraspis; dd = 8.5 mm) ....... 139

Figure 7.9. Amphipholis pugetana (Lyman): A, disk, dorsal view; B, disk, ventral view (LACM 75-

i 297.2; dd = 4.1 ram) ....................................................................................................... 142 Figure 7.10. Amphipholis squamata (Delle Chiaje): A, disk, dorsal view (LACM 41-5.12; dd = 3.6 ram);

B, disk, ventral view (LACM 41-5.12; dd = 3.3 mm) ................................................... 144

I Figure 7.11. Amphiura arcystata H.L. Clark: A, disk, dorsal view (LACM 39-79.11; dd = 6.4 mm); B,

disk, ventral view (LACM 39-100.7; dd = 10.1 ram); C, disk, dorsal view; D, disk, ventral

i view (LACM 39-79.11; dd = 6.4 ram). Note variability in disk scalation .................... 146 Figure 7.12. Amphiura diomedeae Liitken and Mortensen: A, disk, dorsal view; B, disk, ventral view; C,

oral frame (USNM 19532, syntype; dd = 15.1 mm); D, oral frame (USNM 19532, syntype;

I dd = 11.0 mm). Note variability of Amphioplus-like oral papillae ................................ 149

Figure 7.13. Dougaloplus amphacanthus (McClendon): A, disk, dorsal view; B, disk, ventral view; C,

i oral frame; D, arm, dorsal view; E, arm, ventral view; F, arm spines, showing glassy teeth (LACM 58-140.7; dd = 7.4 mm) .................................................................................... 152

Figure 7.14. Amphioplus hexacanthus H. L. Clark: A, entire, dorsal view, disk missing; B, entire, ventral

I vE,iearmw; C, oral frame; D, arm, dorsal view, showing elongate, flattened, middle arm spines; spines, showing glassy teeth (USNM 25583, syntype; ofl = 2.5 ram) ............... 153

Figure 7.15. Amphiuridae sp. juvenile: A, disk, dorsal view; B, disk, ventral view (USNM, primary

I voucher; dd = 3.3 mm) ................................................................................................... 155

Figure 7.16. Ophiacantha diplasia H.L. Clark: A, disk, dorsal view; B, disk, ventral view (USNM, primary

I voucher; dd = 14.8 ram) ................................................................................................. 157

Figure 7.17. Ophiopholis bakeri McClendon: A, disk, dorsal view; B, disk, ventral view CLACM 40-

341.2; dd = 7.9 mm) ....................................................................................................... 159

I Figure 7.18. Ophiuroconis bispinosa Ziesenherme: A, disk, dorsal view; B, disk, ventral view (LACM

92-53.2; dd = 2.5 ram) .................................................................................................... 161

I Figure Ophiothrix (Ophiothrix) spiculata A, disk, view; B, disk,

7.19. Le Conte: dorsal ventral view

(LACM 37-85.1; dd = 6.6 mm) ...................................................................................... 163

I Figure 7.20. Ophio 41-205.2; sphalmajollien dd = 12.0seram).(MOphio cClensphalma don): A,variabile disk, dorsal (Liitken view;andB,Mortensen): disk, ventralA,viewdisk,(LACM dorsal

view; B, disk, ventral view (USN/Vl 19502, syntype, Ophiomusium variabile; dd = 13.9

i ram) ................................................................................................................................ 166

Figure 7.21. Ophiura leptoctenia H. L. Clark: A, disk, dorsal view; B, disk, ventral view (LACM 64-

I 240.2; dd = 6.1 mm) ....................................................................................................... 168

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Figure 7.22. Ophiura luetkenii (Lyman): A, disk, dorsal view; B, disk, ventral view; C, disk edge, showing i

blunt, close-set scales of the arm comb (LACM 38-155.5; dd = 13.9 ram) .................. 171

Figure 8.1. Typical aboral morphology of regular echinoid (after Word and Charwat, 1975) ........ 182 I

Figure 8.2. Typical aboral morphology of irregular echinoid (after Word and Charwat, 1975) ..... 183

Figure 8.3. Oral (upper) and aboral (lower) views of regular echinoid (after Pierce and Maugel, 1987) I

• .... ** .................. ..,,,,., ........... , ............. • ................. . ........ , ............................. , .................... 184

Figure 8.4. Aboral view of undisturbed Lytechinus pictus (upper) and cleaned test (Lower) of test and i

spines (after Word and Charwat, 1975) ......................................................................... 187 I

Figure 8.5. Aboral view of undisturbed test and spines (upper) and cleaned test (lower) of Allocentrotus

fragilis (after Word and Charwat, 1975) ........................................................................ 188 H

Figure 8.6. A. Aboral view of cleaned test of Brisaster latifrons (after Word and Charwat, 1975) ...... 190

Figure 8.7. Aboral views of Brisaster latifrons (left) showing sunken petaloids, notched anterior I

ambulacrum, and faint peripetalous fasciole; and Brissopsis pacifica (fight) indicating i

"shallow" petaloids and prominent peripetalous fasciole. Specimens are from Stations 088

and 025, respectively ...................................................................................................... 190 H

Figure 8.8• Aboral view (left) of undisturbed test and spines (left-side) and cleaned test (right-side),

and oral view (right) of undisturbed test and spines of Brissopsis pacifica. Source: Word i

and Charwat, 1975 .......................................................................................................... 192 _1

Figure 8•9. Posterior views ofBrissopsis pacifica(fight) showing prominent subanal fasciole, contrasted

with Brisaster latifrons (left). Specimens are from Stations 025 and 088, respectively 192 H

Figure 9.1. Types of tentacles: A, peltate; B, digitate; C, tentacles with 2 pairs of digits and no terminal

digit ................................................................................................................................. 196

II Figure 9.2. Whole animal, dissected to show internal morphology (from Pawson, 1971) .............. 197

Figure 9.3. Types of ossicles: A, anchor; B, anchor plate; C, wheel; D, miliary body or granule; E, cup; I

F, table; G, supporting table; H, plates; I, buttons ......................................................... 198 |

Figure 9.4. Types of calcareous rings: A, simple ring; B, ring with short processes; C, ring with long

processes ......................................................................................................................... 201 HI

Figure 9.5. Leptosynapta spp.: A, anchors; B, anchor plates; C, miliary granules .......................... 208

Figure 9.9. Chiridota spp.: Wheels from body wall ........................................................................ 209 H

Figure 9.7. Molpadia intermedia: A, tables from caudal appendage; B, tables from body wall of small

individual, table at right forming phosphatic bodies; C, rosette of raquet-shaped plates with a

anchor (not to scale, copied from H.L. Clark, 1907); D. racquet-shaped plate ............. 212 II

Figure 9.8. Hedingia californica: ossictes from caudal appendage of syntype ................................ 213

Figure 9.9. Parastichopusjohnsoni: ossicles from the body wall of the holotype .......................... 215 H/

Figure 9.10. Psolus chitinoides: buttons from sole ........................................................................... 217

m

Figure 9.11. Psolus squamatus: plates from sole .............................................................................. 218 H

Figure 9.12. Lissothuria nutriens: ossicles from sole, A, plates; B, cups ......................................... 219

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I Figure 9.13. Cucumaria curata: ossicles from body wall .................................................................. 221

Figure 9.14. Cucumaria miniata: ossicles from body wall ................................................................ 223

Figure 9.15. Cucumaria piperata: ossicles from body wall .............................................................. 224

Figure 9.16. Cucumariapseudocurata: ossicles from body wall ....................................................... 225

Figure 9.17. Cucumaria salma: ossicles from body wall ................................................................... 226

Figure 9.18. Pseudocnus lubricus: ossicles from body wall .............................................................. 228

I Figure 9.19. Eupentacta quinquesemita: ossicles from body wall, A, plates; B, cups; C, supporting tables

from tube feet ................................................................................................................. 230

i Figure 9.20. Pachythyone rubra: ossicles from body wall, A, reticulated plates; B, 4-holed buttons; C, supporting tables from tube feet ..................................................................................... 232

I Figure 9.21. Havelockia from introvert. bentiHavelockia : A, tables benti from var. bodyzacae wall;: D, B tables supporting from tables body wall; fromE,tubesupporting feet; C, tables

from tube feet. Havelockia variant: F, biscuit-shaped plates and bar-shaped deposit .. 235

I Figure 9.22. Havelockia charlottae: A, tables from body wall; B, tables from introvert 237

Figure 9.23. Pentamera lissoplaca: A, lozenge-shaped plates, B, small tables, C, large tables, D,

I supporting tables; E, large plate from specimen from San Clemente ............................ 238 Figure 9.24. Pentamerapopulifera: A, ossicles from body wall; B, supporing tables ...................... 241

i Figure 9.25. Pentamera supporting tables; psuedocalcigera D, plate :fromA,introvert plates from ...................................................................... body wall; B, tables from body wall;242C,

Figure 9.26. Pentamera pseudopopulifera: A, ossicles from body wall; B, supporting tables ......... 244

I Figure 9.27. Stolus trachyplaca: A, plates from body wall; B, supporting table; C, plates from introvert

.............................................. , ........... ,,.o,,._.o,,o.,. .............................................................. 245

I Figure 9.28. Thyone montereyensis: A, buttons from body wall; B, supporting tables; C, irregular plate from specimen from Long Beach; D, plates from introvert ........................................... 246

I Figure 10.1. Plate key .................................................................................................................................. showing anterior ends of four different genera of enteropneusts. Letters explained256in

Figure 10.2. Plate showing cross sections of the proboscis in four different genera of enteropneusts.

I Letters explained in key ................................................................................................. 257

Figure 11.1. A, ascidian tadpole larva greatly enlarged (from Kozloff, 1990); B, a typical compound

I ascidian zooid with postabdomen, x16. (from Abbott, 1975) ....................................... 263 Figure 11.2. Pyura haustor, an example of a typical solitary ascidian (approximately life size) showing

major body parts. A, method of dissection; B, animal spread open to show inner surfaces of

I oralsiphon and branchial sac (pharynx); C, same view asin B but with branchial sac removed, oral tentacles not shown; D, a small part of branchial sac, enlarged. (from Abbott, 1975)

.............................. ............................................... ........................................................... 264

I Figure 11.3. Didemnum carnulentum var. lacteolum. A, zooid, x50 (a.o.: atrial opening; m.b.: muscle

band; o.: ovary; s.: stomach; t.: testis; v.d.: vas deferens); B, spicule, x650 (from Ritter and

i Forsyth, 1917). C-D, Trididemnum opacum. C, zooid, X30; D, spicules, x900. (from Van Name, 1945) ................................................................................................................... 270
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Figure 11.4. Distaplia occidentalis, A, zooid, x42 (b.s. - branchial siphon, e - esophagus, end. - endostyle,

i.p. - incubatory pouch, o. - ovary, ov.d. - oviduct, p.gl. - pylofic gland, s. - stomach, t. -

testis, v.d. - vas deferens) (from Ritter and Forsyth, 1917); Eudistoma carolinense, B, 1

colony, xl.3 (from Van Name, 1945); C, zooid, left and fight sides of body, x3.2 (from I!

Plough, 1978) ................................................................................................................. 272

Figure 11.5. Aplidium californicum, A, zooid, x22 (fp - fecal pellet, sd - sperm duct, st - stomach) (from /

Van Name, 1945, adapted from Ritter and Forsyth, 1917); Aplidium spauldingi, B, zooid, ill

x8 (al. - alrial languet, ov. - ovary, t. - testis) (from Ritter, I907) ................................. 274

/m

Figure 11.6. Euherdmania claviformis. A, young colony; B, older colony, x2; C, zooid with embryos I

and larvae in oviduct, x9; D, cross section of stomach; E, part of branchial sac with dorsal

languets and adjacent stigmata, greatly enlarged. (A, C, D, E from Van Name, 1945, after Ill

Ritter, 1903; B from Johnson and Snook, 1927) ............................................................ 276 |

Figure I1.7. Chelyosoma columbianum. A, whole animal, x2.4 (redrawn from Huntsman, 1912a); B,

plates of disk surrounding apertures, showing connecting muscles, x3 (from Van Name, I

1945, after Huntsman, 1912a); C, plates of disk from another animal, with a slightly different !

pattern of plates, x2.2 (redrawn from Huntsman, 1912a); Molgula regularis. D, left side of

body; E, fight side; x5. (a - atrial siphon, b - branchial siphon, k - kidney) (from Van Name,

1945) ............................................................................................................................... 278 It

Figure 11.8. Molgula napiformis. A-B, whole animals, x13 (a - atrial siphon, b - branchial siphon); C, I

dorsal tubercle; D, three branchial tentacles; E, detail of branchial sac. C-E greatly enlarged |

........................................................................................................................................ 280

Figure 11.9. Molgula napiformis. A, fight side of animal removed from tunic; B, left side. (a - atrial

siphon, b - branchial siphon, d - dorsal ganglion, e - endostyle, 1- liver, o - ovary, r - rectum, II

t- testis), x32 ................................................................................................................. 281

Figure 11.10. Boltenia polyplacoderma. A, anterior surface of tunic, x7.2; B, detail of branchial sac; C, I

dorsal tubercle; D, branchial tentacle (B-D greatly enlarged); E, anterior surface of mantle

showing the siphons and musculature; F, posterior view of body removed from tunic; E-F /

x9.7.(a - atrial siphon, b - branchial siphon, e - endostyle, 1- liver, lv - longitudinal vessels,

o - ovary, od - oviduct, s - stigmata, st - stomach, r - rectum, t - testis) ........................ 282

Figure 11.11. Halocynthia igaboja. A, whole animal, x2.3 (redrawn from Abbott and Newberry, 1980);

B, a large tunic spine, x4; C, a smaller spine without terminal whorl of spinelets, x4; D, two II

of the short whorled spines, x27; E, a single branched oral tentacle, greatly enlarged. (B-E

from Ritter, 1907) ........................................................................................................... 284

J

Figure 11.12. Pyura haustor. A, whole animal, life size (redrawn from Kozloff, 1990) (a - atrial siphon,

b - branchial siphon); B, a clump of animals, x0.4 (from Herdman, 1898) .................. 285 I

Figure 11.13. Styelagibbsii. A, whole animal, xl .4 (redrawn from Huntsman, 1912a);B, pieceofbranchial I

sac, x31; C, alimentary canal and gonads, xl.6 (B and C from Herdman 1898); Styela milled

(from Van Name, 1945). D, body outline, x2; E, terminal part of gonad, x6. (a - atrial siphon,

iv - longitudinal vessels, o - ovary, r - rectum, s - stomach, sd - sperm duet, t - testis ........ 288 II

Figure 11.14. Styela tesseris. A, whole animal, x7; B, detail of tunic, x23; C, dorsal tubercle; D, small m

piece of branchial sac (C-D greatly enlarged); E, body removed from tunic, fight side, x7.

(a - atrial siphon, b - branchial siphon, lv - longitudinal vessel, s - stigmata) ............... 290

I
!

I Figure 11.15. Styela tesseris. A, animal opened to show internal anatomy, x9; B, enlarged detail of gonad;

C, detail of stomach and anterior intestine showing the delicate connection between them.

I (aes- atrial siphon, at - atrial tentacles, b - branchial siphon, bt - branchial tentacles, e - endocarps, - esophagus, i - intestine, o - ovary, pd - pytoric duct, s - stomach, sd - sperm duct, t - testis)

............ .................................................. ................. .............. ...................... ..................... 291

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Acknowledgments I

This study was funded by the Pacific Outer Continental Shelf Region of the U.S. Department of the I

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 1

assistance for this volume. Ms. Adele Smith, also of the Santa Barbara Museum, assisted with editing and M

production of this volume.

i

List of Acronyms i

AMNH American Museum of Natural History, New York, USA. I

ANSP Academy of Natural Sciences, Philadephia, USA.

BLM Bureau of Land Management. !

i

BMNH The Natural History Museum, London, U.K.

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

II transect A/B.

BRC Refers to a station designation from the MMS Phase I Reconnaissance: Benthic Rocky, 1

transect C/T).

CAMP California Phase 1I Monitoring Program (by MMS). iI

CASIZ California Academy of Sciences, Department of Invertebrate Zoology, San Francisco, I

California, USA.

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

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

MMS United States Minerals Management Service. i

SCAMIT Southern California Association of Marine Invertebrate Taxonomists.

Natural History, Santa Barbara, California, USA. i

i

SBMNH Santa Barbara Museum of

SCCWRP Southern California Coastal Waters Research Project

Diego Natural History Museum, San Diego, California, USA. I

SDNHM San

SSB S. Stillman Berry Collection, now housed at the Santa Barbar Museum of Natural History.

Scripps Institute of Oceanography, La Jolla, California, USA. I [

SIO

UCSB University of California, Santa Barbara, USA.

/

United States National Museum. A historical designation for the National Museum of B

USNM

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

ZMUC Zoological Museum, University of Copenhagen, Denmark. I

xxii I
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I 1. THE BcmovooA

I by

F.G. Hochberg 1

I

I Introduction

Brachiopods or "lamp shells" are a phylum of exclusively marine invertebrates. The phylum is

I characterized by the presence of a lophophore, a tentacle bearing structure used for feeding and respiration. The lophophore unites the brachiopods with several other phyla, namely the Phomnida and the Bryozoa or

Ectoprocta. Brachiopods are solitary animals enclosed in a bivalved shell and typically are either attached to

I a hard substmte or anchored in asoft substmte by a tough, flexible stalk or pedicle. A diversity of brachiopods in the two classes, Inarticulata and Articulata, is illustrated in Figure 1.1.

Approximately 350 species of living brachiopods are known which is but a small fraction of the

I than 20,000 extinct described in the literature. The fossil record for dates back at

more species brachiopods

least 600 million years. Off the coast of California brachiopods range in depths from the intertidal to several

thousand meters. Adults range in length from 1-100 mm. I

Morphology and Anatomy

!

Shell morphology with a focus on fossil forms is documented in an extensive literature summarized

in Moore (1965). Details of internal anatomy are know for relatively few living species. Most anatomical

I and histological studies are 50-100 years old but still remain the principal original sources

of information

and illustrations (for reviews see Hyman, 1959; Williams and Rowell, 1965; and Rudwick, 1970). The

literature on functional morphology is reviewed in James et al. (1992). Some of the key features used in

I briefly summarized below. Additional details can be found in the references listed above. systematics are

Shells. The brachiopod shell is composed of two separate valves that are bilaterally symmetrical

but dissimilar. The valves are oriented dorsal and ventral in reference to the body unlike in bivalve mollusks

i in which the valves are oriented lateral to the animal inside. In life the ventral valve in articulates typically is uppermost in reference to the substrate to which they are attached. The ventral valve, often termed the

pedicle valve, is generally larger and deeper than the dorsal valve. It contains the hole or foramen, through

I which the pedicle protrudes. The dorsal, or brachial valve, contains the lophophore and the supporting loop.

Brachiopods are identified principally on descriptions of the size, shape, sinuosity and sculpture of

the valves. Visually they are identified by figures which illustrate dorsal, ventral, lateral and end views.

i Shapes are expressed by a comparison ratios lengths to or depths to sinuosity

of of widths widths. The of

valves is described based on the anterior and lateral commissures. Simple folding patterns as used in the

following text are illustrated in Figure 1.3B. External valve surfaces may be smooth or textured with radiating

I ribs, concentric growth lines, spines and a diversity of other features. Valves be pitted with canals or

may

punctae which are seen as tiny holes in the shell (Figure 1.12E). The presence of muscle attachment scars

leave characteristic patterns on the internal surfaces of valves (Figs. 1.2C and D, 1.5D).

I

JDepartment of Invertebrate Zoology, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road,

I Santa Barbara, CA 93105-2936

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II

The valves are either locked together posteriorly by teeth and sockets (Class Articulata) or held l_

together solely by muscle (Class Inarticulata). The articulation of teeth and sockets in the hinge of articulate

brachiopods is important in diagnosing various taxa in the class, m

There is a complex and rather formidable terminology that covers in minute detail every feature II

used to describe brachiopod shells (see Moore, I965).

Pediele. Most brachiopods are attached to a hard substrate or anchored in soft sediments by a i

cylindrical stalk or pedicle which emerges from the posterior end of the shell (Figs. 1.1, 1.2). In a few cases, II

such as Neocrania, the pedicle is absent. In inarticulates the pedicle emerges between the two valves but in

the articulates it emerges from the beak of the ventral valve through a hole or foramen. The foramen is i

formed by the partial closure of the delthyrium by the deltidial plates. The opening is described as being IN

either complete or incomplete.

Lophophore. The lophophore develops as an outgrowth of the anterior body wall. It projects into i

and fills the greater part of the mantle cavity. The lophophore is lined along its entire length with ciliated II

tentacles. It functions for both feeding and respiration.

Compared with other lophophorate phyla (e.g., Phoronida, Bryozoa) brachiopod lophophores come i

in a tremendous diversity of shapes and sizes. Specific named patterns characterize the various higher taxa.

"lB

The evolution of lophophore shape and efficiency involves increasing lengths and complexity in bending to

accommodate more tentacles (Figure 1.3A). The lophophore is supported by a calcareous loop or brachidium

(Figs. 1.1D, 1.2D).

u

Mantle. The mantle of brachiopods is patterned with a series of coelomic channels or sinuses. I

These mantle canals may leave impressions on the inner surfaces of the valves or may be seen through the |

shell in live animals or wet preserved specimens. Mantle canal patterns are characteristic generic features

(Figure 1.2E). m

In addition, spicules may be present in the mantle tissue of some brachiopods. The presence or

absence of spicules is another character used at the generic level and above.

!

Reproduction

Sexes are separate in most brachiopods, although a few genera, such asPlatidia, are hermaphrodites. I

Sexual dimorphism appears to be absent. Two pairs of gonads (dorsal and ventral) develop from cells lining

the coelom. When mature they expand greatly to fill the space within the mantle. The majority of brachiopods Im

studied do not appear to have a breeding cycle, although a few exhibit distinct spawning seasons. When II

spawning occurs sperm and eggs are released and pass into the mantle cavity where they are then carried to

the exterior in exhalent currents. Fertilization in these cases is external. In a number of cases ova are retained

in the mantle cavity and following internal fertilization the embryos undergo extended development in a II

brood pouch.

Details of larval development are known in only a few cases (see Reed, 1987 and Long and Stricker, I

1994 for reviews of reproduction and development in west coast species). Free swimming, ciliated brachiopod I

larvae, known as "protegula," spend varying periods in the plankton depending on the class. In articulates

the protegula are comprised of three segments. Following a brief phase in the plankton, they settle at an In

early stage of growth. The protegula of inardculates consist of two segments. These larvae spend a much It

longer period in the plankton and are much further advanced when they settle to the bottom. At the point of

settlement in both classes the valves are present, the lophophore is well developed and used for both feeding I

and swimming, and the pedicle is evident at the edge of the posterior mantle. Settlement may be delayed if It

the bottom is too deep or if the subslxate first contacted is unsuitable.

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I Biology and Classification

I The paper by James et al. (1992) provides the most comprehensive recent review of the biology and ecology of living bmchiopods.

Overviews of brachiopod classification variously are treated in Moore (1965) and Foster (1982).

I Evolution of the phylum is discussed in Williams and Hurst (1977) and Wright (1979).

The identification of brachiopods from the Eastern North Pacific is in need of critical reevaluation

i which is beyond the scope of this chapter. Many species appear to be extremely variable not only in color but shape and sculpture which has created considerable confusion in the literature. There is a morass of

synonyms in which the validity of many Recent and fossil variety and subspecies names have not been

adequately reviewed in the recent literature. Bernard (1972) provides the most recent comprehensive,

I illustrated of forms from the Eastern North Pacific.

SUffi/_ living Deep-water brachiopods, including

several species treated here, are covered in Cooper (1972, I982). West coast fossils are reviewed in Hertlein

and Grant (1944, 1960).

I Brachiopod shells are comparable to the shells of mollusks in attracting collectors which has resulted

in a proliferation of names. Brachiopods are often listed with mollusks in earlier faunal reports from a

i variety of locations in the Eastern North Pacific. For the purposes of this paper I have not attempted to review the extensive fossil literature. In the synonymy section for each species treated below I have attempted

to list the key works to provide a chronology of spellings or mispellings of the morass of names that have

been used for west coast species. For more complete citations the reader is referred to Dall (1920) and to

I Hertlein and Grant (1944, 1960).

I Collection and Preservation

I FollowingDeep- colwater brachiopods typically are collected utilizing bottom trawls, benthic sleds and dredges. lection specimens should be sorted out from the remainder of the catch and cleaned of any

debris. Before epifaunal specimens are removed from their substrates observations should be made to

i determine whether the substrate is biological (type of organism to which they are attached; whether the organic substrate is living or dead) or inorganic (rock type). If possible, preserve small pieces of substrate

along with the sample for later identification. Avoid storing large rocks with brachiopods because their

i shells are very fragile and subject to breakage if handled or placed together with heavy objects. Additional notes should be taken to indicate whether a given species is attached singly, in small clusters or in dense

aggregations.

I based solMost ely onbrachiopod shell morpholog speciesy are and dried do notforrelease y onofintset

rna oralgeanatom and sy.tudy. HowIden evertifications , in any givmost en col oftenlectingare

program where material might be utilized for taxonomic study a subsample should be preserved in alcohol.

I

them Smallinbrachiopods alcoholizedmaysea bewater placedfor directly several hours into 70before % ethythey l alcohol are placed . Largeintospecimens alcohol are for narcotized preservation.by placing Prior to

preservation place a toothpick or small piece of cork between the valves to keep them open. Brachiopods

i should not Specimens be fixed orto stored be prepared in formalin for anatomical unless it isstudy buffermust edofirst prevenbet decalcified damage to the in Bouin's shell. solution or

another appropriate decalcifying agent. Stained whole mounts of small, fiat brachiopods (e.g., Glottidia,

I Neocrania, prior to examination. Platidia) are easily prepared. Larger, deep-bodied specimens need to be sectioned and stained

!

! 3

I

I DORSAL

stomach digestive diverticulum

blind intestine spiral part of lophophore

I dpeo_rsicalleadjustormuscle lophophore cross section (side arm)

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! , POSTERIOR

I adductor mmde ANTERIOR

I #t nephridium PEDICLE VALVE

m°U_°nad mande c_v.'e_ mantle VENTRAL

I / foramen

deltidium _/._ _ tooth ,. _ / pedicle hinge plate _ _ cardinal process

I _ _ ._<//tooth //__ socket wall _ socket

_1 :_ "Quick" erura

I ((/ ®_\ll * II ])'"-(_'I( II

I 2/\ Ill//loop

I

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Lingula

I Figure 1.2. Diagramatic representations of brachiopod anatomy and morphology. A. principal organs, lateral

view; B, articulation and muscle system, median section; C, D, muscle scars, shell and brachidium

terminology, interior views; C, ventral or pedicle valve; D, dorsal or bracMal valve; E, mantle canal

I paRowtterns, ell, 1of965; ten seen B, modifi throeugh d ftrheom sheRudwick, ll, in a dive196rsi1ty; C,of D,bracmohiopod dified gefxomnera.Davidson, (A, E, from1886).Williams and
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trocholophe[ I

I

,

zygolophe _ shizolophe _ early spirolophe _

plectolophe _ ptycholophe spirolophe I

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rectimarginate e antiplica_e episculate 1

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Figure 1.3. Diagramatic representations of brachiopod anatomy and morphology. A, lophophore loop patterns; IB

B, simple types of folding patterns in the anterior commissure of the shell, anterior end views.

Abbreviations: m, mouth; tb, tips ofbrachia. (A, from Rudwick, 1962; B, from Williams and Rowell, It

1965). |

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I Glossary

I Adductor muscles. Paired muscles which, on Crura. Two processes which extend from septum

contraction, close shell, to give support to posterior end of

lophophore; distal ends may also be

i Adjustor muscles. Two pairs of muscles in many prolonged into primary lamellae of spire or

articulate brachiopods attached to base of descending branches of loop (singular =

pedicle and responsible for moving shell

relative to pedicle, crus).

I Articulation. Interlocking of 2 valves by Crural plates. Shelly plates extending mediaty

from the crura, sometimes fusing to floor of

projections along their posterior margins; the dorsal valve.

I typically effected in Articulata by 2 ventral

teeth filling sockets of brachial valve. Delthyrium. Median triangular aperture in margin

of the ventral valve, wholly or partially used

i Beak. Extremity of umbo, commonly pointed, as the pedicle aperture.

Body cavity. Posterior coelomic space between Deltidial plates. Pair of shelly plates, growing

the 2 valves, enclosed by epithelium, medially from margins of delthyrium, partly

I cmuscles, ontainingmetanephridia, body organs (digestive system, etc.), or completely closing pedicle aperture.

Dental plates. Shelly walls supporting hinge teeth

i Braehial cavity. Anterior space between the from the floor of ventral valve.

valves, lined by mantle and body-wall

epithelium, containing lophophore. Diduetor muscles. The paired muscles which, on

contraction, open the shell by pulling on the

I cardinal process, situated on opposite side of

Braehial valve. See dorsal valve.

Brachidium. Calcareous support for lophophore the hinge axis from ventral areas of

in form of loop or spires, attachment.

I Caeca. Outgrowths of outer mantle epithelium Dorsal valve. Valve that contains skeletal support

contained in endopunctae of shell (singular for lophophore andnever wholly

I = caecum), accommoda than pedicletevsalve pedicl (= e;braccommonly hiaI valve).smaller

Cardinal process, Blade or variably shaped boss

of secondary shell situated medially in Endopunctae. Minute canals which perforate

I of brachial valve and interior of the shell but do not extend to posterior

end serving

for separation or attachment of diductor external surface; occupied by caecal

muscles, prolongations of the outer epithelium.

i Commissure. Line of junction between the edges Foramen. Aperature through which the pedicle

of the two valves, passes; typically confined to the ventral

I Co valve; aperature may be complete or

sta. Radial ridge on external surface of shell incomplete.

originating at margin of protegulal node or

any coarse rib. Hinge plates. A general term for skeletal structure

I C connected with the sockets and crura in ostate. Shell radially ornamented by costae, dorsal valve.

i Crenulated. Scalloped or notched valve margin. Hinge teeth. Two principal articulating processes situated at anterolateral margins of

delthyrium and articulating with dental

I sockets in brachial valve.
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II

Impunetate. Shell not perforated by minute Setae. Chitinous bristle arising from invagination II

canals or holes (see punctate), of muscle groove (singular, seta).

Loop. Delicate shelly loop-like support for Sockets. A pair of cavities, near posterior margin 1

lophophore, exending anteriorly from crura of dorsal valve, into which the teeth fit (= m

as closed apparatus; generally ribbon-like dental sockets). It

with or without

supporting septum from

Socket ridges. Ridges of shell, extending I

floor of dorsal valve (= brachial loop), anterolaterally from cardinal process,

Lophophore. Feeding organ with filiform bordering the inner side of tooth sockets on I

appendages, symmetrically disposed about dorsal valve. I

mouth; typically suspended from anterior

body wall but may be attached to dorsal Spicules. Small, irregular bodies of calcite II

mantle; occupies mantle or brachial cavity, secreted within connective tissue of body I wall, mantle and lophophore.

Mantle. Prolongation of body wall as fold of

ectodermal epithelium; secretes shell tissue. Spondylium. Trough-shaped or spoon-like •

apparatus complosed of dental plates in

Mantle canals. Flattened, tube-like extensions of various stages of coalescence, usually with

body cavity radiating into the mantle from median septum, l1

the body cavity (= pallial sinuses). Strophic. Shell with true hinge line parallel to l

Median septum. A vertically disposed shell plate hinge axis.

of variable height or length in median plane I

of either valve, normally present in body Suleate. Form of folding with dorsal valve bearing ==

cavity only. median sulcus; single sulcus in dorsal valve,

opposed by fold in ventral valve (reverse of I1

Palliul sinuses. See mantle canals, uniplicate). I

Pediele. Cuticle'covered stalk, commonly SulcipHeate. Type of fold involving a raised

protruding from ventral valve; attaches uniplicate fold with median sulcation I

animal to substratum; controlled by (= biplicate; reverse of intraplicate) I

muscles. Sulcus. Depression of valve surface, externally I

Pediele aperture. See foramen, concave in transverse profile and radial g

Pediele valve. See ventral valve, from umbo.

Teeth. Two principal articulatory

Periostraeum. The thin organic layer on exterior situated on ventral valve at aprntoerola cesses, teral I

of the shell, margins of delthyrium (= hinge tooth;

Protegulum. The first-formed valves secreted by singular, tooth). 1

the juvenile brachiopod; commonly Tubercles. Large nodose protuberances on shell

II

distinguished from later shell by absence of

surface.

ornamentation. Umbo. Poste I rior median, apical region of either

Punctate. Used to describe a shell perforated by valve, containing the beak. minute holes or canals known as mm

endopunctae. Uniplicate. Single fold in dorsal valve opposing a 1

sulcus in ventral valve (reverse of sulcate). Ribs. A form of external ornamentation in which

the shell surface is radially ridged; very f'me Ventral valve. Valve through which pedicle I

radial ridges termed striations, commonly emerges; usually larger than m

dorsal valve (= pedicle valve).

Septum. Relatively long, narrow elevation of

secondary shell, commonly blade-like. •

II

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I List of Species

I ClassOrdInarticulata Huxley, 1869 erLinguiida Waagen, 1885

Superfamily Linguloidea Menke, 1828

I Family Lingulidae Menke, 1828

Glottidia albida (Hinds, 1844)

I Order Acrotretida Kuhn, 1949

Suborder Craniidina Waagen, 1885

I Superfamily Cranioidea Menke, 1828

Family Craniidae Menke, 1828

I Neocrania californica (Berry, 1921)

i ClassOrder Artieulata RhynchHuxley, onellida 1Kuhn, 869 1949

Superfamily Rhynchonelloidea Gray, 1848

I Family Frieleiidae Cooper, 1959

Frieleia halli Dall, 1895

I Order Terebratulida Waagen, 1883

Suborder Terebratulidina Waagen, 1883

I Superfarnily Cancellothyridoidea Cooper, 1973

Family Cancellothyddidae Thompson, 1926

I TeTerebratulina rebratuIina cunguicula rossei Davi(Carpenter, dson, 18821865)

I Suborder Terebratellidina Muir-Woods, 1955 Superfamity Terebratelloidea King, 1850

Family Laqueidae Hatai, 1965

I Laqueus califomianus (Koch, 1848)

Family Dallinidae Beecher, 1893

I TerebrataIia occidentalis (Dall, 1871)

Terebratalia transversa (Sowerby, 1846)

i FamilyPlatidia Platidiidae hornii Th(Gabb, ompson1861) , 1927

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Key to Genera of West Coast Brachiopods I

IA. Shell flattened dorsal ventrally (depth less than one-half width) .................................................... 2 I

lB. Shell deep, not flattened (depth greater than one-half width) ......................................................... 4

2A. Pedicle absent; lower valve cemented directly to substrate ............................................. Neocrania I

2B. Pedicle present ................................................................................................................................. 3

3A. Shell large, elongate, lateral margins subparallet; pedicle very long; infaunal, in soft substrates ... I

............................................................................................................................................. GIottidia

3B. Shell small, round in outline; pedicle very short; foramen extending to brachial valve; epifaunal, or I

attached to hard substrates .................................................................................................... Platidia

4A. Shell thin, translucent ...................................................................................................................... 5 I

4B. Shell thick, solid, not translucent .................................................................................................... 6

5A. Shell inflated, shape globular to elongate, longer than wide; live in depths less than 500 m ....Laqueus I

5B. Shell flattened, shape triangular, as wide as long; live in depths greater than 500 m ......... Frieleia

I

6A. Shell usually longer than wide; anterior valve margin straight (without undulations) or very slightly

6B. Shell sulcate;usually beak large widr ....................................................................................................... than long; anterior valve margin with pronounced undulations; beak Terebratulina small .... I

....................................................................................................................................... Terebratalia

Descriptions of Species I

Class Inarticulata Huxley, 1869 I

Diagnosis. Valves not locked together posteriorly; hinge teeth and sockets absent; valves held I

together by muscles and body tissues. Lophophore spirolophous or zygolophous; never supported by

bra Embryological chidium. Peddevelopment icle absent ofor man moderately long; originating embryologically from ventral mantle. I tle direct without reversal. Functional anus present. Two groups of

muscles present -- adductors and obliques. Spicules absent in soft parts of body. Shell chitionophosphatic

or calcareous. 'l

Order Lingulida Waagen, 1885

Diagnosis. Valves thin, shape elongate, narrow; partly chitinous with calcium phosphate as principal I

inorganic constituent. Pedicle long, flexible; containing coelomic lumen.

I

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I Superfamily Linguloidea Menke, 1828

I Family Lingulidae Menke, 1828

Diagnosis. With characters of the order.

I Type Genus. Lingula Bmgui_re, 1797

Genus Glottidia Dall, 1870

I Diagnosis. Shell elongate, lateral margins gently convex to subparallel; thin, slightly thickened in

areas of muscle attachment. Pedicle valve with two low septa internally, diverging from beak and extending

I foprojec rwartdinto aboutma1ntle 13length cavityoffromvalve; manbrtale.chial valve with low median septum of similar length. Gill ampullae

Type Species. Lingula albida Hinds, 1845; by original designation.

I Remarks. similar but characterized the of internal

Externally to Lingula by presence septa.

I G/ott/d/a a/b/da (Hinds, 1844)

Figure 1.4 A-C

I Lingula albida Hinds, 1844: 71, pl. 19, fig. 4. --Sowerby, 1847: 339, pl. 67, fig. 6. --Davidson, 1852: 377.

--Reeve, 1859: pl. 1,fig. 4.---Carpenter, 1864: 636.----Cooper, 1867: 3; 1888: 246.--Yates, 1890:

39.--Thompson, 1927: 128.

I GIottidia aIbida: Dall, 1870: 157, pl. 8, figs. 1-6; 1873: 204; 1874: 296; 1877: 160-161; 1878: 28; 1889:

232; 1920: 267.--Davidson, 1888: 221, pl. 28, figs. 2-4.--Beecher, 1892: pl. 1,figs. 1-3.--Arnold,

I Snook, 1903: 941927: . --Kelse 153-154, y, 190fig. 7: 40. 127.--Hatai, --Berry, 1907: 1936: 18.78.--Burch, ---Oldroyd, 1942: t924b:1.221, Hertlein pl. 49,andfig.Gr3.ant, Johnson 1944: and 12-

16, text-figs, la-c, pl. 1, figs. 5-7. ---Smith and Gordon, 1948: 209. --Mattox, 1956: 76-77, 80, 83.

i --Hertlein and Grant, 1960: 89-90, pl. 19, figs. 17, 18, 23-25. --Jones and Barrtard, 1963: 27-52, fig. 1. --Ricketts and Calvin, 1968: 319, fig. 236. --MacGinitie and MacGinitie, 1968: 177-178,

fig. 55. --Zimmer and Haderlie, 1980: 110.

I Glonideafig.[si1. c] albida: Keep, 1904: 14, text-fig. 7; 1911: 19, text-fig. 9. --Keep and Baily, 1935: 19, textGlottidia palmeri: Dall, 1920:267-268 [in part]. --Burch, 1942:1-2. (Not Dall).

,! Material Examined. California: Santa Mafia Basin, Station 21, 1specimen, (24.2 mmTL), USNM;

4 specimens,(2.7-15.9 mm TL); Station 64, 59m; SBMNH.

I Additional material examined: 13 lots (see List of Additional Specimens Examined). Description. Shell smooth, thin; shape narrow and elongate. Valves similar in size, sides parallel to

subparallel. Pedicle long (longer than shell); fleshy. Color of shell - brownish white when alive or freshly

I dead, often with red-brown or brownish markings.

Shell size medium; total lengths to 35 mm; widths, 11-13 ram; depths about one-half width.

i Bay, 7 fmType[13Specimens m]. Not seen. and Type Locality. Holotype, BMNH (Paleontology) ZB. 1. MEXICO, Magdalena

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I

I

I

I

I

I

I

I

I

I

I

I

Figure 1.4. A-C, Glottidia albida (Hinds, 1844); depth unknown, San Pedro Bay, California (SBMNH [SSB

155]); 24.4 mm TL. A, dorsal view; B, lateral view; C, end view, anterior margin. D, E, Glottidia I

palmeri Dal11871; depth unknown, San Felipe, Baja California, Mexico (SBMNH 46125); 42.2 mm

TL. D, dorsal view, pedicle not visible; E, end view, anterior margin.

I

Distribution. Late Eocene to Recent. Eastern North Pacific, known to range from Tomales Bay,

California depths ranging (38°N)fromto the Todoslow Santos intertidalEstero, zone Btoaja150Califor m. nia, Mexico (26°N). Common in soft sediments at I

Etymology. Name derived from the Latin albidus, meaning "white" denoting the characteristic

color of the shell. I

Common Name. White tongue shell. Erroneously called a "tongue clam."

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I Biology. Glottidia albida live in vertical burrows in mud or soft sediments. The long fleshy pedicle

anchor the animal in its burrows. The anatomy of the mantle has been studied by Williams (1977) and

I aspects ofA the reproductive biology, especially oogenesis, are reviewed in Chuang (1983). small crepidulid snail, Crepidula glottidiarum Dall, 1905, is found on some valves of this

brachiopod (DalI, 1905; Hertlein and Grant, 1944; MacGinitie and MacGinitie, 1968; see also McLean,

I 1996, in the Taxonomic burrows in close Atlas, association Vol. 9). Thewith sipunculid, GlottidiaGolfingia albida (MacGinitie hespera (Chamberlain, and MacGinitie, 1919),1968). is reported Nitherto live th

specificity nor the biological aspects of these two relationships are known.

I Remarks. For the purposes of this paper the southern limits of the range of Glottidia aIbida are considered to be close to the tip of Baja California on the Pacific side. In the Gulf of California and south to

Panama G. albida is replaced by G. palmeri Dall, 1871 (see Figure 1.4 D, E). In the southern part of its range

I G. albida may be confused with G. palmeri from which it is distinguished by: 1) smaller maximum shell size; 2) very much smaller pedicle; 3) anterior margin of shell rounded not truncate; 4) anterior commissure

smooth without gaps between shell valves; 5) internal septa more divergent; and 6) whiter shell color (not

I br ofoMexico wn). Thecanrecor notdsbeofconfirmed. G. albida from The records the statesof ofGlottidia Jalisco, palmeri Oaxaca and and Guerrero G. audebarti alongBroderip, the mainland 1835 cfromoast

southern California most likely represent G. albida but locality-specific specimens need to be reexamined.

I' Order Acrotretida Kuhn, 1949

I and moreDiagconvex nosis. than Shellventral chitinophosphatic valve; apex ormarginal calcareous; to subcentral; absent in embryo. outline shape Dorsalcircular valve distinctly to subcircular. larger

Mantle lobe with or without setae. Pedicle absent or very short; if present it emerges through foramen on

I ventral spirolophous. valve; Statocysts without muscle absent. layer, but muscles fill coelomic lumen. Lophophore schizolophous or

I Suborder Craniidina Waagen, 1885

Diagnosis. Shell calcareous; densely punctate; bristles absent on shell margin; shell not present

I until postlarval stage. Dorsal valve conical; ventral valve relatively flat; conforms to shape of substrate to which it is entirely cemented; no gap between valves. Setae absent on mantle edge. Pedicte absent. Lophophore

spirolophous. Muscles - 2 pairs adductors + 2 pairs obliques.

I

Superfamily Cranioidea Menke, 1828

I Family Craniidae Menke, 1828

i Dorsal valve Diagnusually osis. Shell conical, calcare ventral ous;valve punctate, subconlcal punctaor convex in dorsal whenvalve free,branching conformingintosomshape e recent of attachment genera.

surface when fixed. Pedicle absent. Marginal sinus absent. Marginal setae absent.

I Type Genus. Crania Retzius, 1781

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Genus Neocrania Lee and Brunton 1986 I[

Diagnosis. Shell of medium size, subcircular to quadrangular in outline. Ventral valve cemented to m

substmte by entire surface; valve varying from thin, uncalcified organic film to wholly calcified and thickened 1

with antemlateral rim. Valve interior with sometime sunken posterior muscles scars with anterior scars

II

united medially. Dorsal valve smooth, slightly pustulose or finely ribbed, umbo centrally to posteriorly

placed. Valve interior with large, widely separated pad-like posterior muscle scars and smaller diverging V- I

shaped anterior scars. Weak posterior submarginal rim, internal surfaces strongly endopunctate.

Type Species. Patella anomala Miiller, 1776; by subsequent designation. _,

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Neocrania californica (Berry, 1921) l

I

Figure 1.5

Crania californica Berry, 1921:210-212, pl. 11, figs. 1-3.--Thompson, 1927: 136.--Keep and Baily, I

1

1935: 20. --Hatai, 1936: 80. --Burch, 1942: 3. --Hertlein and Grant, 1944: 40, pl. 3, figs. 3, 5, 6.

--Bernard, 1972:75, fig. 2. --Kozloff, 1987: 419-420, fig. 20.1. I1

Neocrania californica: Lee and Brunton, 1986: 151. II

Material Examined. Holotype - California, Los Angeles Co., Santa Monica, from rock at base of m

siliceous sponge, 100 fm [183 m]; coll. fishermen, summer 1918; SBMNH 34501 [SSB 4530]. II

Additional material examined: 2 lots (see List of Additional Specimens Examined).

i

Description. Shell shape subrectangular to rounded-oval, strongly depressed. Ventral valve thick, I [

flat; entirely cemented to substmte, margins free, flared outward; interior with strong submarginal thickenings. I

Dorsal valve smooth, small apex situated subposteriorly. Interior of shell microscopically granulose. Adductor

attachment scars prominent; 4 in number, diamond-shaped. Color of shell - exterior whitish brown; interior I

brownish white. I

Shell size relatively large; total lengths to 20.0 mm; widths to 23.0 ram; depths less than one-fifth

width, i

Type Specimens and Type Locality. Holotype, SBMNH 34501. USA, California, Santa Monica,

100 fm [183 m].

Distribution. Recent. Eastern North Pacific known from type locality and off shore banks (32°N; l

CASIZ collections) to British Columbia, Canada (51°N; Bernard, 1972). Lives attached to hard substrates at

depths ranging from 120-200 m. I

E I tymology. Named after the State of California.

Common Name. None.

1

Biology. Attached to hard substrate. 1

Remarks. This is the only locally occurring brachiopod in which the ventral valve is cemented

directly to the substrate. It is most similar to Crania patagonica Dall, 1902 from the coast of Chile and the •

Straits of Magellan in South America but lacks the short spines on the dorsal valve. II

The species rarely has been reported in the literature and is very uncommon in west coast museum

collections. Kozloff (1983) reported the species to be abundant in the subtidal off the open coast in the I_

northern part of its range. It appears to be either sporadic in occurrence or lives in rock or large cobble II

habitats which are seldom sampled.

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i Figure 1.5, Neocraniacalifornica(Berry, 1921). A-D, Holotype, 180m, off SantaMonica, Califomia(SBMNH

34501); 13.0mmTL. A, dorsal view; B. lateral view; C. end view, anterior margin; D, interior view

i of dorsal valve. E, 124 m, CordellBank, California (CAS 105948); 20.0 nun TL. E, dorsal view.

I Class Articulata Huxley, 1869

i Diagnosis. Shell composed of scleroproteins and calcite; shell absent until postlarval stage. Valves locked together posteriorly with articulated hinge; hinge teeth and sockets present. Lophophore supported

by skeletal structure (brachidium). Pedicle present; originates from foramen in ventral valve; distal end may

i be branched; internal muscles in pedicle absent; coelomic lumen in pedicle absent. Adductor muscles present; oblique muscles absent. Functional anus absent; intestine ends blindly. Spicules absent or present in soft

parts of body. Coelom formation enterocoelous; mantle reversal occurs after settlement. Epifaunal.

I Order Rhynchonellida Kuhn, 1949

Diagnosis, Shell small to moderate size; typically impunctate. Pedicle not used in locomotion.

I Lophophore spirolophous, with dorsally directed cones; partly supported by crura. Mantle canals with 2

main trunks in each mantle lobe. Dental plates typically present. Spicules absent. Metanephridia present, 1

or 2 pairs. I

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Superfamily Rhynchonelloidea Gray, 1848 U

Family Frieleiidae Cooper, 1959 I

Diagnosis. Shell small to medium size. Pedicle relatively short; as broad as foramen. Intestine

straight without terminal enlargement. Dorsal median septum present as low thick ridge or absent. Cmra •

short, straight, spinuliferous, supported by short plates.

Type Genus. FrieIeia Dall, 1895.

Genus Frieleia Dall, 1895

Diagnosis. Shell thin; outline elongate oval to subtriangular; greatest width at or anterior to middle; 1

both valves may be somewhat sulcate; surface smooth to minutely costellate. Ventral valve with greater

depth and convexity; beak short, nearly straight to suberect; foramen incomplete, elongate oval; deltidial

plates thick, disjunct but nearly united; apex marked by small triangular plate elevated above valve floor. •

Ventral valve interior with long, curved corrugated teeth buttressed by prominent dental plates; muscle and

pallial marks lightly impressed. Dorsal valve interior with deep corrugated sockets marginate by thick

socket ridges; crura of spinulifer type, long, thin, divergent, attached directly to socket ridges; inner hinge _•

plates small, rounded, disjunct; media septum long, slender, elevating posteriorly and united to inner hinge

plates forming small V-shaped chamber. Median septum rising to crestjust anterior to apex; cardinal process

small, triangular, transversely striated pit at apex; adductor scars long, narrow, posteriorly situated. I

Type Species. Frieleia halli Dall, 1895; by original designation.

Fr/e/e/a ha///Dall, 1895 I

Figure 1.6

Frieleia halli Dall, 1895:714, pl. 24, figs. 6, 9-13; Dall, 1920: 292-293.--Kelesy, 1907: 40. --Jackson, i

1918: 192, pl. 1, fig. 8.--Oldroyd, 1924a: 224. ---Thompson, 1927: 157-158, text-fig. 48. Nomura

and Hatai, 1934: 16. --Keep and Baily, 1935: 23-24, text-fig. 7. --Hatai, 1936: 225, fig. 24; Hatai, I

1940: 219, text-fig. 21. --Burch, 1942: 4. --Hertlein and GranL 1944: 58-59, text-fig. 11, pl. 4,

figs. 6, 7. ---Smith and Gordon, 1948: 209. ---Cooper, 1959: 54; Cooper, 1969: pl. 2, fig. 19, pl. 5,

figs. 18, 19.--Bernard, 1972:77, fig. 7. I

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Hemithryris psittacea: Keep, 1904, fig. 6.--Keep and Baily, 1935:23 [in part]. (not Gmelin).

Material Examined. 14 lots (see List of Additional Specimens Examined). I

m

Description. Shell shape subtfigonal, smoothly rounded or with medial depression; anterior margin

bilobed. Valves extremely thin; smooth; translucent to transparent. Color of shell light yellow, yellowishgray or colorless. 1

Ill

Shell size medium; total lengths, 18-20 mm; widths, 16-18 mm.

Type Specimens and Type Locality. Holotype, USNM 123148. USA, Califomia, Cortez Bank. I

Distribution. Miocene to Recent. Eastern North Pacific, known from Aleutians and Gulf of Alaska

(58°N; Dall, 1920) to San Diego, California (32°N; SBMNH collections). Western North Pacific, reported

from Japan and Kamchatka. Collected from 40-2200 m, typically occurring between 500-1000 m. 1

Etymology. Named in honor of James Hall, a noted brachiopod expert.

Common Name. Hall's lamp shell. I
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Figure 1.6. Frieleia haUiDall, 1895:1150-1170 m, off San Diego, California (SBMNH [SSB3038]); 17.7mm

I TL. posterior A, dorsal margin. view; B, ventral view; C, lateral view; D, end view, anteriormargin; E, end view,

!

Biology. Exclusively attached to organic substrates, i.e., dead bivalve shells, living sponges,

I polychaete tubes and pogonophoran tubes. Valentine and Ayala (1975) documented genetic variation and

Rokop (1977) discussed seasonal reproduction ofFrieleia halli in the deep sea. A number of dead specimens

examined for the latter paper had tiny holes in the shell, and probably had been drilled by a predator such as

I an octopus (see fig. 1.6A). Nothing additional has been reported on the biology of this species.

Remarks. Frieleia halli is a northern boreal species that was formerly considered to be the young

ofHemithryspsittacea (Chemnitz, 1785) from which it differs principally in its lighter color. Records of H.

I psittacea from south of Oregon are probably Frieleia halli. In addition, young F. halli have been confused

with Cryptophora hesperis Cooper 1982, a cryptoporid known only from the type locality off southern

Oregon in 170 m. Cryptophora, in contrast to Freilia, has unmodified marginal deltidial plates, a gently

I sulcate anterior commissure and the outline shape is rounded. Fossil brachiopods from the Tertiary of

California and Oregon, presently referred to Hemithrys may belong in the genus Freilia.

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Order Terebratulida Waagen, 1883 •

Diagnosis, Shell small to large; punctate; fight colored often with bright colors. Pedicle normal and i

functional, vestigal and non-functional, or absent. Mantle canals with 2 or 4 main trunks in each mantle i

lobe. Dental plates present or absent. Metanephxidia present, 1 pair. Lophophore in call configurations, i

supported by calcareous loop. Spicules present or absent.

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Suborder Terebratulidina Waagen, 1883 II

Diagnosis. Shell small to large. Dorsal valve lacks median septum. Lophophore supported by short B

skeletal loop; developed directly from cardinal process. Pedicle distal end often split into large rootlets.

Spicules commonly present. Gonads reticulate, located in pair of pillared spaces in each mantle lobe. •

Remarks. There are no representatives of the Superfamily Terebratuloidea off California. The II

Superfamily Cancellothyridoidea is represented by two species of Terebratulina.

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Superfamily Cancellothyridoidea Cooper, 1973 i

Diagnosis. Shell size small to moderately large; costellate to costate or secondarily smooth; color fi

drab to bright. Dorsal pedicel adjustor muscles attached to dorsal valve floor. Lophophore spirolophorous

through pletophorous. Dental plates rarely developed. Cardinal process weak to moderately developed.

Hinge plate absent; socket ridges present. Dorsal median septum absent. Crura united to form ring or open. I

Family Cancellothyrididae Thompson, 1926 i

m

Diagnosis. Shell size medium to moderately large; costellate or capillate; punctate with externally

branching pores. Dental lameltae absent. Hinge plates absent; socket ridges fused with cmral bases; cardinal •

process developed. Deltidial plates disjunet. Pedicle very short to moderately long, muscles attached to

floor valve. Loop short, crural processes united to form ring with transverse band. Lophophore typically

plectolophous, may be spirolophous or subplectolophous. Spicules present, i

Type Genus. TerebratuIina d'Orbigny, 1847

Genus Terebratulina d'Orbigny, 1847 Ii

Diagnosis. Shell outline oval or elongate, inflated; finely striated; auriculate; deltidium usually

rudimental; foramen incomplete. Loop short, appears annular in adult through union of oral processes; crura •

forming wide ring; lophophore plectolophous.

Type Species. Anomia caputserpentis Linn6, 1767; by original designation.

I

Key to Species of the Genus Terebratulina

1A. Shell smooth or with faint riblets; anterior valve margin smooth; shape round to elongate; umbo i

orbicular ........................................................................................................... Terebratulina crossei

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lB. Shell with conspicuous ribs; anterior valve margin crenulated; shape distinctly triangular; umbo I

narrow ......................................................................................................... Terebratulina unguicula

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I Terebratulina crossei Davidson, 1882

Figure 1.7

I Terebratulina crossei Davidson, 1882:106, pl. H-7, fig. 1.--Dall, 1910: 96; Dall, 1920: 303. --Burch,

1942: 5-6. ---Bernard, 1972:77, 79, fig. 9. --Roth, 1972: 5.

I Terebratulina crossii [sic] Davidson, 1886: 33, pl. 3, figs. 4-6.

Terebratulina crossei not Davidson. --Fischer and Oehlert, 1892: 5, pl. 8, figs. 1-8.

I TerebratuIina 32, figs. kiiensis 8, 9.--Dall, not Dall1894: and 720, Pilsbry. pl. 32, --Dall figs.and8, 9;Pi1920: lsbry, 307. 1891----Oldroyd, b: 152, pl. I1924a: 1,figs. 203, 9, 10;pl.1895: 29, figs. 720,13- pl.

14.-Keep and Baily, 1935: 23, fig. 6.--Hatai, 1936: 100-101; 1938:112, 128, pl. 19, figs. 5, 6, 8,

I 71, 9.Nomura text-fig.and16,Hatai, pl. 6, 1figs. 937: 11, 141,12,pl.pl.21,17,fig.figs. 17.--Burch, 3, 6, 10. --Smith 1942: 5.and --Hertlein Gordon,and1948: Grant, 209.1944: --Roth, 68-

1972: 8.

I Terebratulina (unquicula var.) Kiiensis Dall and Pilsbry, 1891a: 18, pl. 1, figs. 4, 5, 1894: 720, pl. 32, figs. 8, 9. Terebratulina kiiensis Dall and Pilsbry. --Dall, 19t0: 96, 1920: 307. ---Oldroyd, 1924a: 203, pl. 29, figs.

13-14, 1924b: 226, pl. 16,figs. 13, 14. Keep and Baily, 1935: 23, text-fig. 6. --Hatai, 1940: 235, pl.

' 7, figs. 18-21.--Hertlein and Grant, 1944: 71-73, text-fig. 18a-c, pl. 6, figs. 10, 13 [in part]. -- Nomura and Hatai, 1983: 63. --Zezina, 1970: 443, pl. 1, fig. 3. --Roth, 1972: 8. ----Cooper, 1982:

6-8, pl. 1, figs. 8-18.

I and USNM 128463. JAPAN, coast of Kii Province. Not Type Specimens Type Locality: Holotype 1

seen.

i Terebratulina species. ---Cooper, 1982: 8, pl. 1, figs. 5-7.

Material Examined. California: Santa Mafia Basin, Station BRA-16, 591m; 17 specimens, 4.3-

37.5 mm TL; USNM.--3 specimens., 12.7-30.3 mm TL; SBMNH.

I Additional material examined: 28 lots (see List of Additional Specimens Examined).

Description. Shell thin; shape elongate to nearly circular; rostrum inflated, very short; smooth or

. with sculpture of fine radial riblets; anterior valve margins smooth. Color of shell waxen white, often with brownish stains.

Shell size medium; total lengths, reported to 44 ram, observed to 37 mm; widths, reported to 40

I mm, observed to 33 mm. Type Specimens and Type Locality. Types in BMNH; not searched. JAPAN, Sagami Bay.

Distribution. Miocene to Recent. A boreal species known to range from Japan to southern California

I and possibly asfar south as Peru and Chile. Eastern North Pacific, from Vancouver Island, British Columbia,

Canada, (5 I°N; Bernard, 1972) to La Jolla, California (32°N; SBMNH) and the California Islands.--Westem

North Pacific, Japan. Depth off California reported to range from 100-450 m.

i Etymology. Named in honor of T. Crosse, a noted brachiopod expert.

Common Name. Cross's lamp shell.

I Biology. The species appears to be solitary and is considered rare in its occurrence. Nothing is reported in the literature about the biology of this species.

Remarks. Under the synonymy for TerebratuIina crossei I also reviewed references to California

I records of T. kiiensis. Terebratulina crossei almost identical to T. kiiensis Dall and 1891 appears Pilsbry,

(see Figure 1.7F). When comparing the two species T. kiiensis has: 1) a larger shell size; 2) a flatter and

more discoid shell shape; 3); a coarser shell sculpture with larger, more conspicuous longitudinal fibs; 4) a

I darker, more brownish color when fresh; and 5) lives in deeper water.

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Figure 1,7. A-E, Terebratulina crossei Davidson, 1882; 88 m, off Santa Monica, California (SBMNH); 30.7 I

mm TL. A, dorsal view; B, ventral view; C,lateral view; D, end view, anterior margin; E, end view,

posteriormargin. F, Terebratulina kiiensis Dalt and Pilsbry, 1891; 183 m, off La Jolla, California I

(SBMNH [SSB 3265]); 31.8 mm TL. F, dorsal view.

After examining a large number of lots from California I cannot reliably separate the two species. I

Both species are widely distributed in the Eastern North Pacific Ocean, reportedly ranging from Alaska to

Calif of T ornia (Dall and Pilsbry, 1891; Dall, 1920; present study). Cooper (1982) additionally noted the presence I . kiiensis in South America. Cooper (1982: pl. 1, figs. 5-7) illustrated a single specimen from 439 m off

Santa Barbara which he identified as intermediate between Terebratulina crossei and T. kiiensis. In the next

sentence seen in otheherindic brachiopod ated thatspecies the specimen (principally "seemsshell to beshape a newandspecies." sculpture) In lighit would t of theappear morphologic that theal validity variationsof I

the closely related crossei-kiiensis complex should be critically reevaluated. For the purposes of this Atlas

Idescribed have chosen of theto two use the species. name T. crossei for all California specimens in this complex because it is the first I

In comparison with Terebratulina unquicula the shells ofT. crossei are: 1) considerably larger, and

proportionally wider and deeper; 2) smoother; and 3) the beak proportionally shorter. I

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I Terebratulina unguicula (Carpenter, 1864)

Figure 1.8

I Terebratula unguiculus Carpenter 1864: 607.

i Terebratula unguicula Carpenter. Carpenter, 1865: 201, text-figs. 1-4.--Cooper, 1867: 3.--Dull, 1870:

102.

Terebratulina unguicula (Carpenter). ---Dull, 1873: t80; 1877a: 155; 1920: 300-301. --Whiteaves, 1878:

I 468.--Dull 27, figs. 2-5;and1924b: Pitsbry, 226,1891: pl. 38,19,figs. pl. 2-5. 1,figs. --Johnson 6, 7.--Du andBois, Snook, t916:178.- 1927:--156, Oldroyd, text-fig.1924a: 130. --Hatai, 203, pl.

1936: 83. --Burch, 1942: 5. --Hertlein and Grant, 1944: 79-81, pl. 5, figs. 9, 10, pl. 6, figs. 1, 6, 7,

I pl. 79, 17, figs. 9, 13, 14, 18, 19. --Smith and Gordon, 1948: 210. --Mattox, 1956: --Bernard, 1972: fig. 8.--Roth, 1972: 14-15.

Terebratulina unguiculata [sic] (Carpenter): Davidson, 1880: 14.

I Terebratulina caput-serpentis vat. unguiculata (Carpenter): Davidson, 1886: 25-26, pl. 5, figs. 38, 40. -- Kelsey, 1907: 50.

Terebratulina caput-serpentis not Linnaeus: Cooper, 1894: (pages not numbered). --Dull, 1895: 719, pl.

I 32(figs. 2, 5). --Kelsey, ---Willett, --Keep Baily, 22-23, fig.

1907: 50. 1918: 66. and 1935: 5.

Material Examined. California: Santa Mafia Basin, Station BRA-21, 75-90m, 27 spec., 2.4-18.5

I mm TL, USNM; 54-63m, ca. 25 spec., 2.3-18.8 mmTL, SBMNH.

Additional material examined: 25 lots (see List of Additional Specimens Examined).

i Description. Shell thick; shape ovate to triangular, longer than wide; valves almost equally convex, rounded laterally and anteriorly; lateral sides of umbo strongly auricular, especially in young and mediumsize specimens. Anterior commissure straight, regularly crenulated; lateral commissure straight or slightly

i sinuous, crenulated. Beaks short; obliquely truncated by relatively large, incomplete foramen; margined anteriorly by umbo of dorsal valve and by 2 small lateral deltidia. PedicIe short. Valve surface marked by

numerous prominent radiating fibs, simple and stronger at their origin, some dichotomously branched;

i number rapidly increases from bifurcations and by interpolation with shorter riblets between larger ones; valves crossed with delicate concentric raised striae. Ventral valve evenly convex, occasionally slightly

depressed anteriorly. Loop short and simple; crura disunited in young, annelidiform in larger specimens.

Color of shell light yellowish-white to greyish-ash.

I Shell size small; maximum total lengths, reported to 25 mm, majority observed measure less than

15 mm; widths, reported to 22 mm, observed to 10 mm.

I USA, California, Type Specimens Monterey.and Type Locality. Types reported to be in the Cumings Museum; not searched.

Bernard (1972) erroneously reported the type locality to be off Vancouver Island, British Columbia,

I Canada.

Distribution. Miocene to Recent. Eastern North Pacific, Priblof Islands, Alaska (57°N; Dull, 1920)

to Cabo San Lucas, Baja California Sur, Mexico (23°N; Dall, 1920).--Western North Pacific, reported

I from the Kamchatka Peninsula.

Depths typically range from 10-850 m; in the northern part of the range the species may be locally

abundant intertidally in areas of broken rock and coralline algae.

I Etymology. From the Latin word unguicula meaning having "claws" in reference to the characteristic

radiating ribs.

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Figure 1.8. Terebratulina unguicula (Carpenter, 1865). A-E, 132 m, off Gaviota, California (SBMNH), 18.3

nun po TL. A, dorsal view; B, ventral view; C, lateral view; D, end view, anterior margin; E, end view, I sterior margin. F, depth unknown, off La Jolla, California (SBMNH [SSB 13507]); 17.0 mm TL.

F, two specimens attached to solitary coral.

I

BCommon Name. None. l iology. This species typically nestles inside dead bivalve and gastropod shells or in the hollow

spaces in dead hexactinellid sponges. It also has been found attached to the hairy periostracum of the large

gastropod Fusitriton oregonensis and to the corallum of solitary corals such as Paracyathus stearnsii (see I Figure 1.8F).

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I Various aspects of the biology of Terebratulina unguicula have been extensively studied, especially

in Washington and Canada where live specimens are easily obtained. Reproduction and development have

i been 1987)reviewed by Reed (1987). Details of embryology are treated by Long (1964). Wilkens (1978a, b, investigated the physiology of muscle activation and contraction. The orientation of this brachiopod

to water currents and functional aspects of the tophophore were examined by LaBarbera (1978, 1981, 1986).

I Fouke (1986) documented the functional significance of spicule-reinforced connective tissues. A number of papers have examined details of distribution and ecology, especially in populations living in British Columbian

fjords which have low ocygen concentrations and high turbidity flows (see Thayer, 1981; Farrow et al.,

1983; Tunnicliffe and Wilson, 1988).

I Remarks. According to Bernard (1972) the records of Terebratulina unguicula from the Western

North Pacific probably represent T. crossei. This shells of this species are distinguished from T. crossei by:

I valve 1) theirmargins. smaller maximum size; 2) narrower umbo; 3) distinct sculpture of large, coarse ribs; and 4) crenulated

The close resemblance of Terebratulina unguicuIa to fossils specimens of T. tejonensis Stanton

I from the Eocene of California warrents additional study.

I Family Laqueidae Hatai, 1965

Diagnosis. Shell size medium to moderately large; shape biconvex; smooth to costate; posterior

I margin parts ofnon-strophic. loop. Hinge Dental plates present. plates present. SpiculesBrachidia typicallywithabsent cross(present union between only indiscending Laqueus). and Lophophore recurved

plectolophous. Cardinal process small or absent.

' Type Genus. Laqueus Dall, 1870.

i Diagnosis. Shell thin, large, inflGen ated;us hinge LaqueuslineDalcurved, l, 1870beak prominent; pedicle foramen small,

complete. Brachidium long.

I Type Species. Terebratula californiana Koch, 1848; by original designation.

I Laqueus cah'fornianus (Koch, 1848) Figure 1.9

I Terebratula Koch, 1848:38 + 1844: 2b, 21-23. --Davidson, t 852: 364.

californiana pl. figs. ----Carpenter,

1864: 568, 574.

Terebratula Waldheimia californica californica:: Sowerby, Gray, 1853:1847: 60.---Carpe 352, pl.nter, 70 figs. 1856: 50,298; 51. 1864: 636.

Laqueus californica: Hatai, 1936: 90.

Laqueus californicus: Dall, 1870: 123, pl. 7, fig. f, pl. 8, figs. 9, 10; I894: 725; 1920:344 [in part]. --

Davidson, 1887: 112, pl. 18, figs. 6-9. --B eecher, 1893b: 383, pl. 1,fig s. F4, G4.--Lowe, 1904:19.

--Berry, 1907: 18. --Arnold and Anderson, 1907: 153. Kelsey, t907: 41.--Oldroyd, 1924a:

205, pl. 28, figs. 1-4, 1924b: 230, pl. 17, figs. 1-4. --Thompson, 1927: 258, fig. 85. --Johnson and

Snook, 1927: 157, fig. 129. --Keep and Baily, 1935: 20-21, fig. 2 [sic Laqueas]. --Nomura and

Hatai, 1937: 187-188, pl. 25, figs. 5-10. --Burch, 1942: 8. ---Smith and Gordon, 1948: 210. --

MacGinitie and MacGinitie, 1968: 178, fig. 56.

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Laqueus californianus: Hertlein and Grant, 1944: 144-147, text-fig. 31, pl. 13, figs. 5, 8, 14, pl. 21, figs. 1- •

7; 1960: 96-97, pl. 20, figs. 1-3, 5-7. --Mattox, 1956: 75, fig. 2. --Bernard, 1972: 81, fig. 1.

Kozloff, 1987: 419-420, fig. 20.2. •

Terebratula kochii K_ster in Martini-Chemnitz, 1848: 39, pl. 2d, figs. 1-3. --Dall, 1920: 348.

Type Specimens and Type Locality: Not searched. m

Laqueus erythraeus Dall, 1920: 350.---Oldroyd, 1924b: 231-232. --Thompson, 1927: 259.----Hatai, 1936: 1

90. ---Cooper, 1969: pl. 1, fig. 2.

all

Type Specimens and Type Locality: Holotype,wUSNM 19395. California, off Santa Catalina Island, •

80 fm [146 m]. Not seen. l

Megerlia Jeffreysi Dall, 1877: 187. wWhiteaves, 1878: 468. _Beecher, 1893: 383.

Type Specimens and Type Locality: Types not searched. British Columbia, Vancouver. I

m

FrenuIa Jeffreysi: Dall, 187 l a: 55 [in part]. (not Dall).

Ismenia Jeffreysi: Dall, 1871d: 65-66, pl. 11, figs. 7-10, 1887: 65, pl. 11, figs. 7-10 [in part]. (not Dall). •

Laqueusjeffreysi (Dall). --Dall, 1894: 725. --Lowe, 1904: 19. --Arnold, 1907: 425. --Du Bois, 1916: l

178. --Willett, 1918: 66. --Keep and Baily, 1935: 21.

Laqueus californicusjeffreysi (Dall). --Oldroyd, 1924a: 205-206. n

i

Terebratula californianus forma vancouveriensis Davidson, 1877: 113, pl. 18, figs. 10-13, 13a, 13b.

Type Specimens and Type Locality: Types not searched. British Columbia, off Vancouver Island. •

Laqueus californicus vancouverensis [sic] (Davidson). ---Oldroyd, 1924a: 206, pl. 28, figs. 5-10. ---Oldroyd, I

1924b: 231, pl. 17, figs. 5-10. Thompson, 1927: 259.

Laqueus vancouverensis [sic] (Davidson). --Hatai, 1936: 90. _Burch, 1942: 8. H

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Laqueus vancouveriensis (Davidson). --Hertlein and Grant, 1944: 147-151, text-fig. 32, pl. 13, (figs. 6, 7,

9, 10, pl. 17, figs. 15-17, pl. 18, figs. 15-17, 19-21. --Bernard, 1972:81, fig. 5. --Roth, 1972: 15. I

Laqueus vancouveriensis diegensis Hertlein and Grant, 1960: 97-98, pl. 20, figs. 4, 8, 21.--Roth, 1972: 5. i

Type Specimens and Type Locality: Holotype,----CASIP 7355. California, San Diego, Pacific Beach

[Pliocene]. I

Material Examined. 21 lots (see List of Additional Specimens Examined).

Description. Shell thin; inflated; shape ovoid to nearly circular, length slightly to much greater than •

width. Dorsal valve uniformly convex, with slight tendency to depression close to frontal margin. Ventral

valve slightly deeper than dorsal valve, sometimes with slight indication of masial longitudial elevation or

flattened fold. Beaks curved; truncated by small circular foramen; margined anteriorly by 2 wide delthidial •

plates; beak-ridges sharply def'med, a distinct flattened space present between ridges and hinge-line. Pedicle l

elongate, long and narrow. Valve surfaces smooth, with faint concentric growth lines. Crura short, curved.

Color yellowish-brown to light reddish-brown to almost white; often with darker markings where mantle •

canals show through thin shell. I

Shell size large; maximum total lengths to 53 ram; widths to 42 mm; depths to 25 mm.

Type Specimens and Type Locality. Types not searched. California. 1

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Distribution. Pliocene to Recent. Eastern North Pacific, reported from the Cook Inlet, Alaska (59°N)

to La Jolla, California and the offshore islands (32°N; SBMNH). Western North Pacific, from Sea of Japan. m

Depths reported from the intertidal to 1500m; abundant intertidally in northern part of range, lill

Etymology. Named for the State of California where the species was first found.

Common Name. California lamp shell •

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l Biology. One of the most abundant mid-depth species in southern California. It occurs in dense

populations often with individuals attached to each other forming large clusters. Found on clean rock or

I gravel boThe ttoms. Many shells examined appear to have been drilled by Octopus or other molluscan predators. biology ofLaqueus californicus has been extensively studied throughout its range. The functional

morphology of the lophophore, especially in relation to food collection and waste rejection, have been

I dTheocuomented rientationby Strathman of Laqueus(i 973), to water Reynolds movements and McaCammo nd the ndynamics (1977), Gofilmour inhalent (1978and), and exhalent LaBarbera currents(1986). was

elucidated in LaB arbera (1978, 1981). Clarke and Wheeler (1915) analyzed the chemical components of the

i she thosell and of Wilkens Mackay (1978a, and Hewitt b, 1987) (1978)whoexamined investigated the ultrastructure the activationof the andpedic contraction le. Physiological of adductorstudies and diductor include

muscles and Scheid and Awapara (1972) who investigated metabolism.

l low oxygen The concentrations presence of Laqueus and highpoputurbidity lations lflows iving inhasfjorbeends indocumented British Columbia by Thayer which(198areI),known Farrowto ehtaalve.

(1983) and Tunnicliffe and Wilson (1988). The relationship of these ecological conditions to the

I vancouveriensis ecomorph is discussed. Remarks. There is some confusion as to the status of the rounder and smaller Laqueus

vancouveriensis (Davidson, 1877) which typically is found in quiet waters on rocks mixed with mud. Hertlein

I and Grant (1944) considered the differences sufficient to warrant separation at the species level, whereas, Bernard (1972) considered L. vancouveriensis to represent a quiet water ecomorph. In comparison with L.

californianus, the vancouveriensis morph is characterized by: 1) a larger foramen; 2) a smaller shell size; 3)

I valves which are flatter and thicker; 4) an olive drab or yellowish-white color vs pinkish-white. In addition, the animals are reported to live singly or in small clumps versus large, dense colonies. Reports in the earlier

literature of L. vancouveriensis in California should be refered to L. californianus. Laqueus jeffreysi is a

I senior synonym of this species but this name has not been commonly used in the literature. Dall considered this the young of the former species (see Oldroyd, 1924b).

I Suborder Terebratellidina Muir-Woods, 1955

I skeletal Dloop; iagnosdeveloped is. Shell small with both to large. cardinal Dorsaprocess l valve with and memedian dian sesepptum. tum. Lophophore Pedicle distalsuppported ends seldomby lsplit ong t

into rootlets. Spicules rarely present. Gonads not located in pilleard mantle. I

I Superfamily Terebratelloidea King, 1850

I Family Dallinidae Beecher, 1893

Diagnosis. Loop passing through all or part of pre-campagiform, campagiform, frenuliniform,

I terebrataliiform, and dalliniform growth stages, or modifications of these. Spicules in some forms

present

but never abundant. Dental plates typically present.

i Type Genus. DaUina Beecher, 1893

Genus Terebratalia Beecher, 1893

I Diagnosis. large, or ribbed, margin sulcate; incomplete;

Shell smooth anterior deltidium foramen

large. Punctae developed. Beak short, suberect. Hinge teeth with poorly developed dental plates. Brachidium

highly developed and complex; attached to septum.

i Type Species. Terebratula transversa Sowerby, 1846; by original designation.
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Figure 1.9. Laqueus 45.7 mm californ TL. A,ianus dorsal (view; Koch,B1,848): ventralA-E, view;90 C, m, lateral off Santaview; Catalina D, endIslan view, d, Ca anterior liforniamargin; , (SBMNH); E, end I

view, posterior margin. F, 155 m, off Morro Bay, California (SBMNH [SSB 18432]); 42.5 mm TL.

F, dorsal view. I

Key to Species of Terebratalia I

1A. Anterior commissure uniplicate, medial flexure of dorsal valve convex; shell typically slightly I

longer than wide; with few, broad, low-relief ribs .................................... Terebratalia occidentalis

1B. Anterior commissure sulcate, medial flexure of dorsal valve concave; shell wider than long; smooth

or with numerous, distinct, narrow ribs ...................................................... Terebratalia transversa i

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Terebratalia occidentalis (Daft, 1871)

i Figure 1.10

TerebrateIla occidentalis: Dall, 1871a: 182, pl. 1,fig. 7, 1891: 172-173, pl. 4, figs. 8, 9 only.--Davidson,

1887:79-81 [in part], pl. 16, fig. 13 only. --Lowe, 1904: 20.

I Terebratalia occidentalis: Dall, 1894 [1895]' 729, pl. 31, figs. 7, 8; 1920: 341.--Kelsey, 1907: 50. Eldridge

and Arnold, 1907: pl. 39, figs. 6-9. --Arnold, 1907: 534, pl. 49, figs. 6-9a. --Arnold and Anderson,

I 1907: pl. 22, figs. 4a, b.--Oldroyd, 1924a: 229. --Hertlein and Crickman, 1925: 267, 272. --Keep and Badly, 1935: 21, fig. 3.--Burch, 1942: 7.--Hertlein and Grant, 1944: 127-131, text-fig. 27, pl.

10,fig. e, pl. 17, figs. 2, 5, 8, 11, 12; 1960: 94-95, pl. 19, figs. 16, 22, 31.--Smith and Gordon, 1948:

210.--Mattox, 1956: 75-76, figs 4A-M. --Roth, 1972: 9.

I Dallinella occidentaIis: Thompson, 1927: 246.--Woodring, 1930: 60.--Hatai, 1936: 89; 1940: pl. 9. igs.

22-24, 26, 27).--Nomura and Hatai, 1937: 185-187, pl. 25, figs. 15-19.

I TerebratelIa occidentalis var. obsoleta Dall, 1891: 186. Type Specimens and Type Locality: Syntype, USNM 123142. MEXICO, Baja California, NW of

Cerros [Cedros] Island, 113 fm [207 m]; USBCF Albatross, station 2984. Not seen.

I Terebratalia occidentaIis obsoleta: Hertlein and Grant, 1944: 131-133, text-fig. 28, pl. 12, figs. 5, 9-11. --

Roth, 1972: 9.

i Terebratalia obsoleta: Dall, 1893: 382, pl. 2, figs. 4-12, pl. 3, figs. 1-15.--Beecher, 1893a: 382.--Beecher, 1893b: 392-394, pl. 2, figs. 4-12, pl. 3, figs. 1-15. --Dall, 1894 [1895]: 726, pl. 30, fig. 70. --Dall,

1920: 341-343.----Oldroyd, 1924b: 229-230. --Burch, 1942: 7. --Roth, 1972: 9.

i Terebratella obsoleta: Lowe, 1904: 20. Terebratula obsoleta: Dall, 1910: 96.

Dallinella obsoleta: Thompson, 1915: 75.--Thompson, 1927: 246.--Hatai, 1936: 90.

Terebratulia obsoIeta: Etherington, 1931" 49.

Terebratalia arnoldi Hertlein and Grant, 1944: 119, pl. 11, figs. 1-3, 10-15, 1960: 92-93, pl. 19, fig. 30.-

i RType oth, 1972: 3. Specimens and Type Locality: Holotype, CASIP 7313. USA, California, Ventura Co., Tapo

Canyon, center of southwest one-fourth of Section 23, T. 3 N., R. 18 W., S.B.B. and M. [Middle

I Pliocene]. Not seen. Waldheimia Kennedyi Dall, 1874: 299; 1903: 1540.

Type Specimens and Type Locality: Syntype, USNM, 370827. Mexico, Baja California, Cerros

I [Cedros] Island [Miocene].

TerebrataIia kennedyi: Dall, 1907: 33, pl. 17, figs. 4a-d.--Thompson, 1927: 246. --Hatai, 1938: 717; --

i Miogryphus 1940:willetti pl. 9, figs. Hertlein39-42. and--Hertlein Grant, 1944: and95-96, Grant,pl.1944: 1 125-126, pl. 13, figs. 1-4.--Roth, 1972: 8. l(figs. 4-9).--Mattox, 1956: 76.--Roth, 1972: 16.

Type Specimens and Type Locality: Holotype CASIP 7361. USA, California, Santa Barbara Co.,

I 6.4 km S of Lompoc, west side of San Miguelito Creek [Miocene, Temblor Formation]. Not seen.

Material Examined. 19 lots (see List of Additional Specimens Examined).

Description. Shell deep; ribs wide undulations of shell. Pedicle short. Color of shell whitish, ribs often carmine.

Shell size large; total lengths to 42 ram; widths to 46 mm; depths to 28 ram. !
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Figure 1.10. Terebratalia occidentalis (-Dall, 1871): off Santa Catalina Island, California (SBMNH 32400); 38.5 i

mm TL. A, dorsal view; B, ventral view; C, lateral view; D, end view, anterior margin; E, end view,

posterior margin. 1

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Type Specimens and Type Locality. Holotype, -- USNM 95850. California, off San Clemente "

Island, 45 fm [82 m], Albatross station 2981.

I

Distribution. Oligocene to Recent. Monterey, California (36°N) southward to Cabo San Lucas, l

Baja California, Mexico (23°N, SBMNH) and out to Cedros Island. Ranges in depth of collection from 50-

250 m.

I

Etymology. Specific name derived from the Latin word occidentalis, meaning "western or westerly." W

Common Name. Western lamp shell.

Biology. Nothing is known about the biology of this species. In southern California the species is I

abundant in collections that also include Laqueus californianus. The species typically occurs attached to

II

rocks or other brachiopod shells either singly or in small clusters (Mattox, 1956). I

Woodring (1950) and later Mattox (1956) questioned the distincmess of Terebratalia i

Remarks.

arnoldi Hertlein and Grant, 1944, a Pliocene fossil which is very similar to T. occidentalis in general shape

and ornamentation. It is considered here to represent a junior synonym. Other fossil taxa, such as T. smithi m

Arnold, 1917, need to be reevaluated and their taxonomic placement critically assessed. The smooth subspecies I

Terebratalia occidentalis obsoleta Dall, 1891 currently is considered to represent an ecomorph ofoccidentalis.

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I Terebratalia transversa (Sowerby, 1846)

Figure 1.11

Terebratula transversa Sowerby, 1846: 94; 1847: 361, pl. 72, figs. 114, 115. --Gould, 1860: 120.

TerebrateIla transversal: Gray, 1853: 89.--Reeve, 1860: pl. 5, fig. 22.--Dall, 1873: t85. _Whiteaves,

i Terebratalia 1878:transversa 468. --Davidson, : Dall, 1891: 1887:7172, 9 [in526,part], pl. 4,pl.figs. 16, figs. 6, 7;6-1894 9 only. [1895]: 727 [in part]; 1920: 339.--

Beecher, 1893a: 382.--Beecher, 1893b: 377.--Kelsey, 1907: 50.--Willett, 1918: 66.---Oldroyd,

I Thompson, 1924a: 204, pl. 1927: 29,245, figs.text-fig. 1-3; 1924b: 79. --Johnson 227, pl. 16,and figs.Snook, 1-3. --Hertlein 1927: 157,andfigs.Cri128, ckman, 131.1925: --Keep256.and--

Baily, 1935: 21-22, fig. 4.--Hatal, 1936: 89.--Nomura and Hatai, 1937: 180-181, pl. 25, fig. 20-

Burch, figs. 1, 2,195-7, 42: 6-7. pl. 21, --Hert figs.lein8, 9.and ---Smith Grant,and1944: Gordon, 135-11948: 38, te210. xt-figs--Mattox, 8, 9, pl. 81956: , figs.76, 10,figs. 15, 5A-J. 16, pl. --9,

Rickets and Calvin, 1968: 262-264, fig. 195. --MacGinitie and MacGinitie, 1968: 178. --Bernard,

1972:81, figs. 11-14. --Roth, 1972: 14. --Kozloff, 1973: 177. --Zimmer and Haderlie, 1980: 109-

1 110, pl. 39, figs. 7.1 a, b.

Terebratulina transversa: DaU, 1907:18.

I Magasella radiata Dall, 1877a: 159; 1877b: 49.--Davidson, 1887: 101, pl. 18, fig. 1.

Type Specimens and Type Locality: Holotype, USNM 110913. Alaska, Popoff Strait, Shumagin

Islands, low tide.

Terebratula transversa caurina Gould, 1850: 347; 1857: 468, pl. 44, fig. 582. 1862: 97.

Type Specimens and Type Locality: Holotype,--USNM 5964. Washington, Puget Sound. Not seen.

i Terebratula caurina: Gould, 1852: 468.--Carpenter, 1856 [1857]: 210, 298 [canrena, sic]

Terebratella caurina: Gould, 1862: 97.--Dall, 1870: 119, pl. 6, figs. 1-3. --Thompson, 1927: 246.

i Terebratella TerebrataIia transversa transversa var. caurina caurina : Dall, : Davidson, 1894[ 1895]: 1887: 733,80,pl.pl.31,16,figs. figs.12,10-12, 13; 1920: 14, 14a 340.only. --Oldroyd, 1924a:

204, pl. 29, figs. 4-7; 1924b: 228, pl. 16, figs. 4-7.--Fenton, 1932: 53-54, figs. 1-3. --Hatai, 1936:

89. 1944:--No138-143, mura and text-figs. Hatai,29,1937: 30, pl.181, 8, figs. pl. 25, 11-14, figs.pl.1-4. 9, figs. --Burch, 9, 10, 1942: 12, pl. 1.12,--Hertlein figs. 2, 8, pl. and15, Grant, figs.

7, 12, 13, 15, 16, pl. 21, figs. 12, 14.---Smith and Gordon, 1948: 210. --Mattox, 1956: 76, figs. 5A-D.

I Terebratalia caurina: Thompson, 1927: 246. --Hatai, 1936: 89. TerebrateUa coreanica: Carpenter, 1864: 636. [see Dall, 1920: 345]. (not Adams and Reeve, 1850).

Terebratalia transversa var. rubescens Dall, 1910: 96.

I Type Specimens and Type Locality: H01otype, --USNM, 253820. California, San Pedro. Not seen

TerebrataIia caurina rubescens: Dall, 1920: 341. --Burch, 1942: 7.

I Terebratalia rubescens: Hatai, 1936: 89.

Waldheimia grayi: Carpenter, 1864:636 [see Dall, 1920: footnote 350, 360]. (not Davidson, 1852).

I Terebratal1907: ia he425, mphill pl.i 58, Dalfigs. l 1904a, 2: 561 b. --Smith , pl. 40, fi1919: gs. 8,150-151. 10. --Arno--Hertlein ld, 1903:and92,Grant, pl. 17,1944: figs. 1123-125, -3. --Arnol pl.d,9,

figs. 3, 4, pl. 10, figs. 1, 2; 1960: 93-94, pl. 19, figs. 26, 27. --Roth, 1972: 7.

I Type ArroyoSpecimens Buero [Band urro],Typein Hope Locality: RanchHolotype, 112-1 mi--USNM inland from108495. sea [Pliocene]. California, west Not seen. of Santa Barbara,

Terebratalia hemphillii [sic]: Carson, 1925: 268.

Terebratalia obsoIeta: DuBois, 1916: 32, 1-8.

178, pl. figs. (not Sowerby).
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Material Examined. California: Santa Mafia Basin, Station BRC- 1, 73.5-78m, 2 specimens, 11.8, !

19.5 mm TL; USNM (w). Station BRA-13, 92-100m, 1 specimen, 5.5 mm TL; SBMNH (w).

Additional material examined: 22 lots (see List of Additional Specimens Examined). l

I

Description. Shell shape subovate to transversely produced, wider than long. Anterior commissure

sulcate, sometime sinuated; valve margin smooth to crenulated. Valves smooth, wrinkles with growth lines /

toward margin; valves surface of carina morph ornamented with 12 or more obtusely angular, radiating i

ribs, gradually enlarging and sometimes bifurcating; interspaces and elevations about equal; valves punctate. Ill

Beaks short, concavely flattened beneath each side; foramen very large; deltidium small, widely divided.

Color typically red, occasionally pale yellow to white or light gray. i

i

Shell size large; total lengths to 45 ram; widths to 55 mm; depths to 37 ram.

Type Specimens and Type Locality. BMNH, not searched. Washington, Puget Sound. Bernard •

(1972) indicated that a type locality was not designated by Sowerby. l

Distribution. Miocene to Recent. Eastern Pacific Ocean, Kodiak Islands, Alaska (58°N; Dall, 1920)

to Ensenada, Baja California, Mexico (32°N; Burch, 1942). Reported from the lower intertidal zone in the I

northern part of its distribution to 1800 m in the south. I

Etymology. Specific name derived fi'om the Latin transvers meaning "situated or lying across" in

obvious reference to the distinct ribs seen in some morphs of this species, i

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Common Name. Transverse lamp shell.

Biology. A voluminous literature documents the biology of Terebratalia transversa. The species in

has been studied extensively in Puget Sound and other areas in Washington. Ecological studies by Paine |

(1969, 1974) and Thayer (1975) have documented growth, size distribution, and the role and structure of T.

transversa populations in intertidal communities. This species is often extremely abundant intertidally in i

the northern part of its range, generally on underside of large boulders. I

Details of the ontogeny of shell secretion, such as development of the mantle and formation of the

juvenile shell and the development of the pedicle are elucidated in several papers by Stricker and Reed I_

(1985a-d). Details of metabolism and shell growth are discussed by Williams (1971), Hughes et aI. (1988), i

Rosenberg et al. (1988) and Tkachuck et al. (1989).

Reed (1987) and later Long and Stuckey (1994) reviewed reproduction and development in ill

Terebratalia transversa and several other west coast brachiopod species. In part they review the earlier II

embryology studies in the dissertation of Long (1964).

The ultrastructure and physiology of muscles, especially slow and rapid contraction mechanisms I

are treated in papers by Wilkens (1978a, b, 1987), Eshleman and Wilkens (1979), Eshelman et al. (1982), II

and Ackerly (1991 a-c).

The ultrastructure of a variety of organs are treated in papers by Gustus and Cloney (1972; setae), I

Reed and Cloney (1977; tentacles), and Cavey and Wilkens (1982; statocysts). Aspects of the functional s

morphology of the lophophore, including oxygen consumption, feeding and nutrient assimilation, are I

discussed by Strathman (1973), McCammon and Reynolds (1976), Reynolds and McCammon (1977) and i

LaBarbera (1986). The orientation of animals in relation to currents and the dynamics of ventilation are

I

included in LaBarbera (1978b, 1981). Thayer (1986) and later Rhodes and Thayer (1991) investigated the

effects of turbidity on suspension feeding in Terebratalia transversa and compared the efficiency of feeding i

in bivalves and articulate brachiopods. I1

Bradbury (1970) described Urceolaria kozloffi, a mobile ciliophoran from the tentacles of the

lophophore of two brachiopods from Puget Sound, Terebratalia transversa and Hemithyris psittacea. The i

protozoan is thought to feed at the expense of the host's food supply.

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Figure 1.11, Terebratalia transversa (Sowerby, 1846). A-E, depth unknown, off Point Conception, California

(SBMNH 46074); 34.4 mm TL. A, dorsal view; B, ventral view; C, lateral view; D, end view,

I anterior margin; E, end view, posterior margin. F, G, 0.5 m, Mission Bay, San Diego, California (SBMNH); 22.5 mm TL. F, dorsal view; G, end view, anterior margin.

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Remarks. The environmental variation of this highly plastic species has been discussed by Shimer

I (1905) and Du Bois (1916). Bernard (1972) stated that the subspecies Terebratalia transversa caurina (Gould) is not valid but is simply a southern heavily ribbed, gibbous form with a tendency to a reddish hue.

Terebratalia hemphilli, a widely distributed Pliocene fossil in Central California, is similar to the

I large smooth form of T. transversa, but generally is longer and broader (length 64mm, width 58ram and

depth 40 mm). It should be regarded as a synonym.

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Family Platidiidae Thomson, 1927 II

Diagnosis. Shell size small, plain colored; piano-convex, amphithyridid. Hinge plates absent; socket 1

ridges present. Pedicle short. Foramen large, occupies posterior area of both valves (= amphithyridid). Loop

rudimentary or absent. Spicules conspicuously present. Dental plates absent or low and heavy. Lophophore

shizolophous or plectolophous. Cardinal process absent. I

Ty It pe Genus. Platidia Costa, 1852.

Genus Platidia Costa, 1852 n

Diagnosis. Shell minute, thin; penetrated by numerous microscopic punctae. Hinge teeth weak,

crura long and curved; septum a triangular plate. I

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Type Species. Orthis anomioides Sacchi and Philippi, 1844; by monotypy.

Platidia hornii (Gabb, 1861) N

Figure 1.12

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Morrisia hornii Gabb, 1861:371. ---Carpenter, 1864: 632. ---Gabb, 1869: 36, pl. 12, fig. 63. ---Cooper, I

1871: 4. --Hertlein and Grant, 1944: 110-112, text-figs. 24, 25, pl. 7, fig. 30, pl. 19, figs. 14-18. --

Smith and Gordon, 1948: 210, 233. ]l

Platidia hornii: Thompson, 1927: 219. --Hatai, 1938: 715. --Bernard, 1972:79, fig. 4. --Zimmer and m

Haderlie, 1980: 110.

Platidia anomioides var. radiata Dall (in Orcutt), 1885: 552.--Dall, 1886: 204.--Thompson, 1927: 219. i

Type Specimens and Type Locality: Syntypes, --USNM 860930. California, San Diego, beach

drift. Not seen. I}

Platidia seminula var. radiata: Dall, 1889: 28. l

Platidia seminula radiata Dall, 1885: 551; 1920: 332-333. --Oldroyd, 1924b: 226, pl. 49, figs. 2, 7. --

Burch, 1942: 6. I

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Platidia radiata: Hatai, 1936: 87; 1938: 715.

Not Platidea aneminoides [sic]: Berry, 1907: 39. ---Gripp, 1909: 137. ---Oldroyd, 1914: 81. --Keep and it

Balty, 1935: 22. I

Not Platidia anomioides: Kelsey, 1907: 47. --Dall, 1907: 86.

Not Platidea anomioides: Berry, 1908: 39. ---Gripp, 1909: 137. --Oldroyd, 1914: 81. I

Material Examined. California: Santa Maria Basin, Station BRA-13, 92-100m, 44 specimens,

1.1-2.8 mm TL; USNM(w). Station BRA-25, 64.5-72m, ca. 30 spec., 1.3-2.7 mm TL; SBMNH(w). l

Additional material examined: 16 lots (see List of Additional Specimens Examined).

Description. Shell flattened; shape uniformly circular; valves thick; inner surface with numerous _

small punctae; outer surface without distinct sculpture, either smooth or with concentric growth lines. Pedicle II

short; foramen large, occupies posterior area of both valves. Color of shell light brown.

Shell size minute; maximum diameter observed to 4.6 mm. I

Type Specimens and Type Locality. Holotype, presumed to be in ANSP collection. California,

Santa Barbara [Miocene]. See remarks below.

i
I

I

I

i

l

i

I

i

I

i Figure 1.12. Platidia hornii (Gabb, 1861). A, B, 97 m, off Point La Jolla, California (SBMNH [SSB 30052]);

3.6mmTL. A, dorsal view; B, ventralview. C-E, 45 m, off Monterey,California (CASIZ 105953);

i 4.6mm TL. C, ventral view; D, ventral view; E, detail of punctae at edge of shell.

I Distribution. Upper Pliocene to Recent. Eastern North Pacific, British Columbia, (5 I°N; Bernard,

1972) to Todos Santos Bay, Baja California, Mexico (23°N; Dall, 1885). Depths range from low intertidal to

I 500 m. Biology. Attached to stones or dead shells in the low intertidal zone along semiprotected rocky

shores and in subtital waters. Additional information on the biology of this species is not not known. Tiny,

i young or immature specimens appear very similar to foraminiferans and often are confused as such. Remarks. A lot (ANSP 4495) labeled in Gabb's hand as the type ofMorrisia horni was obtained on

loan from Philadelphia. Unfortunately the specimen does not correspond to Platidia. It is presumed that the

i type Grant,is not extant. Cooper considered the species to belong in the genus Amphithyris (see Hertlein and 1944:112).

Reports of this species from Santa Cruz Island in the West Indies are presumed to instead refer to

Santa Cruz Island off the south central coast of California (see DaU, 1889). This apparent mistake has been

perpetuated in all subsequent papers which discuss distribution of this species.

I

!
I

Acknowledgments l

The following individuals provided valuable assistance in completing this publication: Andrea Moe cataloged I'

II

brachiopod material in the SBMNH collection; Elizabeth Kool and Terry Gosliner facilitated access to the

brachiopod collections at CASIZ; Mike Sweeney helped locate literature and provided assistance with

collections at the NMNH; Joann Rounds prepared illustrations for the introductory sections of this paper; []

Paul Scott and Marie Murphy helped to prepare plates of digitized images of brachiopod shells.

Ill

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List of Additional Specimens Examined n

Additional brachiopod specimens examined. Specimens preserved in alcohol are indicated by (w) I

following the catalog number, all other specimen lots are dry. Abbreviations: Co. = county; coll. = collector I

(s); spec. = specimen(s); TL = total length. I

n

G/ott/d/a a/b/da

California: 1 specimen, 12.0 mm TL; Ventura Co., Point Mugu, USNMC Base, 112mi N of Mugu t

Pier, 3/8 mi offshore, 40-50 ft [12-15 m]; coll. M. Conboy and G.E. MacGinitie, 13 August 1963; SBMNH Q

44477M.--2 spec., 16.4, 16.6 mm TL; Ventura Co., Point Mugu USNMC Base, 3/4 mi off Mugu Pier, 50-

60 ft [15-18 m]; coll. B. Spronce and party, 06-07 October 1963; SBMNH 46150 (w). --2 spec., 23.4 mm B

TL; Ventura Co., off Ventura, 20 fm [36 m]; coll. P. Brophy, June 1967; SBMNH. --2 spec., 5.0, 11.5 mm m

TL; Santa Barbara Co., Santa Barbara Channel, bearing 187°T, 2.5 mi from Platform Hilda; coll. R/V

Velero, station 12747, 06 March 1969; SBMNH 43528. --2 spec., 5.2 mm TL; Santa Barbara Co., Santa 1

Barbara Channel, beating 123°T, 2.7 mi from Steams Wharf; coll. RN Velero, station 12861, 03 April U

1969; SBMNH 43604.--1 spec., 15.7 mm TL; Santa Barbara Co., Santa Barbara Channel, bearing 119°T,

2.8 mi from Stearns Wharf; coll. RN Velero, station 12857, 04 April 1969; SBMNH 43566.----4 spec., 3.9- n,

6.6 mm TL; Santa Barbara Co., Santa Barbara Channel, bearing 117°T, 3 mi from Stearns Wharf; coll. RN I!1

Velero, station 12858, 03 April 1969; SBMNH 43558. Ventura Co., bearing 150°T, 4 mi off Port Hueneme

Lighthouse, 20 fm [36 m]; coll. RN Velero, station 12792, 20 March 1969; SBMNH 43786. --9 spec.,

22.1-31.7 mm TL; Los Angeles Co., off San Pedro; coll. and date unknown; SBMNH 28265 [ex F.S. U

VanDenBergh]. --10 spec., 14.5-19.9 mm TL; Los Angeles Co., Santa Catalina Island, off Avalon, 50 fm

[91 m]; coll. and date unknown; SBMNH [SSB 686].--1 spec., 33.3 mm TL; Orange Co., Newport Harbor, •

depth unknown; coll. W.S. Smith,- 1967; SBMNH 47644 (w).----4 spec., 12.7-22.2 mm TL; Orange Co., il

Newport Bay, depth unknown; coll. and date unknown; SBMNH. --2 spec.; San Diego Co., off La Jolla

Cove, 32°52'N, 117°17.4'W, 73 m; coll. F. Rokop and S. Luke, RN Oconostota, station 1, Van Veen grab, I

04 November 1969; BIM/SIO B22.

Neocrania cah'fornica I

California: I specimen + 9 valves, 11.4-14.4 mm TL; Coronado Bank, 32°32.8'N, 117°21.1%V, 60

fm [110 m]; coll. H.V. Hamilton, U.S. Bureau of Mines, station 58, date unknown; CASIZ 105949. --i 1

spec., 20.0 mm TL; Cordell Bank, bearing 347°T, 6.2 mi from distant rocks, 37°55'N, 123°24a_V,68 fm [124 |

m]; coll. USS Mulberry, station 19, 28 September 1950; CASIZ 105948.

Frieleia haUi I

California: 13 specimens, 12.8-19.0 mm TL; San Diego Co., off San Diego, 628-640 fm [1149- lit

1170 m]; coll. USBCF Albatross, station 4353 date unknown; SBMNH [SSB 3038]. --17 spec., 9.5-17.2 _

mm TL; San Diego Co., S of San Diego, "depth 1nu....., coll. C. Snell, December 1969; SBMNH 25850. g

24 spec., 6.1-20.0 mm TL; San Diego Co., San Diego Trough, 32°37'N, 117°32.6N_r, 1171-1217 m; coll. F.

Rokop, R. McConnaughey and S. Luke, RN Agassiz, station Rokop 70-1-12, 25 ft otter trawl, 29 October I

1970, attached to sponges; SBMNH [ex SIO] (w). --2 spec., 15.3-16.3 mm TL, Monterey Bay, 700 fm U

[1280 m]; coll. and date unknown; SBMNH [SSB 1183 or 1883?]. --1 spec., 18.0 mm TL; San Diego Co.,

San Diego Bay; coll. F.W. Kelsey, date unknown; CASIZ 029804 [ex Hemphill 19206; voucher, Hertlein I

and Grant, 1944, text-fig. 11, pl. 4, figs. 6, 7]. --4 spec., 13.0-15.5 mm TL; Monterey Bay; coll. USBCF U

Albatross; CASIZ 106304. --3 spec., 15.7-18.0 mm TL; San Diego Co., off San Diego, 32°40'N, 117°31'W,

822 fm [1503 m]; coll. USBCFAlbatross, station 2923, 19 January 1889; CASIZ 106306. ---4 spec., 14.4- I

42 I
!

I 18.0 mm TL; San Diego Co., San Clemente Island, beating 74°W, 5.8 mi from E point of Northwest Harbor,

542-599 fm [991-1096 m]; coll. USBCF Albatross, station D4402, 08 April 1904; CASIZ 106305. --1

I T. spec., Matsui, 16.7SIO71-29, mm TL; San25Diego ft otterCo., trawl, offSan18-19Diego, January32_24.7'N, 1971; BIM117°27.7'W, /SIO B42 658-680 (w). --200+fm [1243- spec.,1203 6.8-18.6 m]; coll. mm

TL; San Diego Trough, 32°35'N, 117°28'W, 1097-1180 m; coll. F. Rokop and S. Luke, R/V Oconostota,

I station 32°37.0'N, 4, 07117°32.6'W, November 1170-1216 1969; BIMm;/SIOcoll.B27F. Rokop (w). --200+ and S. Luke, spec., R5.0-18.0 mm TL; San Diego Trough, IVAgassiz, station Rokop 70-1-12, 25

ft otter trawl, 29 October 1970; BIM/SIO B53 (w). --140 spec., 7.2-18.2 mm TL; San Diego Trough,

i 32035.0'N, 117°33.6'W, 1183-1216 m; coll. F. Rokop and S. Luke, R/VAgassiz, station Rokop 71-II-28, 25 ft otter trawl, 21 January 1971; BIM/SIO B55 (d). --2 spec., 14.0, 16.4 mm TL; off La Jolla, 32°55.5'N,

117°37.4'W, 896 m; coll. J. F. Siebenaller, R/V Oconostota, station 10, 16 ft otter trawl, 06 September 1974;

i BIM/SIO B116 (w). --27 spec., 7.8-19.9 mmTL; San Diego Trough, 32°52.1'N, 117°42.1'W, 1057 m; coll. J.F. Siebenalter, R/V Agassiz, station A-24 Afterthot U, 25 ft otter trawl, 08 May 1975; BIM/SIO B117 (w).

Terebramlina crossei (also included here are specimens previously identified as T. kiiensis)

m

California: 1 specimen (broken), 27.4 mm TL; bearing WSW 3.5 mi off La Jolla Point, 53 fm [97

m]; coll. H. Barrington, F/V Christine, 10 April 1962; SBMNH [SSB 30049].--2 spec., 18.1, 24.0 mm TL;

I off Monterey, "deep water"; coll. H.R. Turner (from fishermen), 02 April 1943; SBMNH [SSB 16908]. -- 2 spec., 21.9, 27.2 mm TL; San Diego Co., off Tijuana River, 70-80 fm [128-146 m]; coll. R. McAllister,

October 1953; SBMNH [SSB 25547 and 24789].--3 spec., 5.6-31.4 mmTL; off La Jolla, 100fm [183 m];

i coll. San Diego Marine Biological Association, 1901; SBMNI-t [SSB 3265]. --1 spec., 27.4 mm TL; San Diego Co., beating SW, 9.7 mi off Point Loma, 34°34.2'N, 117° 24'W, 120 fm [220 m]; coll. W. Dahlstrom,

prawn trap, 29 March 1969; SBMNH [SSB 36310]. --35 spec., 6.6-25.0 mm TL; Cortez Bank, 125 fm [229

I m]; Bank,coll. L. Zermatten, hook and line, April 1965; SBMNH [SSB 33321]. --1 spec., 4.0 mm TL; Cortez 100 fm; coil. L. Zermatten, December 1962; SBMNH [SSB 30019]. --1 spec., 30.6 mm TL; bearing

S, 9 mi off Santa Monica, 48 fm [88 m]; coll. P. Brophy, ballon trawl, August 1971; SBMNH. --2 spec.,

I 11.9, 15.2 mm TL; Santa Barbara Channel, off Gaviota, 73 fm [133 m]; coll. P. Brophy, June 1966; SBMNH. --8 spec., 10.2-12.8 mm TL; Santa Barbara Channel, off Gaviota, 72 fm [132 m]; coll. P. Brophy, June

1968; SBMNH.--2 spec., 11.0, 18.0 mm TL; Santa Catalina Island, off Isthmus, 50 fm [91 m]; coll. P.

I Brophy, (between 2Point m beamMugutrawl, and June Point1972; Dume), SBMNH 50 fm (w). [91 m]; --1 coll. spec., H. 31.3 Barrington, mm TL;F/Venture V Christine Co.,, Sycamore 24 March Bank 1962;

SBMNH [SSB 30069]. --1 spec., 27.0 mm TL; San Francisco Co., Farallon Islands, W of Middle Farallon

I Island, USS Mulberry bearing, 101°T, 26 August 9.2 mi 1944; fromCASIZ Farallon 105951 Light, [GE37°43.8'N, 32934]. --1126°04.6'W, spec., 30.070-126 mm TL;fmSan[128-230 Diego m]; Co., coll. San

Diego region, 32°32'N, 117°19.0'W, 104-250 m; coll. San Diego Marine Biological Association, 27 July

I Point, 1901; CASIZ 64-80 fm106307. [117-146--3m];spec., coll. 18.1-28.0 G.D. Hanna, mmRTL; /V NSanta .B. Scofield Barbara, 6 Co., June bearing 1953; CASIZ 233"I',106308.----4 3 mi off Coalspec., Oil

17.3-28.5 mm TL; San Diego Co., San Diego region, 32052'N, 117°17'W, 100-195 m; coll. San Diego

I Marine RedondoBiological Beach, deep Association, water; coll. 23and Julydate 1901; unknown; CASIZ CASIZ 106309. 106336. --1 spec., --131.4spec., mm35.0TL;mmLos TL; Angeles Los ACo., ngelesoff

Co., off Redondo Beach, deep water; coll. H. Hill, pre 1938; CASIZ 029801 [voucher, Hertlein and Grant,

i A. 1944, Sorensen, pl. 17(figs. date 3,unknown, 6 10)]. --2attached spec., 34.9, 37.2 mm TL; Monterey Co., Lopez Point, 70 fm [128 m]; coll. to solitary coral, Paracyathus stearnsii; CASIZ 105950 [GE 37188].

--2 spec., 28.8, 35.5 mm TL; Monterey Co., off Point Sur, 36_20.45'N, 122_06.15"9¢, 208 fm [380 m]; coll.

i R. Diego, Bolin and Budd, 12 November 1935; CASIZ 016592. --1 spec. 29.5 mm TL; San Diego Co., off San 50 fm [91 m]; coll. and date unknown; CASIZ 029806 [voucher for T. kiiensis, Hertlein and Grant,

1944, text-fig. 18].--3 spec., 32.0-34.1 mm TL; Monterey Bay, 50-60 fm [91-110 m]; coll. unknown, 28

i June 1932; CASIZ 106335. --3 spec., 21.8-34.0 mm TL; Los Angeles Co., off San Pedro, deepwater; coll.

I 43
!

T.S. Oldroyd, date unknown; CASIZ 106334. --3 spec., 20.0-21.0 mm TL; San Diego Co., La Jolla, 100 fm I

[183 m]; coll. University of California, pre 1912; CASIZ 105952. --2 spec., 28.2, 29.1 mm TL; off San

Diego, 137 m; coll. and date unknown; BIM/SIO B65 [ex Baker-Kelsey collection]. --3 spec., 21.5-29.6 I

mm TL; Forty-mile Bank, 32"N, 117°56.2N¢, 183 m; coll. F.P. Shepard, R/V E.W. Scripps, station Shepard U

69, dredge, 10 June 1938; BIM/SIO B118. --5 spec., 25.0-31.0 mm TL; Forty-mile Bank, 32°40'N, 118°12'W,

60 fm [110 m]; coll. F.P. Shepard, RN E.W. Scripps, dredge, 22 March 1938; BIM/SIO Bl]9. --2 spec., •

33.0, 34.2 mm TL; San Diego Co., Nine-mile Bank, 9 mi W by SW off Point Loma, 32°40'N, 117°28'W, II

depth unknown; coll. Capt. J. LaGrange, F/V Cloud Nine, spring 1990; SBMNH.

Terebratulina unguicula !

California: 7 specimens, 2.8-17.7 mm TL; Monterey Co., off Point Pifios, 46-56 fm [84-102 m]; II

coll. USBCF Albatross, station 4551, date unknown; SBMNH [SSB 3034]. --10 spec., 5.4-13.9 mm TL; II

Santa Barbara Co, Anacapa passage, 26-27 fm [47-49 m]; coll. C.L. Hubbs and party, R/V Orca, station

H51-252, 02 July 1951, on coral colony; SBMNH [SSB 23577]. --17 spec., 3.9-14.7 mm TL; Santa Rosa I

Island, offE point, 80 fm [146 m]; coll. F. Fricia, 15 May 1953; SBMNH [SSB 18889].--10 spec., 5.2-11.2 n

mm TL; San Diego Co., off Oceanside; coll. H.G. Eaton, ca. 1906, from kelp holdfasts; SBMNH [SSB

9096]. --9 spec., 5.9-11.7 mm TL; San Diego Co., off La Jolla, La Jolla Canyon; coll. Scripps Institution of I

Oceanography, 22 March 1938, on solitary corals; SBMNH [SSB 13507]. ---6 spec., 3.8-8.9 mmTL; Ventura il

Co., Sycamore Bank (between Point Mugu and Point Dume), 34_00'N, 119°02'W, 50 fm [91 m]; coll. H.

Barrington, F/V Christine, 24 March 1962; SBMNH [SSB ??]. ----6spec., 3.0-14.9 mm TL; San Diego Co., I

bearing WSW, 3.5 mi offLa Jolla Point, 53 fm [97 m]; coll. H. Barrington, F/V Christine, 10 April 1962; I

SBMNH [SSB 30050]. ----41 spec., 3.3-15.8 mm TL; Monterey Bay, 15-30 fm [27-55 m]; coll. and date

unknown, station 3-18, attached to solitary coral; SBMNH 28272. --1 spec., 9.8 mmTL; 195°'1"10 mi from m

Marina det Rey Harbor, 59-62 fm [t08-113 m]; coll. P. Brophy, shrimp trawl, 9 September 1974, on coral !

head; SBMNH (w). --1 spec., 12.5 mm TL; Humboldt Co., off Trinidad, 180 fm [329 m]; coll. unknown,

September 1975; SBMNH. --1 spec., 10.0 mm TL; San Diego Co., off San Diego; coll. and date unknown; /

CASIZ 29820 [ex Hemphill 6073; voucher, Hertlein and Grant, 1944, pl. 5(figs. 9, 10)].--14 spec., 11.5- _l

14.0 mm TL; Monterey Bay, Monterey Co., off Del Monte, 35-40 fm [64-73 m]; coll. J.Q. Butch and A.G.

I

Smith, 20 August 1937, attached to solitary corals; CASIZ 106319.---6 spec., 13.5-23.0 mmTL; Humboldt in

Co., off Eureka, 67 fm [122 m]; coll. Capt. J. Riley, M/V Inez, March 1969; CASIZ 106333.----6 spec., 4.2- i

13.4 mm TL; Ven_ra Co., off San Nicolas Island; coll. and date unknown; CASIZ 27603. ---8 spec., 9.0- J

18.5 mm TL; Humboldt Co., Eel River Canyon, 40°28'N, 200 fm [366 m]; coll. Capt. K. Einhem, F/V

Stephanie, 19 September 1975, attached to calcitic rock; CASIZ 106321. --1 spec. + 1 valve, 6.0-6.8 mm B

TL; Monterey Bay, Monterey Co., off Mussel Point, 10 fm[ 18 m]; coll. A.G. Smith and M. Gordon, station II

4, 27 September 1932; CASIZ 106320. --18 spec., 8.5-13.2 mm TL; Monterey Bay, Monterey Co., off

Monterey, ca. 2-112 mi offshore, 35-40 fm [64-73 m]; coll. A.G. Smith and J. Burch, 20 August 1937; I

CASIZ 28916. --22 spec., 3.3-13.0 mm TL; Monterey Bay, Monterey Co., 1t2 mi off Hopkins Marine g

Station, between Pacific Grove bell bouy and Del Monte wharf, 20-30 fm [37-55 m]; coll. G.D. Hanna, A.G.

Smith and J.L. Nicholson, July 1930; CASIZ 106318.--102 spec., 2.0-10.5 mm TL; Monterey Bay; coll. I

and date unknown, attached to solitary corals, Paracyathus stearnsii; CASIZ 106322. --2 spec., 24.0, 25.0 I

mm TL; Monterey Bay; coll. J. Strohbeen, March 1956; CASIZ 106338. --7 spec. + 3 valves; Monterey

Bay, Monterey Co., Pacific Grove, 20 ft [6m]; coll. Harrington; CASIZ 106316 [ex SU 53167].--2 spec., I

5.4-7.5 mm TL; off Point Loma, 64 m; coll. USBCF Albatross, 12 August 1916, attached to solitary coral,

Paracyathus stearnsii; BIM/SIO B102. --1 spec., 15.6 mm TL; offLa Jolla, in Scripps Canyon, 1/2 mi off

SIO pier, 32°52.12'N, 117°15.58'W, 600 ft [183 m]; coll. R. McConnaughey, trap, 03 February 1984; BIM/ 1

SIO B156 (w). IB

I
!

I Washington: 4 specimens, 8.3-25.0 TL; Clallam Co., 50 mi W of Umatilla Rocks, 72 fm [132 m];

coll. Capt. E. Anderson, F/V Marie II, March 1960; CASIZ 106315. --1 spec., 21.0 mm TL; Puget Sound,

I 25 fm [46 m]; coll. and date unknown; CASIZ 106314.

Laqueus cah'fornianus

i California: 7 specimens, 23.2-40.0 mm TL; Los Angeles County, off San Pedro, 62-79 m; coll.

W.E. Ritter and W.J.Raymond, San Diego Marine Biological Association, station 23, 22 June 1901; SBMNH

i 28267 V. Stevens, and 46052. 01 August --1 spe1930; c., 41SBMNH .5 mmTL;28273. Santa Barbara Co., Santa Barbara Channel, 70 fin [I28 m]; coll. --1 spec., 32.2 mm TL; Santa Barbara Co., Santa Barbara

Channel, off Gaviota, 130fm [238 m]; coll. Capt.R. Hazard, FtVKildee, March 1974; SBMNH. --26 spec.,

i 14.3-50.5 mm TL; Los Angeles Co., Santa Catalina Island, off White's Landing, 30 fm [55 m]; coll. G.Willett, 1927; SBMNH [SSB 6331]. --15 spec., 11.1-45.5 mm TL; Los Angeles Co., off Santa Catalina Island; colt.

SIS Anton Dohrn, 31 March 1915; SBMNH 46050. --9 spec., 12.7-45.9 mm TL; Los Angeles Co., Santa

I Catafina Island, off Isthmus, 30 fm [55 m]; coll. P. Brophy, beam trawl, May 1969; SBMNH. --5 spec.,

41.1-49.0 mm TL; San Luis Opispo Co., SSE of San Simeon Point, 35029.4'N, 121°08.8'W, 116-118 fm

[212-216 m]; coll. W. Dahtstrom, 19 October 1960; SBMNH [SSB 30077]. --1 spec., 40.5 mm TL; San

I Luis Opispo Co., off San Simeon, 90 fm [165 m]; coll. P. Brophy, shrimp trawl, November 1969; SBMNH. --21 spec., 10.0-49.3 mm TL; San Luis Obispo Co., off Morro Bay, 85 fm [155 m]; coll. J. Squire, R/V

Mama II, 02 February 1953; SBMNH. --24 spec., 4.9-51.6 mm TL; Monterey Co., Monterey Bay, 36°40.3'N,

122°07.1'W, 75 fm [320 m]; coll. R.L. Bolin, 22 December 1937; SBMNH [SSB 8948]. --16 spec., 3.8-

I mm TL; Angeles Co., Island, Avalon, [91 m]; coll. J.H. Paine, 1903;

47.4 Los Santa Catalina off 50 fm

SBMNH [SSB 93]. ---ca. 10 spec., 3.2-6.1 mm TL; Los Angeles Co., Santa Catalina Island, off Isthmus, 42

fm [77 m]; coll. P. Brophy, beam trawl, September 1974; SBMNH. --3 spec., 8.0-38.2 mm TL; San Francisco

i Co., off Moritero Point, 180 fm [329 coll. P. 1968; SBMNH --10 13.8-45.0

m]; Gregory, July (w). spec.,

mm TL; Monterey Co., Monterey Bay, 36°40'30''N, 122°07'15"W, 75 fm [137 m]; coll. R. Bolin, 22 December

1937; SBMNH (w). --28 spec., 12.0-49.5 mm TL; Los Angeles Co., Santa Catalina Island, 2.8 mi E of

I Long Point, 33°26'00"N, 118°24'40"W, 68-88 fm [124-161 m]; coll. R/V Velero III, station 1307-41, 03 April 1941; SBMNI-I (w). --108 spec., 4.5-31.8 mm TL; coll. R/V Velero III, station 2953-54; SBMNH [ex

AHF] (w). --1 spec., 32.3 mm TL; San Luis Obispo Co., Morro Bay, mud flats, depth unknown; coll.

I unknown, 29 June 1957; SBMNH. --1 spec., 46.0 mmTL; San Luis Obispo Co., Morro Bay, depth unknown; coll. M. Smith, 1969; SBMNH.--3 spec., 13.3-37.8 mm TL; LOsAngeles Co., San Pedro, off Point Vicente,

40 fm [73 m]; coll. unknown, 1924; SBMNH. --13 spec., 5.0-29.6 mm TL; Monterey Co., Monterey Bay,

I 2-3 mi SW of Soberanes Point, 85-110 fm [155-201 m]; coll. R/V Searcher, station C-71-59, 22 July 1971; CASIZ 106313. --9 spec., 6.4-29.6 mm TL; Mendocino Co., bearing W by S 1t4 W, 3 mi from Fort Bragg

bouy, 60-65 fm [110-119 m]; coll. G.D. Hanna, R/V. N.B. Scofield, station NBS 277, 02 August 1940;

CASIZ 106312. --3 spec., 22.4-44.3 mm TL; LOs Angeles Co., W of Santa Catalina Island, 33°23.0'N, 118°30.9'W, 165-230 m; coll. C.A. Kofoid and W.J. Raymond, San Diego Marine Biological Association,

station 34, 28 June 1901; CASIZ 106328.

! Terebratalia occidentalis

I unknown;CalSBMNH ifornia: 46068. 3 specimen --2 sspec., , 37.4-39.9 38.1, 39.9mmmmTL;TL;southern LOs Angeles California, Co., off"deepSantawater"; Catalinacoll.Island; and date coll.

unknown, ca. 1933; SBMNH 32400. --3 spec., 12.3-39.5 mm TL; LOs Angeles Co., San Pedro Bay; coll.

I and end,date 33°26'N, unknown; 118°25'W, SBMNH33-43 32380. fm --1[60-79 spec.,m];33.2coll.mmunknown, TL; Los Angeles 09 February Co., Santa 1949;Catali SBMNH na Island, 46064.--1 off SE

spec., 42.8 mm TL; Los Angeles Co., Santa Catalina Island, off Avalon, 50 fm [91 m]; coll. J.H. Paine,

!
!

II

1903; SBMNH [SSB 394]. --1 spec., 32.4 mm TL; Los Angeles Co., off San Pedro; coll. W.H. Golisch _l

(from fishermen), date unknown; SBMNH [SSB 458].--19 spec., 13.3-37.6 mm TL; Los Angeles Co., off

Santa Catalina Island, 40 fm [73 in]; coLl.USC, station D103, 19 July 1924; SBMNH. --1 spec., 32.7 mm I/

TL; LOs Angeles Co., Santa Catalina Island, off Isthmus Cove; coll. USC, date unknown; SBMNH. --18 II

spec., 14.4-42.3 mm TL; LOs Angeles Co., Santa Catalina Island, offWhite's Landing, 30 fm [55 m]; coll.

G. Willett, 1927; SBMNH [SSB 63332, 6333, 6334, 6335]. --7 spec., 9.9-28.6 mm TL; LOs Angeles Co., l1

off San Pedro, 62-79 m; coll. W.E. Ritter and W.J. Raymond, San Diego Marine Biological Association, 22 |

June 1901; SBMNH 46079. --3 spec., 11.0-20.4 mm TL; LOs Angeles Co., off Santa Catalina Island; coil.

SIS Anton Dohrn, dredge, 31 March 1915; SBMNH 46078. --2 spec., 31.6, 41.0 mm TL; Los Angeles Co., II

Santa Catalina Island, off Isthmus, 30 fm [55 m]; coll. P. Brophy, May 1969; SBMNH.--3 spec., 18.5-23.9 II

mm TL; Los Angeles Co., off San Pedro; coll. T.S. Oldmyd, date unknown; SBMNH 46060. ----6spec., 8.8-

32.0 mm TL; San Diego Co., Nine-mile Bank, 9 mi W by SW of Point Loma, depth unknown; coll. Capt. J. 11

LaGrange, F/V Cloud Nine, spring 1990; SBMNH. --1 spec., 28.5 mm TL; Los Angeles Co., off Santa |

Catalina Island, 30 fm [55 m]; coll. G. Willett, pre 1934; CASIZ 29805 [voucher for T. o. obsoleta, Hertlein

and Grant, text-fig. 28]. --4 spec., 23.0-41.4 mm TL; Los Angeles Co., off San Pedro; coll. Mrs. M. Baldridge, il

date unknown; CASIZ 106311 [ex A.G. Smith 6352]. --14 spec., 23.0-41.4 mm TL; Los Angeles Co., off !1

Santa Catalina Island, 30 fm [55 m]; coll. G. Willett, date unknown; CASIZ 106310. --1 spec., 34.0 mm

TL; LOs Angeles Co., Santa Monica Bay, bearing W, 6.25 mi off Playa del Rey, 300-378 ft [91-115 m]; coll. am

W. Donnelly, F/V Prowler, 14 March 1962; CASIZ 106337. I

Mexico: 6 specimens, 15.8-29.5 mm TL; Baja California (outer coast), Bahia Asuncion, 27°02.1'N,

114°21.1'W, 57-62 fm [104-113 m]; coll. C. Hubbs and R. Wisner, R/V Horizon, station MV-65-I-29 [SIO i

65-212]; 45 ft otter trawl; 21 June 1965; SBMNH [ex BIM/SIO B21] (w). II

Terebratalia transversa I

California: 1 specimen, 29.2 mm TL; LOs Angeles Co., off E1Segundo, 31 fm [57 m]; coll. P.

Brophy, beam trawl, February 1974; SBMNH.--11 spec., 11.8-28.1 mm TL; southern California, deep il

water; coil. and date unknown; SBMNH 28263. --3 spec., 36.1-42.5 mm TL; Orange Co., Newport, 600 fi I!

[183 m]; coU. fishermen, date unknown; SBMNH. --8 spec., 24.9-30.6 mm TL; Los Angeles Co., San

Pedro; coll. W.H. Eshmaur, date unknown; SBMNH [SSB 154]. --15 spec., 6.7-27.0 mm TL; Los Angeles 1

Co., San Pedro, Government Breakwater, low intertidal; coll. E.P. Chace, November 1914; SBMNH [SSB I!

3409]. --3 spec., 12.4-24.7 mm TL; San Clemente Island, offSW side, 70 ft [21 m]; coll. R. Fay, September

1972; SBMNH.--10 spec., 12.8-34.9 mm TL; ca. 10-314 mi W of Point Dume, 34000.20'N, 119°01.20'W, all

47-48 fm [86-88 m]; coll. R/V Velero III, station 1276-41, 23 March 1941; SBMNH [ex AHF] (w). --12 II

spec., 15.5-25.7 mm TL; Santa Barbara Co., Santa Barbara; coll. and date unknown; SBMNH 28261 [ex

II

F.S. VanDenBergh]. --1 spec., 40.2 mm TL; Santa Barbara Co., Santa Cruz Island, off Gull Island, 5 fm [9 I

m]; coll. C. Snell, November 1967; SBMNH. --1 spec., 19.6 mm TL; Los Angeles Co., Santa Catalina l[m

Island, off Avalon, 30 fm [55 m]; coll. and date unknown; SBMNH [SSB 293]. --7 spec., 28.0-42.1 mm

TL; Santa Barbara Co., off Point Conception; coll. B. McMillen, December 1965; SBMNH 46074. --1

spec., 13.2 mm TL; Santa Barbara Co., off Point Conception; coll. B. McMillen, 09 June 1965; SBMNH 1

46075. --1 spec., 24.6 mm TL; Santa Barbara Co., off Rincon, 45 fm [82 m]; coll. P. Brophy, July 1965;

SBMNH. --2 spec., 21.7, 27.4 turn TL; 9 mi S of Santa Monica_ 48 fm [88 m]; coll. P. Brophy, balloon

trawl, August 1971; SBMNH. --1 spec., 22.5 mm TL; Monterey Co., Monterey Bay, 15 fm [27 m]; coil. I!

Stanford University marine lab, 1908; SBMNH [SSB 2999]. --3 spec., 20.5-35.0 mm TL; Los Angeles Co., 19

off San Pedro; coll. and date unknown; CASIZ 29798 [voucher for T. t. caurina, Hertlein and Grant, 1944,

pl. 15 (figs. 7, 12, 13, 15)].--1 spec., 25.0 mmTL; Los Angeles Co., San Pedro Bay; coll. and date unknown; I

CASIZ 29799 [ex H. Hemphill; voucher for T. t. caurina, Hertlein and Grant, 1944, pl. 15 (fig. 16)]. --10 m

spec., 12.0-34.4 mm TL; Los Angeles Co., San Pedro, Government Breakwater; coll. E.P. Chace, 19 June

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i 1916; CASIZ 106329 + 106330 [ex A.G. Smith 1318 + 1462]. --2 spec.; Monterey Bay, Monterey Co., off

Del Monte, 12 fm [22 m]; coll. A.G. Smith, 05 June 1992 [ex A.G. Smith 2439]. --3 spec., 17.0-36.7 mm

i TL; Cortez Bank, 32°30.2'N, 119°13.7aN, 282 ft [86 m]; coll. F.H. Berry, 16 May 1963; BIM/SIO B47 (d). --1 spec., 39.8 mm TL; Quast Rock, offLa Jolla, depth unknown; colt. W.A. Newman, spring 1968; BIM/

SIO B96.

I CASIZ 29816 Washington [ex H.: Hemphill 2 specimens, 13951; 36.5,voucher, 47.0 mmHertlein TL; Puget and Grant, Sound;1944, coll.pl.T.S. 8(figs. Oldr10, oyd, 15,d16), ate unknown; pt. 9 (figs.

1, 2)].

I Platidia hornii

i m]; coll.Cal L. ifornia Zermatte: 25n, specime hook andns,line, 2.1-4.4 05 December mm TL; S1966; anta BaSBMNH rbara Co., [SSBS o33309]. f San Miguel --10+ Island, spec., 2.3-2.7 65 fm [119 mm

TL; Los Angeles Co., San Pedro Bay; coll. E.P. Chace, date unknown, on empty Mytilus californiensis

i scoll. hell;RS/VBMNH E.W. Scripps 19853. ,--123 January spec., 3.51948; mm TL; SBMNH San Diego [SSB Co., 15351].--20+ off La Jolla,spec., Scripps 2.2-4.8CanyommTL; n, 30 fmSan[55Diego m];

Co., bearing WSW, 3.5 mi off La Jolla Point, 53 fm [97 m]; coll. H. Barrington, F/V Christine, 10 April

I Mugu 1962; SandBMNHPoint[SSB Dume), 30052]. 34°00'N, --20+119°02'W, spec., 1.9-4.4 50 mm TL; Ventura Co., Sycamore Bank (between Point fm [91 m]; coll. H. Barrington, F/V Christine; 24 March

1962; SBMNH [SSB 30068]. --3 spec., 3.5-4.2 mm TL; Ventura Co., midway between Ventura and Anacapa

I Is4.4land, mm125TL;fmSan[229Luism];Obispo coll. H.Co., Barringto off Avila, n, FN35Christine °00.7'N,, 121 -- °1962; 21.5'W, SBMNH 220-240[SSB 30021]. --5 spec., 3.8- fm [402-439 m]; coll. G.

Bickford, station 63-A-2, 23 March 1963; SBMNH [SSB 30086]. --3 spec., 2.5-4.1 mm TL; Los Angeles

i Co., offRedondo Beach, depth unknown; coll. G. Willett, date unknown; CASIZ 106322. ---4 spec., 3.0-4.6 mm TL; Monterey Co., off Carmel, 25 fm [46 m]; colt. M. Gordon, date unknown; CASIZ 105953. --5

spec., 3.2-3.6 mm TL; Monterey Co., Carmel Bay, 25 fm [46 m]; coll. H. Heath, R/V W.L. Scofield, pre

i March 1930; CASIZ 106327. --1 spec., 4.0 mm TL; San Francisco Co., NW of Middle Farallon Island, 70 fm [128 m]; coll. and date unknown; CASIZ 33150. --1 spec., 3.9 mm TL; NW of Cordell Bank, 38003.0'N,

123°33.0'W, 200-400 fm [366-732 m]; coll. USS Mulberry, station 56, 29 March 1950; CASIZ 106326. --

32 spec., 1.7-3.2 mm TL; Los Angeles Co., off San Diego, 32°40'N, 117018'W, 110-240 m; coll. W.J.

I Raymond, Diego Biological Association, 83, August 1901;

San Marine station 05 CASIZ 106325. _2

spec., 3.0, 4.0 mm TL; Los Angeles Co., off San Pedro, 200 fm [366 m]; coll. and date unknown; CASIZ

106324. --29 spec., 1.8-3.0 mmTL; San Diego Co., offSan Diego, depth unknown; coll. T.S. Oldroyd, date

I unknown; CASIZ 106323. --2 2.6, 3.2 mm TL; off San Diego, 50 fm [91 m] colt. and date unknown;

spec.,

BIM/SIO B82 [ex Baker-Kelsey collection].

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I 2. PHYLUMSUPUNCULA

| uy

Paula S. Winchell _

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I Introduction

The Sipuncula, commonly referred to as the "peanut worms," include roughly 250 species in 17

I mol genera. They arebenthic marine animals, burrowing in sediments, hiding under stones and in some cases in lusc shells or granular tubes.

Sipuncula are coelomate, bilateral, and similar to annelids although they are not segmented. The

body is spherical or cylindrical, and can be separated into two distinct areas: the trunk and retractable

a

introvert region. The intestines are coiled and U-shaped, meaning that the anus is not found on the posterior

end of the animal but anteriorly, in the region where the trunk and introvert meet. aa

I During the Phase I reconnaissance only three species were found, each from a different genus, a

fourth genus and species was found in the Phase II monitoring program. It is likely that additional species

i occur in the general area as well. In the last two decades the phylum as a whole has undergone major taxonomic revisions, and it is important to have the recent literature on hand in order to identify the animals

correctly.

I Taxonomic History

i In 1767 Linnaeus included illustrations of Sipuncula in the twelfth edition of his Systema Naturae

under the heading Vermes. In 1828, DeBlainville introduced the name Sipunculida, and grouped the animals

i with parasitic helminths. Quatrefages grouped the sipunculans, echiurans and priapulans together under the heading Gephyrea in 1847. Gephyra is a greek root meaning "bridge". Quatrefages thought the animals in

these three groups were the intermediary between the annelids and echinoderms. Although he based his

t ideas on superficial characteristics, the notion persisted well into the twentieth century despite attempts by several authors to raise the groups to separate phyla. The first suggestion of elevating the Sipuncula to

phylum was made by Sedgwick (1898). The Sipunculida was finally elevated to phylum rank by Hyman

(1959). Stephen (1964) proposed the currently accepted spelling of the phylum. Stephen and Edmonds

I (1972) genera. Gibbs (1985) further classified the phylum into

established four families and 17 Curler and

two classes, four orders, six families and 17 genera. A key to the families and genera is presented by Gibbs

and Cutler (1987), as well as discussions of subgenera. The keys to subgenera and species can be found in a

series of by Cutler (1979, i986); Cutler and Cutler (1982, 1983, 1985a, 1985b, 1986, 1987, papers 1988,

1989, 1990); Cutler et al. (1983); Curler and Jurczak (1975); Cutler and Murina (1977) and Gibbs et al.

(1983). An overview of sipunculan systematics and synoptic keys to all known genera and species is presented

I in the recent treatise by Curler (1994) 2.

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1948 Head of the Bay Road, Plymouth, Massachusetts 02360.

2Cutler's (1994) book was received too late to be fully incorporated into this chapter. This work provides a thorough

I review of the systematics, biology, and phylogeny of the Sipuncula.

I 49
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Chamberlin (1920) produced a list of the sipunculans of Laguna Beach. There were six species, i

four of which were new to science. Ricketts and Calvin (1952) included several species of Sipuncula in theft

discussions of f'mdings on the Californian shore, but it is a listing by habitat and is therefore difficult to use I

as a taxonomic guide. Fisher (1952) was the first to present a guide to the Sipuncula of California complete II

with keys and species descriptions. In this study he reports the species identifiable from the deep-sea

collections of the Albatross in 1904, adding that the majority of this collection was not usable by the time he 1

attempted identifications. Although both are good references for the intertidal region, they have little or no I

information on the deep-sea animals. Rice (1975) is a guide to the sipunculans of the central Californian

coast, but is virtually lacking in information on the deeper water species as well. Hartman and Barnard 1

(1958 and 1960) characterized the fauna of the deep-sea basins off Southern California, but the majority of 1

the sipunculans were not named to species, and were not described sufficiently to allow for a reasonable

identification. Frank (1983)gives a listing of the Sipuncuta from Canadian waters, and some of his species 11

have distributions that extend as far south as Baja California and can be a helpful reference. |

Systematies .i

Class Phascolosomida Cutler and Gibbs, 1985 _.

Order Aspidosiphoniformes Cutler and Gibbs, 1985 II

Family Aspidosiphonidae Baird, 1868

Aspidosiphon Diesing, 1851 i

Cloeosiphon Grube, 1868

Lithacrosiphon Shipley, 1902 I

Order Phascolosomaformes Cutler and Gibbs, 1985

Family Phascolosomatidae Stephen and Edmonds, 1972

Antillesoma Stephen and Edmonds, 1972 I

Apionsoma Sluiter, 1902

Phascolosoma Leuckart, 1828 I

lIB

Class Sipunculida Cutler and Gibbs, 1985

Order Golfingiaformes Cutler and Gibbs, 1985 Ill

Family Golf'mgiidae Stephen and Edmonds, 1972 !1

Golfingia Lankester, 1885 1

Nephasoma Pergament, 1946 !

Thysanocardia Fisher, 1950

Family Phascolionidae Cutler and Gibbs, 1985 II

Onchnesoma Koren and Danielssen, 1875

Phascolion Threl, 1875 I

Family Themistidae Cutler and Gibbs, 1985 I

Themiste Gray, 1828

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i Order Sipunculiformes Cutler and Gibbs, 1985

Family Sipunculidae Baird, 1868

I Phascolopsis Fisher, 1950

Siphonomecus Fisher, 1947

I Siphonosoma Sipunculus Linnaeus, Spengel,1766 1912

I Xenosiphon Fisher, 1947

Morphology

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The Sipunculan body can be broken down into two distinct regions: the trunk and the retractable

introvert (Fig. 2.1A). The introvert is extended by an increase in coelomic pressure when the drcul_

i muscles of the wall contracted At the anterior end of the introvert is the mouth and

body are (Fig. 2.1A).

feeding tentacles (Fig. 2.1A). The tentacles may be highly elaborate, as in Thysanocardia and Themiste, or

greatly reduced as in Nephasoma and Onchnesoma. The tentacles are part of the compensation system.

I This fluid filled is from the coelom of the body, and when the introvert is extended, the

system separate

circular muscles apply pressure to the contractile vessels or compensation sacs (Fig. 2.1B). This pressure

forces the fluid out of the sacs and up into the tentacles, thereby extending them. Posterior to the tentacular

I region of the introvert there be an area of chitinous hooks (Fig. 2.1A). These hooks be arranged in

may may

a pattern or scattered. The introvert is not always extended in preserved specimens, thus hindering the

efforts of taxonomists in the use of the tentacles as taxonomic characters. The introvert is withdrawn by the

i retractor muscles, which number from one to four, and can be seen through the body wall of the smaller abyssal specimens, or by dissection of the larger opaque animals (Fig. 2.1B). The body of the animal is

covered completely, but to varying degrees, by a cuticle. The cuticle of the trunk may have distinctive

i papillae and/or holdfasts, and may appear striated due to a constriction of the longitudinal muscles (Fig. 2.1B). The anterior and/or posterior regions of the trunk can also have a "shield" made of chitin, another

important taxonomic characteristic (Fig. 2.1A). The shape of the trunk is usually not of taxonomic importance

I due to the consla-iction of soft-bodied animals when they are preserved. Because of pressure changes, the bodies of one species can look very different from animal to animal. For taxonomic purposes it is best to use

internal structures along with any exterior characteristics which are unique.

I The nephridla are paired in most animals, occasionally numbering only one (Fig. 2.1B), and are located ventrally on the anterior portion of the trunk. In close proximity to the nephridiopores is the anus

(except in the genus Onchnesoma where it can be found on the introvert) (Fig. 2.1A). In the area of the anus

I a strand of muscle attached to the body wall extends down the center of the intestinal coil adding to its stability. This spindle muscle may end along the length of the gut, or continue on towards the posterior end

of the trunk (Fig. 2.1B). There may also be fastening or fixing muscles present to attach the oesophagus or

I the posterior region of the intestine to the body wall (Fig. 2.1B). The nerve cord runs ventrally down the entire length of the trunk, and off this cord branch the lateral nerves which extend to the body wall muscles

as well as to sensory receptors in the epidermis (Fig. 2.1B).

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I

m

L.'--':_ h

•--i I

! nc

_- cv I

it "r "- sm |

_, o -- reIrC i

• vr - fm

I

Im - in I

- Im

lm

PP I

CS

I

Figure 2.1. Morphology of a generalized sipunculan: A, external morphology; B, internal anatomy of dissected

specimen. (modified from Frank, 1983). I

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I Biology

I Sipunculans regenerate tentacles, as well as introvert, trunk and digestive tract parts. They can reproduce asexually through transverse fission of the body, although this is not common. Most sipunculans

are dioecious. The gametes arise from the coelomic lining and are released into the coelom for maturation.

I Obody. nce mature The males theyspawn are picked first. uThe p sesperm lectivelyin bythe thewaternephri prompts dia where the female they areto stored spawn.unAfter til released fertilization from the the

zygotes generally follow a prostomous development. Cleavage is spiral and holoblastic. Some species develop

I asecond free-living larvallecith stage,otrophic a planktonic troehopho pelagreosplharveraa whic larva,h dwhich evelopselonga intotesa juvandenilesettlesworm. to become Others da evel juvenile op a

sipunculan. After fertilization of the egg, a few species undergo direct development. These fertilized eggs

I asipunculan. ttach to the substratum, and the embryo develops into a vermiform individual which hatches as a juvenile

I Glossary

I

Anus. The opening in the cuticle where the Longitudinal muscles. The inner muscles of the

intestine ends through which solid waste is body wall which is a continuous sheet in

I excreted. trunk, andItinissome locatedcases anteriorly on the introvert. on the In mtogether ost animals, in somebutgecannera. be Itbundled is this or banding banded

most animals it will be in close proximity to that causes the cuticle of some animals to

i the nephridiopores, appear striated. Circular muscles. The outer muscle of the body Nephridia- Saclike organs located in the anterior

wall used to contract the body to evert the portion of the trunk used in excretion and as

I introvert and tentacles as well as to move gonoducts. the animal. The muscle may be grouped into

Nerve cord. The single cord from which lateral bands, or form a continuous sheet.

i nerves arise and extend to the body wall , Compensation system. A separate fluid filled muscles as well as to sensory receptors in

coelomic system of the body which the epidermis.

I regulates the extension of the tentacles. See also contractile vessels. Oesophagus. Region of the intestine between the

pharynx and the descending loop.

Contractile vessels or Compensation sacs. The

I bamuscles lloon-like squeeze structures when which the introvert the circul is atory Papillae. the Projection cuticle forming s of theraised epidermal bumps glands on the into

external body wall. They axe used for cuticle extended, forcing fluid through a closed

I aidsystemin feeding, into the tentacles, extending them for prod organs. uctioThey n, mmay ucus also secrehave tion aanchitinous d as sensory

outline ridge and can be used as holdfasts.

i Cuticle. The outer layer of the body wall, usually Pelagosphera larva. Second larval stage of some

thicker around the trunk and tapering

towards the mouth region of the introvert, sipunculans. Swims by means of a ciliated

metatroch and is often found in plankton

I Introveretractable, rt. The areausedof the for feeding body which and touc is hing. samples.

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Retractor muscles. Muscles connected to the Tentacles. The hollow extensions of the introvert i

body wall as well as to the introvert used to used in feeding, the arrangement of which

withdraw the introvert into the trunk region, may be an important taxonomic character. _l

II

Shield. Thickened cuticular areas or calcareous Troehophore larva. First larval stage of those

deposits on the anterior and sometimes animals not undergoing direct development. I

posterior trunk regions of the Free-living larva which will develop into a II

Aspidosiphonidae, used for protection and juvenile worm or metamorphose into a

I

burrowing, pelagosphera larva. it

Spindle muscle. The strand of muscle attached to Trunk. The region of the body of the animal I

the inner body wall in the area of the anus which is not retractable.

which runs through the intestinal coil and II

reattaches to the inner wall at varying points |

in different animals, giving stability to the

gut coil. i

List of Abbreviations on Figures l!I!

a, anus no, nuchal organ

cm, circular muscle nc, nerve cord l

cv, contractile vessel (compensation sac) np, nephridiopore

cs, caudal shield o, oesophagus I

dr, dorsal retractor pp, papillae

fm, fastening muscle rc, rectal caecum

g, gonad re, rectum !

h, hooks rm, retractor muscle

i, introvert sm, spindle muscle I_

I

in, intestine t, tentacle

lm, longitudinal muscle tr, trunk I

m, mouth vr, ventral retractor I

md, mid-dorsal line wm, wing muscle

n, nephridium I

List of Species from the MMS Surveys I

Nephasoma diaphanes (Gerould, 1913) Thysanocardia nigra (Ikeda, 1904) I

Phascolion lutense Selenka, 1885 Siphonosoma (Siphonosoma ) ingens (Fisher, 1947)

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I Key to the Sipuncula of the Santa Maria Basin

I 1A. Longitudinal muscles of the body wall gathered into separate or anastomosing bands ...................

........................................................................... Family Sipunculidae ............ Siphonosoma ingens

i 1B. Longitudinal muscle of body wall in a uniform continuous layer .................................................. 2

2A. A single nephridium present ............................. Family Phascolionidae ........... Phascolion lutense

m 2B. Two nephridia present Family Golfmgiidae... 3

I 3B. 3A. Tentacular Numerous tencrown taclesreduced present;to hooks short lobes; absentsmall, ....................................................... scattered hooks present .... Nephasoma Thysanocardia diaphanes nigra

i Descriptions of Species

l Nephasoma diaphanes (Gerould, 1913)

Figure 2.2

n Phascolosoma diaphanes Gerould, 1913:395.

Golfingia diaphanes: Cutler and Cutler, 1980a:425-453; 1980b:201-202.--Frank, 1983:15-16.----Cutler et

n al., 1984:269-270. Phascolosoma improvisum Geroutd, 1913:395-396.--Wesenberg-Lund, 1930:32-34; 1939:22-23.

i Golfingia improvisa: Wesenberg-Lund, 1955:11; 1963:110-111.--Murina, 1958:1625-1628; 1968:197; 1971:42; 1972:300; 1973:69; 1978:123.--Stephen and Edmonds, 1972:145.--Amor, 1975:113-

115.--Edmonds, 1976:222-224.

i Phascolosoma 467 minutum Th_el, 1911:31.--Sluiter, 1912:10-11.--Fischer, 1922a:34; 1922b:237; 1929:464- (part/m).--Wesenberg-Lund, 1930:30; 1932:9-10; 1937:12-13.

Golfingia minuta: Wesenberg-Lund, 1955:11; 1963:110-111.--Murina, 1957a:994-995; 1958:1628-1634;

I 1968:197; 1972:149-150.---Cutler, 1971:42; 1972:1973:155-159; 300; 1973:69; 11977a:143.---Cutler 974:235; 1976:65; 1978: and Cutler, 124.---Stephen 1979:957-958.--Saiz- and Edmonds,

Salinas, 1984:183.

I Golfingia sectiIe Murina, 1974:228-230.

Phascolosoma anceps Sluiter, 1912:10.--Wesenberg-Lund, 1925:90.

i Phascolosoma Golfingia cincta:cinctu Stephen re Geroanduld,Edmonds, 1913:3981972:137-138. -400, pl. 59, fig 6, text-fig. 8.

Phascolosoma sabellariae Gerould, 1913:392-395.

I Material Examined. California: Santa Maria Basin, off Point Buchon, Sta. 12 (3); off Point San

Luis, Sta. R-1 (77); off Point Sal, Sta. PJ-6 (1); off Purisima Point, Sta. 42 (1), Sta. 52 (80); off Point

m Arguello, sta. 65 (71).

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Lt ."*; _

in-- nc .•'"_'.%,,,_':__.__,'_;_'yP/

nc vr =;' :__

I

I

.5 mm I

Figure 2.2 Nephasoma diaphanes: A, internal, dissection of extreme anterior end; B, internal, dissection of

posterior end showing origin of retractor muscles; C, portion of posterior end showing papillae; D, I

hooks; E, reduced tentacles. (A-D after Cutler, 1973; E after Cutler et al., 1984).

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I Description. Small pear-shaped to cylindrical animals, trunks occasionally reaching length of 30

mm, usually 10 mm or less. Introvert approximately equal in length to trunk, but can vary from 50-150% of

I trunk anteriorlenpart gth. ofSkintrunk. translu Somecentanimals to opaque, withmay papillae exhibit on posterior longitudinal end w(Fig. rinkles 2.2C). or folds Tentacular on basecrown of introvert reducedandto

6-8 short lobes and 2 longer dorsal tentacles (Fig. 2.2E). Small, scattered, pale triangular hooks present (Fig.

i 2.2D). Two nephridia short, free at level of, or just posterior to, anus (Fig. 2.2A and B). Pair of ventral retractor muscles originating near posterior end in small animals, in mid-trunk in larger ones. Intestines in

double helix and with weakly developed spindle muscle within coil, not extending onto rectum.

I differs fromRemaNrks . diaphanes . There arediaphanes two subspecies in the nature of Nephasoma of the epidermis. diaphanesNephasoma , of which Ndiaphanes . diaphanesdiaphanes corrugatumhas

smooth, translucent to transparent skin, occasionally with raised pigmented papillae. Nephasoma diaphanes

I is(1986), often confused N. diaphanes taxonomically was often called with NGolfingia . minutum.minuta Prior .toThat the revision paper accep of Nephasoma ts the changebyofCutler statusandproposed Cutler

by Gibbs (1975) to redefine the species Golfingia minuta to include only those hermaphroditic populations

I from the shallow northeastern Atlantic. All other populations of dioecious animals are therefore referred to the species Nephasoma diaphanes.

Distribution. Cosmopolitan, cold water, most from bathyal and abyssal depths. The subspecies

I Nephasoma Red Seas fromdiaphanes 80-5900 corrugatum m. is found in the Atlantic and Pacific oceans and the Mediterranean and

I Phaseolion lutense Selenka, 1885

Figure 2.3

I Phascolion Iutense Selenka, 1885:16-17, pl.4, figs 22-23.--Fischer, 1928:484.--Murina, 1957b: 1781-1790,

figs. 4a-b, 5a-d; 1961:140-142, fig. 1; 1971:43; 1972:305-306; 1974:235; 1978:124.---Stephen and

I PhascolionEdmonds, canum 1972:180-181.----Cutler, Cutler and Curler, 1980a:454-456, 1977:145-146; fig. 2.- Cutler -Gibbs, and Cutler, 1985:321. 1985a:829-830, fig. 7b.

Phascolion species Cutler and Curler, 1980b:197.

I Material Examined. California: Santa Maria Basin, off Point Estero, Sta 2 (1); off Point Sal, Sta.

PJ-1 (3). Western Santa Barbara Channel, Sta. 86 (3).

I Description. External body surface smooth, with small papillae at anterior end and inconspicuous skin bodies in mid-trunk of some animals (Fig. 2.3B, D). Holdfast papillae absent. Tentacular crown reduced

to lobes (Fig. 2.3E). Most specimens beating pale hooks, 40-I50 m long (Fig. 2.3C). One nephridium

I present. Retractor muscles fused for most of length, with 3 or 4 distinct origins (Fig. 2.3A). Biology. This species is commonly found residing in gastropod shells.

Remarks. All material examined was originally named Onchnesoma sp. A.

I Type Location. South Pacific 53°55'S, 108°35'E at 3658m. Holotype: BMNH, no. 1885.12.3.22.

Distribution. Eastern Pacific; a deep-water species, in southern hemisphere (36-66°S) in Pacific

I Ocean off Argentina, southeastern Indian Ocean, northwestern Pacific and Bay of Biscay.

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_:..._-..-:.-_:...-.. . _

__ I

rm _" _._-':...d

-:"'..__,r;_

c D |

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Figure 2.3 Phascolion luten.r8: A, dissection showing internal anatomy; B, external view, C, hooks; ]3, glands

withandCutler,papillae;te1985a).E' ntacular lobes (A-C after Murina, 1975b; D, after Selenka, 1885; E, after Cutler I

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I Siphonosoma (Siphonosoma) ingens (Fisher, 1947)

Figure 2.4

I Siphonomecus ingens Fisher, 1947:365-368, pl. 15, fig. 15.

i Siphonosoma ingens: Fisher, 1952:382-385, pl. 20, fig. 21. Material Examined. California: Santa Maria Basin, off Point Sal, Sta. R-8 (1).

i without Description. Body long and slender; introvert much shorter than trunk; introvert with papillae but hooks. Body wall with coelomic extensions, longitudinal muscle gathered into bands (Fig. 2.4A).

Four retractor muscles and 2 nephridia present (Fig. 2.4B). Nuchal tentacles lacking, but oral disk with 12

double rows of tentacles around mouth (Fig. 2.4C). Spindle muscle attached near anus and at posterior end

I body. Villi present on contractile vessel. Posterior end of trunk with tubular, fusiform glandular bodies

of

(Fig. 2.4D). i

i

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"1 '\' \"

i

| .- :fro

tic

I

nc _

in r

| ,m

I Figure2.4 Siphonosoma ingens: A, internal, dissection of anterior portion of trunk; B, internal, dissection of

anterior end of paratype showing contracted retractor muscles, partly invaginated introvert; C,

I t19entacles; 47 D, internal, posterior trunk showing tubular, fusiform glandular bodies. (A-C after Fisher,

; D after Cutler and Cutler, t982).
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Remarks. This species is very similar toSiphonosoma mourense except that it has posterior fusiform •

bodies on the posterior end of the trunk. Both species occur in the North Pacific.

Type Locality. Morro Bay, San Luis Obispo County, California; intertidal. Holotype: USNM 20910.

Dis Itribution. Southern California.

Thysanoeardia nigra (Ikeda, 1904) I

Figure 2.5

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Phascolosoma nigrum Ikeda, 1904:3, text figs. 1, 25-27.--Sat6, 1934:247; 1939:409.---Chin, 1947:101.

Golfingia (Thysanocardia) nigra: Cutler and Cutler, 1981:68. Ill

Phascolosoma catherinae Sat6, 1937:154; 1939:406. II

Golfin gia (thysanocardia ) catherinae : Cutler, 1977b: 152. Illl

?Golfingia procera Fisher, 1952:402. I

Phascolosoma zenibakense Ikeda, 1924:29-30, fig. I.---Sat6, 1930:17-20, pl. 3 figs. 11-12, text-figs. 5;

1937:153-157; 1939:410.---Okuda, 1946:223. 1

Phascolosoma hyugensis Sat6, 1934:12-14, pl. 1, fig. 6, text-figs. 15-16. II

Phascolosoma hozawai Sat6, 1937:158-160, pl. 4, fig. 15, text-figs. 5-8.

Phascolosoma onagawa Sat6, 1937:156-158, pl. 4, fig. 14, text-figs. 3-4. Dwl

Golfingia macginitiei Fisher, 1952:402-404, pl. 26.

Golfingia pugettensis Fisher, 1952:401 .--Rice, 1967:144-170. 1

Material Examined. California: Santa Mafia Basin, off Point Arguello, Sta. 66 (3), Sta. 74 (3).

Description. A fairly large species with trunk up to 70 mm long. Introvert approximately twice as I

long as trunk, ranging from 1.5-2.5 times as long, difficult to measure when trunk constricted. Numerous

tentacles present, hooks absent (Fig. 2.5B). Number of tentacles varying with size of animal, but in large

specimens up to 24 festoons of 60 tentacles each, with another 30 tentacles surrounding nuchal organ. •

Tentacles darkly pigmented violet-mauve in life, fading to dark brown in preservative, sometimes fading II

completely after long preservation. Skin rough, often with wavy wrinkles, usually colored grey or rarely

black. Posterior end of trunk constricted, resembling a tail or a point. •

| Internally a pair of retractor muscles present originating between 55-75% of the distance to posterior

of trunk. Two nephxidia unattached, 25-70% of the trunk length, opening anteriorly to anus. Rectal caecum

usually present. Contractile vessel villi present along oesophagus, increasing in size and complexity, and i

becoming long and branched posteriorly (Fig. 2.5A).

Remarks. The pigmentation of the tentacles differentiates T. nigra from T. catherinae, though as

stated above, the use of pigmentation is not an ideal taxonomic character because it tends to fade in alcohol 1

and may be lacking entirely in smaller specimens. I

Type LocatiOn. According to Cutler and Cutler (1981) a lectotype had been deposited at the

University of Tokyo Zoological Museum, paralectotypes at the National Science Museum, Tokyo, and II

reference material at the National Museum of Natural History, Washington, D.C. I

Distribution. North Pacific: Philippines and Japan to California; Singapore. 1

I

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Figure 2.5 Thysanocardia nigra: A, internalview of dissected specimen;B,tentacles; C,papillafrom posterior

i of trunk. (A after Ikeda, 1924;B after Cutler and Cutler, 1981;C afterIkeda, 1904).

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Literature Cited I

Amor, A. 1975. Notas sobre Sipuncula de la Argentina, Brasil y Peru. Physis 34(88): 113-120. D1

Chamberlin, R.V. 1920. Notes on the sipunculids of Laguna Beach. Journal of Entymology and Zoology

12:30-31. i

Cutler, E.B. 1973. Sipuncula of the western North Atlantic. Bulletin of the American Museum of Natural i

History 152(3): 105-204.

i

Cutler, E.B. 1977a. The bathyal and abyssal Sipuncula. Galathea Report 14:135 -156. B

Curler, E.B. 1977b. Shallow water sipunculans collected by the Galathea Expedition (Sipuncula). Steenstrupia

4:151-155. I

Cutler, E.B. 1979. A reconsideration of the Golfingia subgenera Fisherana Stephen, Mitosiphon Fisher, and

Apionsoma Sluiter (Sipuncula). Zoological Journal of the Linnean Society 65:367-384.

Cutler, E.B. 1986. The family Sipunculidae (Sipuncula): body wall structure and phylogenetic relationships. J

Bulletin of Marine Science 38(3):488-497.

i

Cutler, E.B. 1994. The Sipuncula: Their Systematics, Biology, and Evolution. Comell University Press, I

Ithaca, New York. xvii + 453 pp.

Cutler, E.B., and N.J. Cutler. 1979. Madagascar and Indian Ocean Sipuncula. Bulletin du Mus6um National i

d'Histoire Naturelle, Paris 4e(1):941-990.

Curler, E.B., and N.J. Cutler. 1980a. Deep water Sipuncula from the Gulf of Gascogne. Journal of the i

Marine Biological Association of the United Kingdom 60:449-459. B

Cutler, E.B., and N.J. Cutler. 1980b. Sipuncula collected by the R/V "Vema." Journal of Zoology, London

190:193 -209. n

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Cutler, E.B., and N.J. Curler. 1981. A reconsideration of Sipunculanamed by I.Ikeda and H.Sat6. Publications

of the Seto Marine Biological Laboratory 26(113)51-93.

Cutler, E.B., and N.J. Cutler. 1982. A revision of the genus Siphonosoma (Sipuncula). Proceedings of the I_

Biological Society of Washington 95(4):748-762.

Cutler, E.B., and N.J. Cutler. 1983. An examination of the Phascolosoma subgenera Antillesoma, 1

Rueppellisoma, and Satonus (Sipuncula). Zoological Journal of the Linnean Society 77:175-187.

i

Cutler, E.B., and N.J. Cutler. 1985a. A revision of the generaPhascolion Th6el and Onchnesoma Koren and l

Danielssen (Sipuncula). Proceedings of the Biological Society of Washington 98(4):809-850. II

Curler, E.B., and N.J. Cutler. 1985b. A revision of the genera Sipunculus and Xenosiphon (Sipuncula). I

Zoological Journal of the Linnean Society 85:219-246. 1

Curler, E.B., and N.J. Cutler. 1987. A revision of the genus Golfingia (Sipuncula:Golf'mgiidae). Proceedings

of the Biological Society of Washington 100(4):735-761. i

i

Cutler, E.B., and N.J. Cutler. 1988. A revision of the genus Themiste (Sipuncula). Proceedings of the Biological

Society of Washington 101(4):741-766. i

Curler, E.B., and N.J. Curler. 1989. A revision of the genus Aspidosiphon (Sipuncula). Proceedings of the i

Biological Society of Washington 102(4):826-865.

i
I Cutler, E.B., N.J. Cutler, and P.E. Gibbs. 1983. A reconsideration of the Golfingia subgenera Golfingiella

and Siphonoides (Sipuncula). Proceedings of the Biological Society of Washington 96(4):669-694.

I Cutler, E.B., N.J. Cutler, and T. Nishikawa. 1984. The Sipuncula of Japan: Their systematics and distribution. Publications of the Seto Marine Biological Laboratory 29(4t6):249-322.

I Cutler, E.B., Zoology and P.E. 34(2): Gibbs. 162-173. 1985. A phylogenetic analysis of higher taxa in the phylum Sipuncula. Systematic

Cutler, E.B., and E. Jurczak. 1975. The sipunculan genus Lithacrosiphon Shipley, a taxonomic review.

I Zoological Journal of the Linnean Society 56(3):235-248.

Cutler, E.B., and V.V. Murina. 1977. On the sipunculan genus Nephasoma (Sipuncula, Golfingiidae).

i Proceedings of the Biological Society of Washington 99(4):547-573. Cutler, N.J. and E.B. Cutler. 1990. A revision of the subgenus Phascolosoma (Sipuncula: Phascolosoma).

Proceedings of the Biological Society of Washington 103(3):691-730. II

i Edmonds, S.J. 1976. Three sipunculan species (two new) from New Zealand. New Zealand Journal of

Marine and Freshwater Research 10(1):217-224.

I Fischer, W. 1922a. Gephyreen des Reichsmuseums zu Stockholm. Arkiv fSr Zoologi 14(19):1-39.

Fischer, W. 1922b. Gephyreen der Arktischen Meere. Wissenschaftliche Meeresuntersuchungen, Abteilung

I Helgoland 13:229-246. Fischer, W. 1928.1I. Uber zwei neue Siphonosoma-Arten der Wfirttembergischen Naturalien-Sammlung zu

Stuttgart. Zoologischer Anzeiger 76:138-143.

I Fischer, W. 1929. Die Sipunculiden, Priapuliden und Echiuriden der Arktis. In: F. Romer and F. Schaudinn

(eds.), Fauna Arctica, pp. 451-490.

I Fisher, W.K. 1947. New genera and species of Echiuridae and Sipunculid worms. Proceedings of the United States National Museum97:351-372.

I Fisher, W.K. 1952. The sipunculid worms of California and Baja California. Proceedings of the United States National Museum 102:371-450.

Frank, P.G. 1983. A checklist and bibliography of the Sipuncula from Canadian and adjacent waters. Syllogeus

I 46:1-47.

Gerould, J.H. 1913. The sipunculids of the eastern coast of North America. Proceedings of the United States

i National Museum 44:373-437. Gibbs, P.E. 1975. Gametogenesis and spawning in a hermaphroditic population of Golfingia minuta

(Sipuncula). Journal of the Marine Biological Association of the United Kingdom 55:69-82.

I Gibbs, P.E. 1985. On the genus Phascolion (Sipuncula) with particular reference to the North-East Atlantic

species. Journal of the Marine Biological Association of the United Kingdom 65:311-323.

I Gibbs, P.E., of Natural and E.BHistory . Cutler.(Zoology) 1987. A classification 52(1 ):43-58. of the phylum Sipuncula. Bulletin of the British Museum

i Gibbs, P.E., Zoologica E.B. Cutler, and N.J. Cutler. 1983. A review of the genus Thysanocardia Fisher (Sipuncula). Scripta 12(4):295-304.

Hartman, O., and J.L. Barnard. 1958. The benthic fauna of the deep basins off Southern California. Allan

I Hancock Pacific Expeditions 22(I): 1-67.
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Hartman, O., and J.L. Barnard. 1960. The benthic fauna of the deep basins off Southern California. Allan •

Hancock Pacific Expeditions 22(2):69-297.

Hyman, L.H. 1959. Phylum Sipunculida. Pp. 610-696 in: The Invertebrates. Volume 5. McGraw-Hill Book i

Co., New York. I

Ikeda, I. 1904. The Gephyrea of Japan. Journal of the College of Science, Imperial University, Tokyo •

20(4):1-87. I

Ikeda, I. 1924.2. Further notes on the Gephyrea of Japan, with descriptions of some new species from the

Marshall, Caroline and Palau Islands. Japanese Journal of Zoology 1:23-44. I

Murina, V.V. 1957a. Sipunculids collected on the fast trip of the complex Antarctic expedition on the "Ob"

in 1956. Zoologicheskii Zhurnal 36(7):992-998. i

Murina,V.V. 1957b. Abyssal sipunculids (genus Phascolion Th6el) of the north-western part of the Pacific

collected by "Vitj az" expeditions in 1950-1955. Zoologicheskii Zhumal 36(12): 1777-1791. I

Murina, V.V. 1958. On the systematics of two closely related species of the deep-water sipunculids of the I

genus Golfingia according to the materials of the "Vitj az" expeditions in 1950-1955. Zoologicheskii

Zhumal 37:1624-1633. •

I

Murina, V.V. 1961. On the geographic distribution of the abyssal sipunculidPhascolion lutense. Okeanologii

(Akademii Nauk SSSR) 1:140-142.

Murina V.V. 1968. New data concerning the occurrence of abyssal sipunculids in the Atlantic Ocean. i

Zoologicheskii Zhurnal 47(2): 195-199.

Murina V.V. 1971. On the occurrence of deep-sea sipunculids and priapulids in the Kurile Kamchatka I

Trench. Trudy Instituta Okeanologii 92:41-45. i

Murina V.V. 1972. Contribution of the sipunculid fauna of the Southern Hemisphere. Zoologii Instituta •

Akademii Nauk SSSR 11(19):294-314. I

Murina, V.V. 1973. The fauna of sipuncutids from the Peruvian-Chilean Trench. Zoologicheskii Zhurnal

5:66-71. I

Murina, V.V. 1974. Conlributions to the knowledge of the fauna of sipunculid worms from the South Atlantic

based on the data of the "Akademik Kurchatov" expedition in 1971. Trudy Instituta Okeanologii m

98:228-239. I

Murina, V.V. 1976. New and rare species of sipunculids from the East China Sea. Vestnik Zoologii 2:62-67. i

Murina, V.V. 1978. On the sipunculids and priapulids fauna of the southern Pacific. Trudy Instituta I

Okeanologii 113:120-131.

Rice, M.E. 1967. A comparative study of the development ofPhascoIosoma agassizii, Golfingia, and Themiste i

pyroides with a discussion of developmental patterns in the Sipuncula. Ophelia 4:143-171. D

Rice, M.E. 1975. Phylum Sipuncula. In: Smith, R.I. and J.T. Carlton (eds.), Light's Manual: Intertidal •

Invertebrates of the California Coast. Third Edition. University of California Press. pp 128-132. I

Ricketts, E.F., and J. Calvin. 1952. Between Pacific Tides. Stanford University, California.

Saiz-Salinas, J.I. 1984. Gusanos sipunculidos (Sipuncula) de varias localidades de la costa de Portugal. In I

V. Monteiro-Marques (ed.), ACTAS do 4th Simposio Iberico de Estudos do Benthos Marinho,

Associac_o des Estudantes da Faculdade de Ciencias de Lisboa, pp. 177-188. I
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I Sedgwick, A. 1898. A student's textbook of zoology. Swan Sonnenchien, London.

Selenka, E. 1885. Report on the Gephyrea collected by H.M.S. "Challenger" during 1873-1876. Report of

I the scientific results of the of Challenger, Zoology 13(36): 1-25.

voyage

Sluiter, C.P. 1912. Grphydens (Sipunculides et Echiurides) provenant des campagnes de l'hirondelle Alice,

I 1898-1910. Rrsultants des Campagnes Scientifique de Prince Albert 1 36:1-36. Stephen, A.C. 1964. A revision of the phylum Sipuncula. Annals and Magazine of Natural History (ser 13),

7:457-462.

I Stephen, A.C., and S.J. Edmonds. 1972. The phyla Sipuncula and Echiura. Trustees British Museum (Natural

History), London, 528pp.

I Threl, H. 1911. Priapulids and Sipunculids dredged by the Swedish Antarctic Expedition 1901-1903 and the phenomenon of bipolarity. Kungliga Svenska Vetenskaps-Akademiens Hartdlingar 47(1):3-36.

I Wesenberg-Lund, KcbenhavnE. 1925. Gephyreer. Conspectus Faunae Groenlandicae. Bianco Lunos Bogtrykkeri. 23:81-91.

Wesenberg-Lund, E. 1930. Priapulidae and Sipunculidae. The Danish "Ingolf' Expedition 4(7):1-44.

I Wesenberg-Lund, E. 1932. The Godthaab Expedition 1928 Gephyrea. Meddelelser om GrCnland 79(3): 1-

18.

I Wesenberg-Lund, E. 1937. The zoology of East Greenland Gephyreans. Meddeleser om Gr_nland 121(1):1- 25.

I Wesenberg-Lund, Gcteborgs E.Kungliga 1939. Gephyreans VetenskapsfromVitterhets-Samhalles Swedish waters in theHandlingar Museum ofFemte NaturalFoljden History6(6):3-35. of Gothenburg

Wesenberg-Lund, E. 1955. Reports of Lund University Chile Expedition, 1948-49. 19. Gephyrea from

I Chile. Lunds Universitets 2krsskrift 51(10): 1-23.

Wesenberg-Lund, E. 1963. South African sipurtculids and echiuroids from coastal waters. Videnskabelige

i Meddelelser fra Dansk Naturhistorisk Forening 125:101-146.

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I'IIIIIIlI
I

| 3. EcmtraA

I by

I John F. Pilger 1

I Introduction

The Echiura are small group of unsegmented, coelomate marine organisms. Although their

I research. distribution is cosmopolitan, they are uncommon and only a few species are known well through basic

I Morphology

I body or sausage-shaped typically an proboscis (homologous

The echiuran is ovoid and has extensible

withtheannelidprostomium).The proboscisdeciduousinmany specieasnd mustbetreatedwithcare

when foundattachedtothetrunk.The mouthisanterioron thetrunkatthebaseoftheproboscis.Few

I external characters are available for use in identification save the of setae and nephridiopores

presence

adjacent to the mid-ventral line near the anterior end. Longitudinal muscle bands may be present and

visible in the body wall of the think.

I The spacious fluid-filled coelom contains the loosely-suspended, elongated digestive tract, usually

filled with sediment pellets. A small collateral tube, the intestinal siphon, is present in most species. The

posterior end of the intestine may bear an enlarged bump called the intestinal caecum. Thin-walled, paired

I hind-gut sacs, vesicles, are present species, may simple or branched and frequently

anal inall The vesicles be

bear eiliated funnels. The proximal ends of the ventral setae project into the coelom and are equipped with

muscle bands that connect to the body wall and aid in manipulation. A strong interbasal muscle typically

I connects the proximal ends of the two setae. Nephridia modified for collection,

are use as gamete storage

and spawning organs, metanephridia. They axe attached to the body wall on the anterior ventral region

adjacent to the ventral nerve cord. The nephrostome frequently is modified into a characteristic form such

I as elongated, coiled strands or as fmabriae. Presumably these features aid in the selection of developed

gametes from the coelomic fluid so that they may be stored prior to spawning. Nephfidia filled with gametes

have been called storage organs and may be quite large relative to the length of the trunk.

I A blood vascular system is present in all echiurans except members of the Order Xenopneusta

(genus Urechis). The proboscis has a median vessel that carries blood anteriorly to the tip. At the tip the

median vessel splits to form paired lateral vessels and these return blood to the trunk. These vessels emerge

I in the trunk ventral vessel and follow the ventral cord the end of the

as a

single nerve tO posterior body. Just

anterior to the setae, theneuro-inteslinal vessel branches fromthe ventral vessel and extends to the esophagus.

On the way it may form a loop around the interbasal muscle. At the esophagus the vessel may surroundthe

I as a ring vessel. The dorsal vessel extends from here along the anterior and enters the base of the

gut gut

proboscis as the median vessel.

I

I t Department of Biology, Agnes Scott College, Decatur, Georgia 30030-3797.
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The location of the gonad is not always known in echiurans but where it is, it is unpaired and on the •

posterior region of the ventral vessel. Gametogenesis begins here but shortly the gametes become free

within the coelomic fluid where gametogenesis continues. The most mature gametes are removed selectively •

from the coelom by the nephrostome and stored in the modified nephridia until spawning. Fertilization is II

external and a trochophore larva is produced.

Taxonomic History I

l

Quatrcfages (1865) classified cchiurans with sipunculans and priapulids as the phylum Gephyrea l

because collectively they represented a "bridge" between the echinoderms and annelids. Later, echiurans

were considered as a class of the annelids (Sedgwick, 1898). Based on the detailed embryological studies II

and the suggestion of Newby (1940), Fisher (1946) erected the phylum Echiuroidea. Stephen (1964) renamed

the phylum Echiura.

The most recent classification system is that of Stephen and Edmonds (1972). It is modified from 1

I

Fisher (1946, 1949) and based on characteristics such as body wall musculature, proboscis morphology, m

vascular system anatomy, anal vesicle morphology, and the number, position, and morphology of setae and mlm

nephridia. They recognized 3 orders, 4 families, 34 genera, and 129 species. Additional species have been II

described.

Biology I

The presence of an elongate proboscis is an indication that the species is adeposit feeding member I

of benthic communities. In all species of this form, aburrowis formed in soft mud, coral, or rock, depending

on the species and habitat. U-shaped burrows are most common and may extend 10-20 cm into the sediment i

and 10 to 40 cm horizontally. When feeding the proboscis is extended from the burrow aperture onto 1

surrounding sediment where it explores the deposit and picks up particles for transport posteriorly to the

mouth. Discrimination and sorting of food by the proboscis has been documented in Bonellia (Jaccarini and 1

Schembri, 1977). Organic matter is extracted from this material in the digestive tract before being released 1

as compacted fecal pellets.

Members of the genus Urechis have a short, stubby proboscis and are not deposit feeders. Instead, •

they produce mucus nets within the burrow and pump water and suspended particulates through it by I

peristalsis. Intermittently, the net becomes clogged with trapped food, is consumed, and a new net is

constructed.

Commensals are common in the burrows of echiurans. Typically, these include polychaetes, small II

crustaceans, small mollusks, and hydroids. Endoparasitic protozoa, platyhelminths, nematodes, and annelids

have been described (see Stephen and Eximonds, 1972 for a list). 1

E I! chiurans may represent ecologically significant members of benthic communities. Listriolobus

pelodes formed dense accumulations in areas offshore from Santa Barbara, California and at the White's

Point sewer outfall, Palos Verdes, California. •

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I Glossary

I Anal vesic lyinglein-theA pair posterior of thin-walled coelom ahindgut nd connecting sacs Nephridiop nephridium. ore - The Gametes externalaxeopening releasedtothrotheugh

to the posterior digestive tract. They may be the pore during spawning.

I simple ciliated orcups branched or funnels. and usTheuallfunction y bear small of the Nephrostome - The coelomic opening to the

vesicles is unclear but may be involved in nephridium. The position of this structure

respiration by "inhaling and exhahng" the and the modification of the lips of the

I nephrostome are important

taxonomic

surrounding seawater, characters. The nephrostome may be basal

Ciliated funnel - A small ciliated cup-like (near the nephridiopore) or distal (away

I structure on the coelomic epithelium from the nephridiopore and towards the

present

of anal vesicles, coelom). The lips are ciliated and may be

modified as fimbriae or elongated threads.

I Caecum - A small pouch on the precloacal region They function to collect the most of the intestine in some echiurans. Its differentiated gametes from the coelom for

function is unknown, storage in the nephridium prior to spawning.

I Dorsal vessel - That portion of the vascular Neuro-intestinal vessel The blood vessel that

system lying on the dorsal side of the branches from the anterior ventral vessel

posterior foregut and carrying blood and carries blood to the dorsal vessel. It may

I form a loop around the interbasai muscle.

anteriorly to the median vessel.

Interbasal muscle - A prominent muscle band Papillae - Prominent tubercles present on the

I connecting the proximal ends of the setae, exterior of the body of most echiurans. Absent in some species. Larger ones usually are present at the

Longitudinal muscle bands - Thickening of the anterior and posterior ends of the trunk.

I longitudinal muscle layer of the body wall Papillae may be glandular. into bands. The bands usually are visible Proboscis - The muscular and highly extensible

externally and are important taxonomic structure that projects anteriorly from the

I area dorsal to the mouth. In some literature

characters.

Lateral setal musdes - Muscles located at the it may be referred to as the "prostomium".

i body wall where the setae emerge into the The organ is used for feeding, burrowing, coelom. They extend laterally and may and perhaps for respiration. The ventral

serve to separate the setae. An uncommon surface has mucus glands, is ciliated and

character, functions to carry sediment particles to the

I mucous strings.

mouth in

Lateral vessels - The paired blood vessels that Although adult echiurans probably do not

return blood from the anterior tip of the leave their burrows intentionally, specimens

I proboscis to the trunk, requiring new burrows will use the

Median vessel - That vessel carrying blood from proboscis in the burrowing process first to

the dorsal vessel and the trunk to the tip of scoop out a shallow depression and later as a

I wedge and terminal anchor.

the proboscis.

Proboscides are morphologically variable

Metanephridium - The modified excretory organ and deciduous in some species. In the genus

I that sorts and stores gametes prior to Urechis the proboscis is reduced to a stubby

spawning. Usually present as one or a few upper lip. In many bonellids, the proboscis

pairs but, in some species, they may be is bifid at the tip while in other bonellids

I unpaired or numerous (as many as 400). anunbranched. d in the echiurids it is elongated and

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Ring vessel - The point where the neuro-intestinal Storage organ - A term used in the literature to •

vessel surrounds the posterior foregut and describe the portion of the metanephridium

connects to the dorsal vessel. The structure used to store gametes prior to spawning. It •

is present in varying degrees of has been referred to as a "segmental organ" II

development and is absent in some species, because it is variable in diameter when it is

Setae - Hooked or bristle-like structures present in filled with gametes. I

the ventral body wall posterior to the mouth. Ventral vessel - The blood vessel that is attached

Typically paired. Setae encircle the anus in to the dorsal surface of the ventral nerve

the genera Echiurus (two rings of setae) and cord and runs posteriorly to the end of the I

Urechis (one ring of setae), trunk. i

Siphon - A small collateral tube associated with •

much of the midgut. The anterior and |

posterior ends join with the alimentary tract

as a ciliated groove. 1

HI

Abbreviations Used in the Figures I

a, anus n, nephridium

av, anal vesicle niv, neuro-intestinal vessel l

cg, ciliated groove nl, anterior nerve loop

dv, dorsal vessel np, nephridiopore m

g, gonad p, proboscis i

gi, gizzard pap, papillae

i, intestine ph, pharynx 1

II ic, intestinal caecum s, setae

im, interbasal muscle ss, secondary seta •

lm, longitudinal muscle vbv, ventral blood vessel I

m, mouth vnc, ventral nerve cord

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I

I Key to the Orders, Families, Genera, and Species of Echiura

from the Santa Maria Basin and Western Santa Barbara Channel

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1A. Open vascular system, coelom with hemoglobin-containing corpuscles; posterior intestine thinwalled and functioning in respiration, anal vesicles present; proboscis present as a stubby upper lip.

I ............................................................................................................................ Order Xenopneusta

lB. Closed vascular system; anal vesicles present but posterior intestine not otherwise modified for

I respiration; proboscis elongate but may be deciduous ............................... Order Echiuroinea ..... 2

2A. Species with sexual dimorphism, males very small and commensal on or in female; proboscis bifid,

I cylindrical or triangular (may be deciduous or absent); anal vesicles usually branched; 1or 2 nephridia present .................................................................................... Family Bonellidae: Nellobia eusoma

2B. Species without sexual dimorphism; proboscis not bifid; anal vesicles saclike; 2 or 4 nephridia

I present; longitudinal muscle bands be present Family Echiuridae 3

may

I 3A. Longitudinal muscles grouped into bands; nephrostomal lips elongate ......................................... 4

3B. Longitudinal muscles not in bands; nephrostome broad funnel with frilled rim ..............................

...................................................................................................................... Arhynchite californicus |

4A. Six longitudinal muscle bands present; with 2 neplu'idia ........................... Listriolobus hexamyotus

I 4B. Eight longitudinal muscle bands present; with 4 nephridia .............................. Listriolobus pelodes

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Description of Species I

listriolobus pelodes Fisher, 1946

Figure 3.1 I

Listriolobus pelodes Fisher, 1946:234-240.--Bamard and Hartman, 1959:1-16.--Pilger, 1980:129-142. 1

Material Examined.California, off Point San Luis, MMS Phase ISta. 21,49 m, 1specimen (USNM);

1 specimen (SBMNH).

Description. Trunk 40-60 mm long, 12-25 mm in diameter (Figure 3.1A). Body wall translucent II

with 8 longitudinal muscle bands; bands may be less distinct in small specimens. Small papillae present in

small circular folds at anterior and posterior ends. Proboscis unbranched and highly extensible. Two hooked •

ventral setae with interbasal muscle (Figure 3.1B). Two pairs of nephridia each with paired, elongated, and

coiled nephrostomal lips. Two anal vesicles capable of great extension and bearing small, scattered ciliated

funnels (Figure 3.1C). Intestinal caecum present near anus. Vascular system with interbasal muscle loop •

and ring vessels. Gonad on posterior-most portion of ventral blood vessel atjunction with intestinal caecum. 1

Small coelomic oocytes present in clusters, large oocytes single.

Biology. Listriolobus pelodes is a deposit feeding echiuran common in fine-grained sediments at 1

depths from 18-155 m. Particularly dense though temporally variable populations have been described from

the shelf off Santa Barbara, California (Barnard and Hartman, 1959) and Palos Verdes, California (Pilger,

1980; Stull, Haydock, and Montague, 1986). The reproductive biology of specimens from southern California I

has been described (Pilger, 1980). M

Remarks. Listriolobus pelodes is distinct from L. hexamyotus in having eight longitudinal muscle 1

bands instead of six. "Small phase" specimens 7-15 mm long and sexually mature were described by Fisher 1

and also found by Hartman (Fisher, 1946). Although Fisher (1946) did not indicate anatomical differences U

sufficient to warrant taxonomic distinction, Pilger (1980) noted that their reproductive biology may be

different from larger specimens. 1

1

Type Locality and Type Specimens. Monterey Bay, California, moderate depth, fine sand, from

the stomach of flounders. Type specimen deposited in National Museum of Natural History (USNM 20608). 1

Distribution. West coast of North America from northern California to Baja California, Mexico. I

l.istriolobus hexamyotus Fisher, 1949 1

I

Figure 3.2

Listriolobus hexamyotus Fisher, 1949:484-485, pl. 29. I

i

Material Examined. California, Santa MariaBasin, off Point Sal, MMS Phase II Sta. R-7, 565 m, I

1 specimen (USNM); offPoint Arguello, MMS Phase I Sta. 62, 582m, 1 specimen (SBMNH).W.-- Western l

Santa Barbara Channel, off Point Conception, MMS Phase 1, Sta. 83,444 m, 1 specimen (USNM).

Description. Trunk elongate, 50 mm x 15 mm (Figure 3.2A). Small papillae present. Six •

inconspicuous longitudinal muscle bands. Proboscis deciduous, often missing (Figure 3.2B). Lateral edges |

of proximal portion surround mouth ventrally. Two setae with hooked tips pointing toward midline. Secondary

setae forming alongside primaries in some specimens. Strong interbasal muscle and lateral setal muscles •

present (Figure 3.2C). Pharynx attached to ventral body wall by mesenteric sheet. Foregut shorter than L.

|

pelodes. Anal vesicles thin-walled. Ciliated funnels not observed. One pair nephridia, each with 2 elongated

but uncoiled nephrostomal lips. Neurointestinal vessel not looping around interbasal muscle. Small, •

incomplete ring vessel present. |

72 !
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Figure 3.1. Listriolobus pelodes: A, external anatomy, ventral view (after Pilger, 1980). B, anterior internal

I ana 1980) tomy, dorsal view (after Fisher, 1946); C, posterior internal anatomy, dorsal view (after Pilger, .

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ph

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Figure 3.2. Listriolobus hexamyotus: A, external anatomy, ventral view; B, proboscis, detached, ventral view; I

C, anterior internal anatomy, dorsal view.

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74 I
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!i Biology. While no direct field observations have been made of this species, from its morphology

one can infer that it is a deposit-feeding species and probably forms U-shaped burrows in the soft muddy

I benthos. Remarks. Listriolobus hexamyotus differs from L pelodes in having one pair of nephridia, six

longitudinal muscle bands instead of eight, a smaller caecum and shorter foregut. The neuro-intestinal

I vessel does not form a loop around the interbasal muscle as in L. pelodes.

Type Locality and Type Specimens. Albatross Sta. 4339, San Diego, California, 574-735 rn, green

mud. Type specimen deposited in National Museum of Natural History (USNM 21079).

I Distribution. Southern California mainland shelf, lower 400-800 bight slope zone, m.

I Arhynchite californicus Fisher, 1949 Figure 3.3

I Arhynchite californicus Fisher, 1949:486-487

Material Examined. California, off Purisima Point, Sta. R-6,410 m (USNM); off Point Conception,

I MMS Phase I Sta. 76, 1 specimen (SBMNH).-- Western Santa Barbara Channel, MMS Phase 1, Sta. 86, 197 m, 1 specimen (USNM).

Description. Trunk elongate with transverse rows of glands present and separated by wrinkles

I (Figure 3.3A). Body wail thickened, especially at ends. Muscle layers distinguishable in cut sections of

body wall. Longitudinal muscle bands absent. Two hooked ventral setae located close to anterior end.

i Proboscis unbranched, fan-shaped distally, deciduous and often missing. Lateral edges meet ventrally at

proximal end (Figure 3.3B). Well-developed interbasal muscle passes through loop of neuro-intestinai vessel.

Gut long, attached to body wall by many mesenteric strands. Presiphonal ciliated groove absent. Two nephridia

attached to body wall close to setae (Figure 3.3C). Extra nephridia reported in some specimens. Nephrostome

within a stalked funnel with frilled rim. Anal vesicles long (up to half body length), anchored proximally by

many filaments. Scattered ciliated cups present. Neurointestinai vessel forms loop a_roundinterbasal muscle

and then attaches to gut. Distinct ring vessel not visible on gut.

I Biology. No direct studies of this species have been conducted. It is a deposit feeding form presumed

to make U-shaped burrows in soft muds. It is rarely found in water as shallow as 50 m. Typically it lives

between 100 and 500 m depth.

I Remarks. The genus was erected because of the many specimens were collected without a proboscis

before the deciduous nature of this structure was known. It is distinctive in that it is the only species in

i southern variable trait California (Thompson, withoutt979). longituDiffer dinals mfromuscleA.bands. pugettensis The presence Fisher 1949 or absence in thisotrait f the and ringthevessel absence may beof aa

ciliated groove in the presiphonal intestine.

I gray mud.TyType pe Locspecimen ality anddeposited Type SpeincimNational ens. Albatross MuseumSta. of Natural 4525, MHistory onterey (USNM Bay, California, 21085). 440 m., soft

Distribution. Monterey Bay, California south. Typically below shelf depths, 100-500 m.

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Figure 3.3. Arhynchite californicus: A, external anatomy, ventral view; B, proboscis, ventral view; C, anterior

internal anatomy, dorsal view. I

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76 I
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I NeUobia eusoma Fisher, 1946

i Figure 3.4

Nellobia eusoma Fisher, 1946:258-259.

l Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. R-5, 154 m (1). Description. Female. Trunk35mm x 19mm. Body wall pigmented, thickest at anterior and posterior

ends (Figure 3.4A). Longitudinal muscle bands absent. Single nephridiopore conspicuous, on left of ventral

I groove setae slightly at tip (Figure 3.4B). Single nephridium

midline. Genital and absent. Proboscis bifid left

with simple, basal, projected posteriorly nephrostome (Figure 3.4C). Pharynx with many radiating muscle

strands to anterior body wall. Cloaca and hind-gut enlarged. Anal vesicles crescent-shaped sacs on each side

I of cloaca, with unequal dendritic branches on lateral border; smaller branches with numerous small digitiform projections, each capped by ciliated funnel. Vascular system as in other bonellids. Dorsal and neuro-intestinal

Ill vessels join through capillary anastomosis on gut rather than through ring vessel. Gonad on venlxal vessel,

II extending as far anterior as junction of neuro-intestinal vessel with ventral nerve cord. II

Male. Three found in nephridiopore of type specimen by Thompson (1979). Males possess ventral

' Biology. Little has been reported about this species. Stephen and Edmonds (1972) report that it is

setae.

only known from the holotype but some local knowledge exists within its range and where benthic studies

i hain vthee been 100-500m regularldepth y conrange. ducted.There Thomispsnooneviden (1979) ce rep to suggest orted itthat fromitslisoanything pe areas wbutithaArhy deposit nchitefeeding californicus species

thatlives in a U-shaped burrow. The specimen examined was collected in January, 1988 and had its nephridium

I full of yoRemark lky eggs, s. indicating Fisher (1946) thaterected spawningthiswgenus as imminent to encompass . the single specimen collected in Japan. It

differed from many other bonellids in that it has a unique anal vesicle morphology and from all other

I bonellids by the presence of an enlarged muscular cloaca and enlarged terminal hindgut. Type Locality and Type Specimen.Albatross Sta. 5021, (48°32'30q_1145 , °08'45_E). Sea of Okhotsk,

Japan. 130 m. Type specimen deposited in National Museum of Natural History (USNM 20605).

I Distribution. Bering Sea (USNM 29998); Puget Sound, Washington; Monterey Bay and southern California mainland bight.

l

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Figure 3.4. Nellobia eusoma: A, external anatomy, ventral view; B, proboscis, ventral view; C, anterior internal i

anatomy, dorsal view. t

I
I Literature Cited

i Barnard,structure. J. L. and O. PacHartman ific Naturalist 1959. The 1:1-16. sea bottom off Santa Barbara, California: Biomass and community

i Fisher, W. Museum K. 1946. Echiuroid worms of the North Pacific Ocean. Proceedings of the United States National 96:215-292, pls. 20-37.

Fisher, W. K. 1949. Additions to the echiuroid fauna of the North Pacific Ocean. Proceedings of the United

I States National Museum 99:479-497, pls. 28-34.

Jaccarini, V. and P. J. Schembri 1977. Locomotory and other movements of the proboscis of Bonellia

i Pilger, viridis. Journal of Zoology, London 182:467-476. J. F. 1980. The annual cycle of oogenesis, spawning, and larval settlement of the echiuran Listriolobus

pelodes off southern California. Pacific Science 34:129-142.

I Quatrefages, A. de 1865. Notes sur la classification des Annelides. Comptes rendus hebdomadaires de

l'Academie des S6ances Paris 40:586-600.

i Sedgwick, A. 1898. A Student's Textbook of Zoology. 1. Swan Sonnenschein, London.

Stephen, A. C. 1964. A revision of the classification of the phylum Sipuncula. Annals and Magazine of

i Natural History, series 13, vol. 7:457-462. Stephen, A. C. and S. Edmonds 1972. The Phyla Sipuncula and Echiura. British Museum (Natural History),

London. 528pp.

I Stull, J. K., I. Haydock, and D. E. Montagne 1968a. The effects ofListriolobuspeIodes (Echiura) on coastal

shelf benthic communities and sediments modified by a major wastewater discharge. Estuarine and

coastal Shelf Science 22:1-17.

I Stull, J. K., I. Haydock, R. W.Smith, and D. E. Montagne 1968b. Long-term changes in the benthic community

on the coastal shelf of Palos Verdes, southern California. Marine Biology 91:539-55 I.

I Thompson, B. 1979. Southern California Echiura. Proceedimas Taxonomic Standardization Program.

Southern California Coastal Water Research Project 7:1-16.

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I 4. PHYLUMECHINODERMATA

| by

Andrew Lissner 1and Debra Hart 2

l Introduction

i The phylum Echinodermata contains some of the more familiar marine invertebrates including

brittlestars (Class Ophiuroidea); seastars (Class Asteroidea); sea cucumbers (Class Holothuroidea); feather

i Primary stars andreferences sea lilies (utilized Class Crinoidea); for this section and searea Ziesenhenne urchins, heart(1941), urchins,Hyman and sand (1955), dollBarnes ars (Class(1980), Echinoide Durhaa).m

et al. (1980), Feder (1980), Muscat (1980), Luke (1982), Lawrence (1987), and Maluf (1988). There are

I approximately 6,700 species of echinoderms, all of which inhabit the marine environment. Of these species, the majority are represented by brittlestars and seastars (approximately 2,000 species each), followed by sea

cucumbers (1,200 species), echinoids (800 species), and crinoids (700 species).

i Echinoderms occur worldwide in all benthic marine habitats from the intertidal to the deep sea. The general body form of the brittlestars and seastars, comprising the subphylum Asterozoa, consists of arms

radiating from a central disc. Compared to seastars, brittlestar arms typically are proportionately longer and

I set off from the disk, and they lack central arm grooves. Echinoids do not possess arms like seastars and brittlestars. Instead, the body has a spherical to greatly flattened appearance, with a characteristic solid test.

Sea cucumbers have elongate, worm-shaped bodies, also without arms, and the body has a less rigid, more

I leathery primitive texture echinoderms. than other Theirechinoderms. body structureTheoften crinoids consists are cons of anidereattachment d the oldest stalkandandprobably a crown theof most arms

similar to those of the seastars and brittlestars; however, many species are unstalked and relatively motile.

i Most echinoderm species axe relatively large (diameters of at least several centimeters) and possess

a distinctive, pentamerous radial symmetry in the adult stages. This symmetry is derived secondarily from

an initial bilateral symmetry apparent during the larval phase. Thus, echinoderms differ distinctly from

I other radiate phyla such as sponges, cnidarians, and ctenophores in that they exhibit a more complex life cycle, typically undergoing a radical metamorphosis from a pelagic, bilaterally symmetric larval stage to a

sedentary, radially symmetrical adult form. In addition, echinoderms are structurally more complex than

I these other radiates, and are true coelomates. All echinoderms retain an internal skeleton composed of calcareous ossicles. Endoskeleton ossicles

may be separate articulations, as in seastars and brittlestars, or fused into a rigid theca or test as seen in sea

i urchins and sand dollars. In most sea cucumber species the ossicles or spicules are reduced to microscopic size within the leathery epidermis. The name echinoderm, or spiny skin, refers to the spines or tubercles

projecting from the endoskeleton, giving the body surface a bumpy or spiny appearance.

I possess aunique or system composed a

Echinoderms water-vascular ambulacral of series of coelomic

canals throughout the body. Seawater enters the echinoderm's water-vascular system through a sieve plate

(madreporite) and is transported through canals via muscular action and hydraulic pressure to the terminal

I podia or tube feet. Historically, the podia were believed to function in food collection and although

transport,

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Science Applications International Corporation, 10260 Campus Point Drive, San Diego, CA 92121

I _Science Applications International Corporation, 1710 Goodridge Drive,McLean, VA 22102
this has evolved additionally to a locomotory function in most taxa. Feeding habits of echinoderms arequite 'it

diverse. Most species of seastars are carnivorous and many are key predators. Regular urchins are mostly

herbivorous, although some function as scavengers or filter their food, while irregular (heart urchins) ingest i

sediments and associated organic material directly from the substrate. Brittlestars may be scavengers, deposit I

feeders, filter feeders or, in some cases, may employ all three methods. Sea cucumbers are primarily

suspension or deposit feeders, while the crinoids are strictly suspension feeders. Digestion in echinoderms /

is carried out in a well-developed digestive tract located in the coelom. They have a complete digestive II

system, although no true excretory system exists. Nitrogenous wastes are diffused through thin surfaces at

the body/seawater interface such as the podia, il

Most echinoderms are dioecious with fertilization occurring externally in the surrounding seawater. I

Reproductive strategies include (1) production of large numbers of small eggs that develop into freeswimming, plankton-feeding larvae (most seastars, brittlestars, sea cucumbers and echinoids); (2) free- i[

swimming, yolk-filled larvae that do not require planktonic food (some brittlestars); and (3) production of a U

few yolky eggs that are brooded to the juvenile stage, thereby eliminating the free-swimming stage (brittlestars

and some cold water species of seastars, sea cucumbers, crinoids, and eckinoids).

The type and size of ossicles or plates, body shape, the number and length of arms, and, in fresh II

specimens, color, often are used as key taxonomic features for echinoderms. Class-specific features are

presented separately by group in the following chapters, i

Sources of Material i

Four species of Echinoidea, eight species of Asteroidea, 28 species of Holothuroidea, 19 species of /

Ophiuroideaan,d one species of Crinoidea are treated in this volume. All specimens were collected during i

I

the MMS Phase I and Phase U studies of the Santa Mafia Basin and western Santa Barbara Channel,

predominantly from soft-substrate stations. Exceptions are one species of asteroid, Henricia leviuscula

annec that was collected from both soft and hard substrate, and four ophiuroids (Gorgonocephalus itens,

eucnemis, OphiophoIis, baked, Ophiacantha diplasia, and Ophiothrix spiculata ) and the crinoid (Florometra

U

serratissima) collected from hard substrate.

General accounts of the study areas and descriptions of characteristic communities are presented in i

Blake (1993), Blake and Lissner (1993), Lissner and Benech (1993), Lissner et al. 1991, and SAIC (1986,

1993). m

A list of stations, navigational coordinates, and bottom depths from which the specimens were i

collected are presented in the appendix to this volume. Soft-substrate and hard-substrate specimens were

collected using a box corer and a manipulator arm on a manned submersible, respectively. /I

Individual samples were selected as voucher specimens by the Phase I and Phase II contractors I

(SAIC and Battelle, respectively). When sufficient material was available, two sets of vouchers were

established: (1) a primary collection deposited with the National Museum of Natural History, Smithsonian !

Institution (designated as USNM on the specimen labels), and (2) a secondary collection deposited with the II

Santa Barbara Museum of Natural History (SBMNH) or the Natural History Museum of Los Angeles County

(designated as LACMNH). I

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I Acknowledgments

I This study was funded by the Pacific Outer Continental Shelf Region of the Minerals Management Service, U.S. Department of the Interior, Washington, D.C., under Contract No. 14-35-0001-30484. Michael

Hart and Danny Heilprin aided in the literature review for species information and Debra Etchen and Millie

I Agostino assisted with editing and text entry.

i Literature Cited

Barnes, R. 1980. Invertebrate Zoology. 4th Edition. Philadelphia, PA., Saunters. 1089 pp.

t Blake, J. 1993. Introduction to the Taxonomic Atlas. Pages 1-16 In J. Blake and A. Lissner (eds.) Taxonomic

Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Volume

i 1.Introduction, Benthic Ecology, Oceanography, Platyhelminthes, and Nemertea. Contract No. 14-

12-0001-30484. Final Report U.S. Department of the Interior, Minerals Management Service, Pacific

OCS Region, Camarillo, CA.

i Blake, J. and A. Lissner. 1993. Physical Description of the Santa Maria Basin and Western Santa Barbara Channel. Pages 21-31 In J. Blake and A. Lissner (eds.) Taxonomic Atlas of the Benthic Fauna of the

Santa Maria Basin and Western Santa Barbara Channel. Volume 1.Introduction, Benthic Ecology,

t Ocean Dep ography, Platyhelminthes, and Nemertea. Contract No. 14-12-0001-30484. Final Report U.S. artment of the Interior, Minerals Management Service, Pacific OCS Region, Camarillo, CA.

i Hyman, Lawrence,L. 1955. The Invertebrates: Echinodermata. McGraw-Hill Book Company. New York, NY. 763 pp. J. 1987. A Functional Biology of Echinoderms. The Johns Hopkins University Press, Baltimore,

MD. 340 pp.

I Lissner, A. andS. Benech. 1993. Benthic Hard-Substrate Community Ecology of the Santa Mafia Basin and

Western Santa Barbara Channel. Pages 47-52 In: J. Blake and A. Lissner (eds.) Taxonomic Atlas of

i the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Volume 1. Introduction, Benthic Ecology, Oceanography, Platyhelminthes, and Nemertea. Contract No. 14-

12-0001-30484. Final Report U.S. Department of the Interior, Minerals Management Service, Pacific

OCS Region, Camarillo, CA.

i Lissner, A., G. Taghon, D. Diener, S. Schroeter, and J. Dixon. 1991. Recolonization of deep-water hardsubstrate communities: Potential impacts from oil and gas development. Ecological Applications

1:258-267.

Lissner, A. and J. Dorsey. 1986. Deep-water biological assemblages of a hard-bottom bank-ridge complex

of the southern California continental borderland. Bulletin of the Southern California of Sciences

i 85:87-101.

• Luke, S. 1982. Catalog of the benthic invertebrate collections: Echinodermata. Scripps Institution of

' I Oceanography Reference Series 82-5:2-24, 44-54. I

Maluf, L. 1988. Composition and Distribution of Central Eastern Pacific Echinoderms. Technical Report

Number 2, Natural History Museum of Los Angeles County, Los Angeles, CA. 242 pp.

I Muscat, A. 1980. Echinodermata. Pages 263-273 In: D. Straughan and R. Klink (eds.) A taxonomic listing

of common marine invertebrate species from southern California. Technical Reports of the Allan

I Hancock Foundation 3.
!

SAIC (Science Applications International Corporation). 1986. Assessment of Long-Term Changes in 1

Biological Communities in the Santa Maria Basin and Western Santa Barbara Channel: Phase I.

CManagement ontract No. Service, 14-12-00Pacific 01-3003OCS 2. Region, Final RepCamarillo, ort to U.CA. S. Department of the Interior, Minerals I

SAIC. 1993. Monitoring Assessment of Long-Term Changes in Biological Communities in the Santa Maria

Basin: Phase nq. Contract No. 14-35-0001-30584. Final Year One Report to U.S. Department of the !

Interior, Minerals Management Service, Pacific OCS Region, Camarillo, CA.

Ziesenhenechinoderms. ne, F. 1941.BulleSome tin ofnothe tes Southern on the dist California ribution Academy records ooff little-know Sciences 40:117-120. n Southern Californian

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i 5. CLASS CRINOIDEA

| uy

Gordon Hendler 1

I

I Introduction

Worldwide there are approximately 700 species of Crinoidea, including 100 species of stalked sea

i lilies which are limited to the deep and 600 species of unstalked feather stars which are most diverse in

sea,

tropical reef habitats. In the Central Eastern Pacific Region, from southern California to southern Peru only

12 crinoid species are known. Six are stalked forms, and the 6 feather stars include 1 thalassometrid and 5

I[ antedonid species (Maluf, 1988). All are restricted to deep water, except for Florometra serratissima, the

m only crinoid collected in the Mineral Management Service (MMS) Phase I and Phase II surveys, which can

occur near shore at the northern end of its range.

I The cornerstones of modern research on crinoids are the monumental Challenger reports (Carpenter,

1884, 1888) and A.H. Clark's invaluable monograph. The latter was published in 5 parts between 1915 and

i 1967, with the last volume coauthored by A.M. Clark, and brings together information on all aspects of the class. A more up-to-date compendium on the Crinoidea is available in the Treatise on Invertebrate

Paleontology (Moore and Teichert, 1978). Messing and Dearborn (1990) and Hendler et al. (1995) provide

a brief introduction to the crinoids and other echinoderm classes. The nutrition of the group is reviewed by

I Jangoux (1982), by Meyer (1983), reproduction by

and Lawrence biotic interactions and Ausich Holland

(1991), and aspects relating to coral reef ecology are covered by Birkeland (1989).

I History

i A.H. Clarkaccompanied the Albatross on a cruise through the Bering Sea to Japan, and he recalled

that "During the 1906 cruise of the Albatross I handled tens of thousands of specimens; several times I saw

i the forward deck of the steamer literally buried under several tons of individuals...everywhere we went we

found crinoids; we dredged them at all depths. My ideas of the comparative importance of the recent forms

underwent a total change; surely a group so abundant...cannot be considered as decadent or degenerate"

(A.H. Clark, 1915:4). In the following years he described the Albatross material, and examined nearly every

t collection of crinoids from around the the results in his major world, compiling monograph.

In the first volume on the Comatulida, A.H. Clark(1915) summarized the history of crinoid studies

i beginning was writtenwith the writings of Fabius Columna in 1592. The earliest publication on Eastern Pacific crinoids by Hartlaub (1895), but as A.H. Clark (1915:56) claimed, the west coast of the Americas

remained "terra incognita so far as crinoids were concerned until 1907, in which year many species were

, In described from the region" (by Clark himself). Developments in the taxonomy of Eastern Pacific crinoids

I! can be traced through his first and last papers on the fauna (Clark, 1907; Clark and Clark, 1967).

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JInvertebrate Zoology Section, Natural History Museum of Los Angeles County, 9130Exposition Boulevard, Los Angeles,

I California 90007.
!

Systematics !

The classification of Crinoidea perhaps in widest use is the system outlined by Ubaghs (1978), and l

a history of crinoid systematics is provided by Lane (1978). Simms (1988) offered a contemporary overview

of crinoid systematics, in which he reduced the living comatutids to infraordinal status "Comatulidia" within

an Order Isocrinida. In the present treatment, Ubagh's scheme is retained, wherein the Class Articulata •

represents all living crinoids and the Order Comatulida includes all the feather stars. In his organization II

there are 16 families of Recent comatulids, and over 130 extant genera. _l

m

Morphology

Featherstars range in size from 3-50 mm body diameter with arms 50-550 mm in length. Most have l

10 arms or more, and depending on the species the number of arms can reach 200.

The soft visceral mass of the crinoid, contained in a naked or weakly calcified tegmen (=disk), rests 1

on supporting rings of ossicles composing a calyx: In the sea lilies the calyx caps a long stalk that supports

the animal above the substrate. In the feather stars (Comatulida) only one ring is evident, made up of five al

radial ossicles, one at the base of each arm. The radials sit on the oral surface of a large discoidal to cylindrical |

centrodorsal ossicle.

Unbranched cirri articulate with the aboral face of the centrodorsal. The cirri are prehensile, composed Ill

of series of ossicles of which the terminal ossicle is usually modified as a claw which can close against an U

opposing spine on the penultimate cirral. Using its cirri, a feather star can firmly attach to a suitable perch.

The rays are long and flexible, made up of series of small cylindrical ossicles (brachials). The rays I

generally branch. The row of brachials between branching points is called a division series (=brachitaxis), I

and terminates in a modified axillary ossicle (=axil) that either bears two unbranched arms or further division

series. II

Most of the brachials are attached to one another by a ligament below a bony fulcral ridge, and by a

paired muscle masses above the ridge. The ligaments serve to lock the extended arms in position. Arm

movements are powered by the contraction of the muscle masses pulling the arms upward, opposing the

tension created by the ligaments. Some brachials are connected by rigid syzygies or by synarthries. A m

syzygy is a ligamentary articulation in which ridges radiating from the center of one brachial are connected

to corresponding ridges on the adjacent brachial. In dried specimens, it appears as a perforated line between Jail

brachials. A synarthry is a Iigamentary connection between two brachials with an aboral/oral fulcral ridge, II

and is capable of limited side-to-side movement. The brachial proximal to a syzygy or a synarthry lacks a

pinnule. Pinnules are the alternating series of unbranched appendages on the sides of the arms, andthey give I

the arm a feathery appearance. From the base to the tip of the ray there are oral, then genital and finally

distal pinnules. Oral pinnules near the mouth are specialized for defense or for cleaning the oral surface. The

genital pinnules bear the gonads. The majority of the pirmules are food gathering organs. They are grooved, a

and the channel (ambulacrum) is bordered by tube feet. The groove of each pinnule unites with an ambulacral

furrow that runs down each arm to the mouth. Microscopic tube feet, grouped in threes with a protective _l

lappet, alternate on opposite sides of the food groove. m

The tegmen, which encloses the visceral mass, is covered by soft skin. The mouth opens on the

tegmen at the meeting point of the ambulacral grooves, and the anus opens at the apex of a small muscular _l

conical papilla (= anal cone). The mouth is central, except in the Comasteridae, the second largest family of

feather stars. The tegmen is perforated by hundreds of microscopic pores, probably serving in respiration,

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i which lead into ciliated channels in the central body. The visceral mass is filled with connective tissue that

contains numerous microscopic ossicles and surrounds the intestine and axial sinus.

I The body plan of crinoids is described in terms of formulae that specify the composition of division series, arrangement of pinnules, number of cirri, and dimensions of ossicles. The formulae are shorthand

expressions incorporating abbreviations for the ossicles: Br = brachial, P = pinnule, + = syzygy. Roman

i numerals positions ofspecify ossicles. division For example, series, /arabic IBr4(3+4) numerals indicates indicate th senumbers cond division of ossicles, series and with subscripts 4 brachials,indicate with a

syzygy between the 3rd and 4th brachial. Division series are absent in an undivided arm; therefore roman

I numerals Interior pinarenules omitted (thosein aitsrising formula; between thustheBr3arms +4 indicates of a ray)theathird re specified and fourthby numerical brachials joined subscripts, by a exterior syzygy.

pinnules by subscript letters. For example, P is directed medially, and on the next distal brachial P2 is

j directed away from the ray. The number of cirri borne on the centrodorsal is specified as a roman numeral.

i Ecology and Behavior

Crinoids are "passive suspension feeders" that extend their arms into the water to form a feeding

i fan. food.Many The crinoids configuration are rheophilic, of the fanpreferring varies depending areas whereoncthe urrents species flowand betwoneenenvironmental 1-2 knots and conditions, carry abundant the

orientation of the arms and pinnules shifting to increase the effectiveness of particle capture. The pinnules

i tvelocity end to align and vortices perpendicucreated lar tobythetheflowarmsofslow water,andwithdirect theparticles food groove towardfacingthe tube down-current, feet. Rheophobic so that reduced species,

which occupy low-velocity habitats, hold their arms in haphazard arrays, with the pinnules oriented

i perpendicular Driftingto oreachswimming other. prey are rapidly flicked by the long outer tube feet toward the food groove,

where they are trapped in mucus, then moved along the grooves in the pinnules and arms to the mouth.

El Crinoids capture zooplankton (forams and actinopods, invertebrate larvae, and small crustaceans),

I phytoplankton (diatoms and unicellular algae), and particulate matter -- most 0.05-0.40 mm in size.

Feather stars can right themselves if overturned, and can pull themselves slowly along the seafloor.

I Some autotomized swim byandpulling damaged alternating arms and armstheupvisceral and downmass. in a coordinated Crinoids arefashion. subject They to predation, are capablemostly of regenerating by fishes.

They support a diverse assemblage of parasites and commensals including gastropods, ophiuroids, clingfish,

I myzostome polychaetes, and crustaceans (including shrimp, copepods, isopods, and cirripedes). The gonads of crinoids consist of several tissue layers enclosing the gametes, and are borne in

specialized basal pirmules. For most feather stars the breeding season is limited to a 1-2 month period in

i spring or summer. Individuals may spawn en masse, and in synchrony, each female feather star releasing thousands or millions of eggs. Spawning events may last an hour or less, or span a period of several days.

While spawning, many individuals whip their arms, dispersing gametes, but fertilized eggs often stick to the

i arms and pinnules. Some species have the capacity to brood their developing young; their eggs adhere to the genital pinnules or enter a brood pouch from which the larvae emerge.

Within a few days after fertilization a uniformly ciliated embryo develops into a doliolaria larva

I encircled by 3-4 ciliated bands. It lacks a mouth and feeds, if at all, by absorbing dissolved chemicals. The doliolaria swims for several days, before selecting a suitable substrate for settlement where it metamorphoses

into a stalked stage called a "cystidean". Since the posterior end of the cystidean becomes the oral surface of

I the adult, its internal structures must rotate 90° during the transformation to a juvenile "pentacrinoid." At that stage it has the appearance of a miniature stalked crinoid. The pentacrinoid begins feeding when the

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mouth and anus have formed, and its arms later begin to branch. Species with numerous arms develop from _l

a five-armed juvenile by repeatedly growing back two arms in the place of arm branches that have been

shed. After some cirri have developed from the centmdorsal ossicle of the pentacrinoid, the feather star Ill

separates from the top of the stalk and attaches directly to the substratum. !1

Preservation i

Details regarding techniques for collecting, anesthetizing, and preserving crinoids and other I

echinoderms are covered in Messing and Dearborn (1990) and Hendler et aI. (1995). The most efficient II

procedure for preservation involves killing individuals with outspread arms by rapid immersion in 95%

ethanol. Specimens killed in buffered 10% formalin or in alcohol should be stored in 70-80% ethanol. /

Often, the details of arm and cirrus structure are best observed on appendages pried from dried specimens I

using f'me forceps. When examination of individual ossicles is required, the structures can be cleaned using

dilute sodium hypochlorite (laundry bleach). I

Glossary t

Terms that are not deemed above can be found in the extensive glossaries provided in A.H. Clark

(1915), Moore and Teichert (1978), and Messing and Dearborn (1990). I

Terminology and Conventions i

Body plan formulae are explained above, in the section on morphology. Abbreviations used are: Ill

AHF, Allan Hancock Foundation of the University of Southern California; CAMP, California Phase II I

Monitoring Program; LACM, Natural History Museum of Los Angeles County; SBMNH, Santa Barbara

Museum of Natural History; USNM, National Museum of Natural History. m

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I Description of Species

I Order Comatulida

Family Antedonidae

i Subfamily Heliometrinae

I

Florometra serratissima (A.tt. Clark, 1907)

I Figure 5.1

Antedon perplexa A.H. Clark, 1907:70, 74.

I Promachocrinus (Florometra) perplexa, A.H. Clark, 1915:140.

Florometraperplexa, A. H. Clark, 1921: 60, 62, 74,79, 303,670, 685, 725, 729, 758, 760, figs. 95-96, 131,

i 590, 1958:47, pl. 5,58.-- figs 1009-1014, Hartman and pl. 1Barnard, 3.fig.1053.-- 1960:Ziesenhenne, 231,281.-- 1A.H. 937: 209, Clark211.--Hartmart and A.M. Clark,and1967:299, Barnard,

313, 317, 318.

i Antedon serratissima A.H. Clark, 1907:71, 77.

Promachocrinus (Florometra)serratissima, A.H. Clark, 1915:140, 142.

i Florometra serratissima, Mortensen, 1920:54, 55, 66, 94, pl. 27, figs. 1-7.-- A.H. Clark and A.M. Clark, 1967:289, 294, 299-309 [see for additional synonymy], 318, 321,738, 741 .--Breimer, 1978:T36-

37, fig. 22.---Gotshall, 1994:77, fig. 185.-- Byrne and Fontaine, 1981:11-17, figs. 1-6.

I Material Examined. California: Santa Maria Basin, Phase I Sta. BRA-14, 96-I05 (1, USNM

m

primary voucher).--Westem Santa Barbara Channel, Phase I Sta. BRA-002, 110-126 m (1, SBMNH

secondary voucher).----Off San Miguel Id., R/V Velero II1 Sta. AHF 1396-41, 104-113 m (4, LACM 41-

I 176.3).----Off Santa Rosa Id., R/V Velero IllSta. AHF 1392-41,104 m (2, LACM 41-172.6), R/V Velero 1II

Sta. AHF 1395-41, 128-135 m (10, LACM 41-175.1).---Off San Clemente Id., R/V Velero II1 Sta. AHF

1023-39, 101-201 m (1, LACM 39-127.3).--Mexico: Pacific Coast, Baja California, off Islas San Benito,

I RIV VeIero III Sta. AHF 1119-40, 159-174 m (67, LACM 40-92.6).

Description. An antedonid with 5 inconspicuous radials,second post-radial ossicle axillary, 10 arms.

i Centro-dorsal hemispherical, cirri 40-50 or more, stout, large, 30 mm long, with approximately 36 joints exhibiting tendency to overlap; nearly all cirrals bearing strong dorsal spines.

First radials concealed; second barely visible; IBr 1 short, IBr 2 (axillaries) very broad, prominent

i synarthrial equilateral; tubercle third irregularly betweenoblong; IBrl anddistal IBr2; brachials first bratochial the tenth very orshotwelfth rt; second wedge-shaped, triangular, then approtriximately angular.

Syzygies between brachials 3+4, 9+10, 16+17, and distally at intervals of 3 muscular articulations. Radials

I ansmall, d lowesharp r brahyaline chials thickly teeth; set longest with proximal small sharpbrachials spines. Br broader achialsthan short,long. overlapping, edges beating numerous

Arm length of large individuals 150-280 mm. P_ 17-21 mm long with 45-60 joints, with long comb

i _ distally; P2 longer than first, 18-22 mm in length with 45-60 joints; P3 20 mm long with 36 joints, bearing

I comb Iike the second, or 12 mm long with elongated joints like the fourth; distal pinnules long and slender,

their joints overlapping, distal edges set with spines.

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Color: Living individuals yellow, brownish yellow, tan, or reddish brown, the cirri whitish to light I

brown, pinnules may be contrasting dark brown to black. In alcohol pigmentation white, brown, or red

(A.H. Clark, 1907; Ziesenhenne, 1937; Gotshall, 1994).

Variation: Abnormal individuals with 9 or 11 arms have been reported. The extent of spine cover !1

varies considerably, with development greatest in specimens from warm water localities (Clark and Clark,

1967). i Remarks. In Clark and Clark (1967) a review of the genus and key to the seven species is provided.

F. perpIexa, originally regarded by A.H. Clark (1907) as a less robust form with smoother, longer arms, is

placed in synonymy with F. serratissima. In addition, F. serratissima is distinguished from F. mariae (A.H.

Clark), which occurs in Japan, by its relatively short bmchials, fromF, tanneri (Hartlaub), found off Panama, I

by oral pinnules with stout, carinate, proximal segments, and from F. asperrima (A.H. Clark), reported

from Monterey Bay to northern Japan, by the position of its 3rd syzygy between brachials 16+17 (sometimes i

between 15+16) rather than 14+15 and by having more spinose division series and brachiats. i

The status of F. serratissima is unclear, particularly with regard to F. asperrima. In their treatment

of F. asperrima, Clark and Clark (1967:321) state that "the ossicles of the division series and arm bases

never show the extreme development of spines frequent in F. serratissima, but the least spinous individuals ql

of that species are less spiny than the extreme specimens of F. asperrima. From Monterey, California, i

northward to the Gulf of Alaska F. asperrima is more variable than farther northward and westward, and •

apparently intergrades with F. serratissima."

The name perplexa has page priority over serratissima based on the original publication. A.H. m

Clark mentioned that his intitial concept ofAntedon perplexa included what he later regarded as F. tanneri, II

an indication that he was not satisfied with the name perplexa. A.M. Clark concluded that "in view of the

very brief description ofperplexa in the form of a comparison withasperrima, I do not dispute the use of the ms

name serratissima for the species" (Clark and Clark, 1967:309). HI

Distribution. Natividad Island, Baja California, Mexico to Shumagins and Sannak Island, Alaska,

at depths of 11-1252 m (A.H. Clark, 1907; Ziesenhenne, 1937; Clark and Clark, 1967).

Biology. Based on trawled samples it appears that in places "there must be many acres, if not miles, tl

pretty well covered with [Florometra serratissima]" (Clark and Clark, 1967:308). Scuba observations reveal

that population densities reach at least 30 individuals/m 2(Byme and Fontaine, 1981). On soft bottom habitats E

of the southern California shelf F. serratissima is infrequent in occurrence (Bamard and Ziesenhenne, |

1960), but on hard bottom habitats it is characterized as an "anchor species" which provides a biogenic

refuge for other fauna (Lissner and Benech, I993). At the southern end of its range it has been collected in a

association with Fariometra parvula (Hartlaub), the young of which it resembles (Clark and Clark, 1967). |

Byrne and Fontaine (1981) found that F. serratissima occurs where currents are of low velocity,

and described how its arms are held in a cone array during slack water, and form a partial fan when flow

increases. They studied the feeding process and activities of the triplet groups of podia, noting that each w

tentacle functions differently in sensing, collecting, and transferring paticles to the food groove. LaBarbera

(1982) characterized metabolic rates of individuals in normal current regimes, finding lower values than

previously estimated. II

The sea star Pycnopodia helianthoides (Brandt) and a crab Oregonia gracilis (Dana) may prey on

F. serratissima (MIadenov, 1983). Its escape behaviors include autotomy and swimming, and the swimming

response has been described (Mortensen, 1920; Mladenov, 1983). Breimer (1978) noted that the species has U

"external obliqueness of radials accompanied by reversion," that is, the pinnules shifting from the low side

of proximal brachials to the high side of distal brachials, a characteristic of swimming comatulids. In the

field, 80% of the individuals encountered have regenerating arms, probably having escaped from predators; U

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arm regeneration takes approximately 9 months (Mladenov, 1983). An external myzostome parasite, N

Myzostomum deani McClendon is associated with individuals off Washington and northern California

(McClendon, 1906; A.H. Clark, 1921).

The reproductive biology and development of F. serratissima are described by Mladenov and Chia

(1983) and Mladenov (1986), and discussed in a comparative context by Holland (1991) and Nichols (1994).

The sexes are separate and the sex ratio of individuals in a population is approximately 1:1. Large specimens B

(arm length 280 mm) have over 400 genital pirmules with a total of approximately 24,000 eggs. The available n

data have been interpreted to indicate that reproduction occurs year round, with individuals spawning

approximately monthly and the level of activity varying with the season. The gonad structure and process of I

gametogenesis are clarified by several authors (Bickel et al., 1980; Mladenov, 1986; Holland; 1991).

The eggs are pale pink, 0.21 mm in diameter, and produce a fertilization membrane with a ridged

ornamentation. The doliolaria larva is yolky and barrel-shaped, with 4 ciliated bands; formation of the

bands and of the larval integument have been described by Chia et al. (1986), Lacalli and West (1986, n

1987), and Lacalli (1993). The larvae swim up to 9 days before settlement, rapidly metamorphose to a

sessile cystidean, and gradually transform to a feeding, stalked pentacrinoid. The feeding response of the

pentacrinoid has been studied (Mladenov and Chia, 1983) and the skeletal ontogeny of the pentacrinoid has J

been depicted (Mortensen, 1920). Between the egg and the prefeeding cystidean stage, 81% of the nutrient

in the egg is depleted. As a result, the pentacrinoid has relatively loe energy reserves, possibly making it a

vulnerable stage in the crinoid's life history (McEdward et al., 1988).

Mitochondrial DNA has been isolated from F. serrafissima, but DNA amplification products have

not successfully been obtained using echinoid or asteroid primers (Smith et al., 1993).

n

Acknowledgements I

This study was funded by the Pacific Outer Continental Shelf Region of the Minerals Management II

Service, U. S. Department of the Interior, Washington, DC. J. Blake and A. Lissner offered the MMS I

collection for study and arranged for its deposition at LACMNH. C. Messing verified specimen identifications.

J. Dearborn and D. Pawson reviewed a draft of the manuscript. I am grateful to all those acknowledged, for m

their assistance with this chapter. I

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I Literature Cited

i Bamard, Naturalist J.L. and F.C. 2:131-152. Ziesenhenne. Ophiuroid communities of Southern California coastal bottoms. Pacific

i Bickell, L.R., F.-S. Chia, and B.J. Crawford. 1980. Fine structural study of the testicular wall and spermatogenesis in the crinoid, Florometra serratissima (Echinodermata). Journal of Morphology

166:109-126.

I BirkelandLawrence , C. 1989. The influence of echinoderms on coral-reef communities. In: M. Jangoux and J.M. (eds.), Echinoderm Studies, Volume 3. Pp. 1-79. A.A. Balkema, Rotterdam.

i Breimer, A. 1978. General morphology: Recent crinoids. In: R.C. Moore and C. Teichert (eds.), Treatise on

Invertebrate Paleontology. Part T. Echinodermata 2, Volume 1. Pp. T9-T58. Geological Society of

America and University of Kansas Press, Lawrence, Kansas.

i Byrne, M. Crinoidea). and A.R.Canadian Fontaine. 1981. The feeding behaviour of Florometra serratissima (Echinodermata: Journal of Zoology 59:11-18.

i Carpenter, P.H. 1884. Report on the Crinoidea collected during the voyage of H.M.S. Challenger, during the years 1873-1876. Part I. General morphology, with descriptions of the stalked crinoids. Report

on the Scientific Results of the Voyage of H.M.S. Challenger during the Years 1873-76. Zoology,

Volume 11 (Part 32):xii, 1-442, pls. 1-62.

i Carpenter, P.H. 1884. Report on the Crinoidea collected during the voyage of H.M.S. Challenger, during

the years 1873-1876. Part II. The Comatulae. Report on the Scientific Results of the Voyage of

I H.M.S. Challenger during the Years 1873-76. Zoology, Volume 26 (Part 60):ix, 1-399, pls. 1-70.

Chia, F.-S., R.D. Burke, R. Koss, P.V. Mladenov, and S.S. Rum_rill. 1986. Fine structure of the doliolaria

larva of the feather starFlorornetra serratissima (Echinodermata: Crinoidea), with special emphasis

I on nervous system. Morphology

the Journal of 189:99-120.

Clark, A. H. 1907. Descriptions of new species of Recent unstalked crinoids from the North Pacific Ocean.

I Proceedings of the United States National Museum 33:69-84.

Clark, A. H. 1915. Die Crinoiden der Antarktis. Deutsche Siidpolar-Expedition 1901-1903. Volume 16,

Zoologie 8:101-209, pls. 2-10.

I Clark, A.H. 1921. A monograph of the existing crinoids. Volume 1,Part 2. United States National Museum

Bulletin 82:xxv, 1-795, 57 pls.

I Clark, A.H. and A.M. Clark (1967) A of the existing crinoids. Volume 1, The comatulids, Part

monograph

5 - Suborders Oligophreata (concluded) and Macrophreata. United States National Museum Bulletin

82:x/v, 1-860.

I Gotshall, D. W. 1994. Guide to Marine Invertebrates: Alaska to Baja California. Sea Challengers, Monterey,

California.

I Harflaub, C. 1895. Reports on the dredging operations offthe West Coast of Central America to the Galapagos,

to the West Coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried

on by the U.S. Fish Commission Steamer "Albatross," during 1891, Lieut.-Commander Z.L. Tanner,

i U.S.N., Commanding. Die Comatuliden. Bulletin of the Museum of Comparative Zo61ogy, at Harvard College 27:129-148, 4 pls.

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Hartman, O. and J. L. Barnard. 1958. The benthic fauna of the deep basins off Southern California. Allan i

Hancock Pacific Expeditions 22:1-67.

Hartman, O. and J. L. Barnard. 1960. The benthic fauna of the deep basins off Southern California. Part II. !

Allan Hancock Pacific Expeditions 22:1-297.

Hendler, G.,J. E. Miller, D. L. Pawson, and P. M. Kier. 1995. Sea Stars, Sea Urchins, and Allies: Echinoderms

of Florida and the Caribbean. Smithsonian Institution Press, Washington, DC. n

Holland, N.D. 1991. Echinodermata: Crinoidea. In: A.C. Giese, J. S. Pearse, and V.B. Pearse (eds.), =m

Reproduction of Marine Invertebrates. Volume 6. Echinoderms and Lophophorates. Pp. 247-299. •

Boxwood Press, Pacific Grove, California. ,m

Jangoux, M. and J.M. Lawrence (eds.). 1982. Echinoderm Nutrition. A.A. Balkema, Rotterdam. I

It

LaBarbera, M. 1982. Metabolic rates of suspension feeding crinoids and ophiuroids (Echinodermata) in a

unidirectional laminar flow. Comparative Biochemistry and Physiology, Part A: Comparative toNi

Physiology and Pharmacology 71A:303-307. mw

Lacalli, T.C. 1993. Ciliary bands in echinoderm larvae: Evidence for structural homologies and a common

plan. Acta Zoologica (Stockholm) 74:127-133. i

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Lacalli, T.C. and J.E. West (1986) Ciliary band formation in the doliolaria larva of Florometra. I. The

development of normal epithelial pattern. Journal of Embryology and Experimental Morphology

96:303-323. N

Lacalli, T.C. and J.E. West (1987) Ciliary band formation in the doliolaria larva of Florometra. II.

Development of anterior and posterior half-embryos and the role of the mesentoderm. Development I

99:273-284. II

Lane, N.G. 1978. Historical review of classification of Crinoidea. In: R.C. Moore and C. Teichert (eds.),

Treatise on Invertebrate Paleontology. Part T. Echinodermata 2, Volume 1. Pp. T348-T359. i

Geological Society of America and University of Kansas Press, Lawrence, Kansas.

Lissner, A. L. and S. W. Benech. 1993. Benthic hard-substrate community ecology of the Santa Maria Basin m

and Western Santa Barbara Channel. In: J. A. Blake and A. L. Lissner (eds.), Taxonomic Atlas of II

the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Volume I.

Introduction, Benthic Ecology, Oceanography, Platyhelminthes, and Nemertea. Pp. 47-73. Santa mR

Barbara Museum of Natural History, Santa Barbara, California. II

Maluf, L.Y. 1988. Composition and distribution of the Central Eastern Pacific Echinoderms. Natural History

Museum of Los Angeles County Technical Reports No. 2:1-242. t

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McClendon, J.F. 1906. The myzostomes of the 'Albatross' Expedition to Japan. Bulletin of the American

Museum of Natural History 22:119-129, pls. 15-18. m

McEdward, l L.R., S.F. Carson, and F.-S. Chin. 1988. Energetic content of eggs, larvae, and juveniles of

Florometra serratissima and the implications for the evolution of crinoid life histories. International

Journal of Invertebrate Reproduction and Development 13:9-22. I

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Messing, C.G. and J.H. Dearborn. 1990. Marine fauna and flora of the Northeastern United States -

Echinodermata:Crinoidea. NOAA Technical Report NMFS 91 :ii, 1-29 III

Meyer, D.L. and W.I. Ausich. 1983. Biotic interactions among Recent and among fossil crinoids. In: M.J.S. m

Tevesz and P.L. McCall (eds.), Biotic Interactions in Recent and Fossil Benthic Communities. Pp.

377-427. Plenum Publishing Company, New York. I

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I Mladenov, P.V. 1983. Rate of arm regeneration andpotential causes of arm loss in the feather starFlorometra

serratissima (Echinodermata: Crinoidea). Canadian Journal of Zoology 61:2873-2879.

I Mladenov, P.V. 1986. Reproductive biology of the feather star Florometra serratissima: Gonadal structure,

breeding pattern, and periodicity of ovulation. Canadian Journal of Zoology 64:1642-1651.

I Mladenov, P.V. and Chia, F.S. 1983. Development, settling behavior, metamorphosis and pentacrinoid feeding and growth of the feather star Florometra serratissima. Marine Biology 73:309-323.

Moore, R.C. and C. Teichert (eds.). 1978. Treatise on Invertebrate Paleontology. Part T. Echinodermata 2,

I 3 vols. Geological Society of America and University of Kansas Press, Lawrence, Kansas.

Mortensen, T. 1920. Studies in the development of crinoids. Carnegie Institution of Washington Publication

294, Papers from the Department of Marine Biology of the Carnegie Institution of Washington

I 16:v, 1-94, 28 pls.

Nichols, D. 1994. Reproductive seasonality in the comatulid crinoid Antedon bifida (Pennant) from the

i English Channel. Philosophical Transactions of the Royal Society of London B343:113-134.

Rasmussen, H.W. 1978. Articulata. In: R.C. Moore and C. Teichert (eds.), Treatise on Invertebrate

i Paleontology. Part T. Echinodermata 2, Volume 3. Pp. T813-T948. Geological Society of America and University of Kansas Press, Lawrence, Kansas.

Simms, M.J. 1988. The phylogeny of post-Palaeozoic crinoids. In: C.R.C. Paul and A.B. Smith (eds.),

I Echinoderm Phylogeny and Evolutionary Biology. Pp. 269-284. Clarendon Press, Oxford.

Smith, M.J., A. Arndt, S. Gorski, and E. Fajber. 1993. The phylogeny of echinoderm classes based on

i mitochondrial gene arrangements. Journal of Molecular Evolution 36:545-554. Ubaghs, G. 1978. Classification of the echinoderms. In: R.C. Moore and C. Teichert (eds.), Treatise on

Invertebrate Paleontology. Part T. Echinodermata 2, Volume 1. Pp. T359-T371. Geological Society

I of America and University of Kansas Press, Lawrence, Kansas.

Ziesenhenne, F. C. 1937. The Templeton Crocker Expedition. X. Echinoderms from the West Coast of

i Lower California, the Gulf of California and Clarion Island. Zoologica 22:209-239.

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I 6. CLASSASTEROIDEA

by

I Andrew Lissner 1and Debra Hart 2

I Introduction

I Asteroids, commonly known as seastars or starfish, are a class of flattened, multli-armed echinoderms

I containing approximately 2,000 living species, with the highest number of species occurring in the Northeast

_l Pacific. General references utilized for this section are Hyman (1955), Barnes (1980), and Feder (1980).

II Most seastars possess five arms arranged around and grading into a central disc. However, species with 4 to

24 arms are also common. Additionally, one species (Heliaster) from the eastern Pacific can have 40 or

_ more arms. The size of most asteroids ranges from 12 to 24 cm in diameter, although Pycnopodia from the

I northwest coast of the United States may reach a diameter of almost one meter.

Typically, the seastar body is dorsoventrally flattened with the oral surface and tube feet (podia)

i positioned Furrows against the substratum, and the anus and madreporite on the exposed upper (aboral) surface. (ambulacral grooves) containing the podia extend radially from the mouth into each arm. Podia

function primarily as locomotory organs but in many species also are used to attach to hard surfaces, to

I cap disksturethatandcreate hold suction prey, orortothrough burrow secretion into sediments. of adhesive Attachment mucus. of podia is by means of terminal adhesive

The seastar endoskeleton is composed of calcareous ossicles embedded in a matrix of connective

I tissues. Each ossicle is composed of a single crystal of magnesium-rich calcite formed inside a dermal cell. The surface of an asteroid is covered by an epidermis of monociliated columnar epithelium. Sensory nerve

cells and mucous gland cells also lie in the epidermis, providing the outer surface with a protective mucous

I layer.

Diet and feeding methods of seastars can vary considerably. Most species are carnivorous, feeding

primarily on molluscs, crustaceans, and other echinoderms. A smaller number are omnivorous scavengers.

i For other, suspension-feeding species, smaller food items such as plankton and organic particulates in the

water column may be trapped by mucus and cilia and moved toward the oral surface for digestion. Seastars

may also use their arms to entrap smaller prey, pulling them toward the everted stomach, where digestion

I begins. Some species such as Pisaster can evert their stomachs into openings in mollusc shells and begin digesting prey tissue externally. Other species, including Luidia andAstropecten, swallow their prey whole,

as generally limited by the size of the mouth opening.

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t Science Applications International Corporation, 10260 Campus Point Drive, San Diego, CA, 92121

i 2Science Applications International Corporation, 1710 Goodridge Drive, McLean, VA, 22102
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External fertilization is characteristic of most seastars; gametes are shed into the sea by both sexes, n

Alternately, some Northeast Pacific species such as Henricia, Pteraster, and Leptasterias lay and brood

large yolky eggs that develop directly into juvenile seastars. Growth and development axe dependent on

water temperature and food supply (cooler temperatures and limited food are directly correlated with slower II

growth). Under natural conditions, growth tends to be slow and life spans are relatively long (e.g., 5-years

for Astropecten and at least 20 years for Pisaster ochraceous). Adult seastars have few natural predators, lm

likely contributing to their longer life spans. l

Systematists primarily classify asteroids according to skeletal characteristics, but soft parts such as

podia and internal organs may also be used. Seastar size may be used for some taxonomic descriptions, such

as the distance between the center of the mouth (or disc) to the tip of the longest arm, and the arm radius.

Taxonomic History I

The distinctive morphology, color, andrelatively large size of most California asteroids have made l

them the subject of numerous scholarly and popular presentations on their taxonomy and distinguishing II

characteristics. Important taxonomic monographs on Pacific Coast Asteroidea include Fisher (191 I, 1928,

1930) and Verritl (1914), with a seminal treatise on their worldwide phylogeny presented in Fell (1963). I

Subsequent papers and reports that provide excellent synopses for many southern California species include U

Feder (1980), Hopkins andCrozier (1966), and Word and Charwat (1975). A classic reference thatsummarizes

key distinctions among Orders and Families of asteroids, as well as considerable species-specific information i

on West Coast taxa is Hyman (1955). A particularly useful, popular reference, including color plates of n

shallow subtidal species, is Gotshall and Laurent (1979). Other relevant summaries of species distributions

and their natural history are presented in Ziesenhenne (1941), Hartman (1955), and Hartman and Bamard II

(1958), Pequegnat (1964), and Lissner and Dorsey (1986). Important synopses of taxonomic collections of I

asteroids, including those from southern California, are provided in Luke (1982), Muscat (1980), and Maluf

(1988). I

Morphology I

Key features that are useful in identifying asteroids include the number and relative length of the

arms compared to the central disc, type and relative size of arm and mouth plates, presence/absence and l

type of pedicellariae and paxillae, number and type of papulae, and, in fresh specimens, body color. il

Specifically, important characters that must be recognizable to use the taxonomic key inthis section include

supramarginal, inframarginal, and ambulacral plates; shape and number of spines on paxillae; type of Ii

pedicellariae; and adambulacral (furrow) spines (Figure 6.1 and General Terms below). II

The taxonomic key for the asteroids was developed in part, and the identifications were made, with

the aid of the monographs of Fisher (1911, 1928, 1930). Many specimens also were compared with the _l

asteroid collection of the Allan Hancock Foundation, University of Southern California. u

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enlarged central paxillae

I

supramarginal plates

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i enlarged view of aboral surface

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inframarginal region

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I enlarged view of ambulacrum

I

F_uLre 6.]1. Common external features of many seastars (Astropecten is depicted). Source: Word and Chazwat,

i 1975.

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Glossary I

Adambulaeral plates. Calcareous plates Pedicellaria. Small, stalked to sessile organs, i

comprising the ambulacrum and located typically on the aboral surface of most sea tl

ventrally along the arm. stars (except most Spinulosida), comprised

of jaw-like valves functioning in defense, B

Adambulaeral spines. Calcareous spines located cleaning, and scavenging. i

within the ambulacrum.

Supra-ambulaeral plates. Calcareous plates

Ambulaerum. Central furrow, containing the located on the dorsal-extending top of the I 

podia, that extends ventrally along the ambulacrum, particularly in the Order

I

length of each arm. Paxillosida.

Inframarginal plates. Calcareous plates located Supramarginal plates. Calcareous plates located i

along the ventral margin of the arm; along the dorsal margin of the arm,

particularly prominent in Orders such as the particularly prominent in the Order I

Paxillosida. Paxillosida. II

Paxillae. Small, peg-like, calcareous plates set in

the aborat epidermis of most sea stars, 1

particularly the Paxillosida, Valvatida, and II

Spinulosida.

i:

Methods of Laboratory Examination i

fill[

Identification of asteroids was accomplished using the unaidedeye to distinguish gross morphological

features such as color, number of arms, and general shape. A hand lens or stereomicroscope was used to l

examine surface features such as paxillae and pedicellariae. Some dissections, aided by the microscope, If

were performed to verify in cross-section some of the arm plate features, as indicated in the taxonomic key.

However, most of the species are very distinct based on external features alone. Specifically, the taxonomic i

orders can be recognized based on external features such as the size of marginal arm plates, presence and It

shape of pedicellaria, and presence or absence of suckers on tube feet. Once the order is determined, the

type of habitat (soft versus hard substrate), color (in fresh specimens), type of pedicellaria, skin thickness, I

presence/absence of spinose plates on aboral surface, and presence/absence of a nidamental (brood) chamber il

(determined by dissection) typically are sufficient to distinguish species from the study area.

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I List of Species

Class Asteroidea Order Spinulosida

Order Paxillosida Family Pterasteridae

I FamilyLuidiaLuidiidae foliolata Grube, 1866 FamilPteraster y Poraniidae tesselatus arctuatus (Ives, 1888)

Family Astropectinidae Poraniopsis inflata (Fisher, 1906)

I Astropecten verriUi (Fisher, 1906) Family Echinasteridae

Henricia leviuscula annectens Fisher, 1910

I OrderFamily ValvaGoniasteridae tida Order Forcipulatida

Hippasteria spinosa Verrill, 1909 Family Zoroasteridae

I Myxoderma pIatyacanthum H.L. Clark, 1913

Family Asteriidae

i Rathbunaster californicus Fisher, 1906

I Key to Orders and Species of Asteroidea Collected from the Santa Maria

Basin and Western Santa Barbara Channel

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1A.Marginal plates (supramarginals and inframarginals) large and conspicuous ....................................... 3

I lB. Marginal plates not large and conspicuous ............................................................................................ 2

2A. Pedicellaria absent or rare, not stalked ............................................................. Order Spinulosida ...... 5

I 2B. Pedicellaria common, stalked ........................................................................... Order Forcipulatida ... 7

3A. Supmmarginal plates smaller than inframarginals and similar in size to paxillae ..................................

II .................................................................................................... Order Paxillosida, Luidiafoliolata

3B. Supramarginal and inframarginal plates similar sized and larger than paxiltae ................................... 4

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4A. Suckers present on tube feet; at least one hyaline or unpaired median tooth common to each pair of mouth

i 4B. Suckers plates, absent or supmon -tube ambufeet; lacralhyaline platesorreduced medianorteeth absentabsent ................ from mouth Order plates; Valvafida, supra-ambula Hippasteriacralspinosa plates3

present, not reduced .............................................................. Order Paxillosida, Astropecten verrilli

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i 3Hippasteria spinosa is the only orange-colored (in fresh specimens), five-armed species with extensive aboral spines

that occurs in the study area. Other orange to brownish-orange, five-armed taxa can include Mediaster aequalis

and Poraniopsis inflata; however, Mediaster has no aboral spines and Poraniopsis has only a few, widely

i spaced spines that typically are white in color.

I 101

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5A. Mouth plates small, not spade-shaped ........................................................ Family Echinasteridae ...... 6 J

5B. Mouth plates large and spade-shaped .................. Family Pterasteridae, Pteraster tesselatus arctuatus 4

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6A. Aboral plates covered with thick skin ...................................................................... Poraniopsis inflata

6B. Aboral plates with small spinelets, not covered with thick skin ......................Henricia leviuscula annectens 5 I

7A. Straight and crossed pedicellaria present; no furrow spine .....................................................................

................................................................................... Family Asteriidae, Rathbunaster californicus 1

III

7B. Only straight pedicellafia present; prominent furrow spine ....................................................................

............................................................................ Family Zorasteridae, Myxoderma platyacanthum 6 I

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Descriptions of Species I

II

OrderPaxillosida

Family Luidiidae I

Luidiafoliolata Grube, 1866

Figure 6.2 I

Luidiafoliolata: Fisher, 1911. 1

Petalasterfoliolata: Fell, 1963. II

Material Examined. California: Santa Mafia Basin, Sta. BRA-26 (USNM), 108-111 m (one Ill

specimen, 150 mm arm length); Sta. BRA-22 (SBMNH), 114-115.5 m (one specimen, 130 mm arm length). I

Description. Large, long-armed, typically five-armed seastar with dull gray or brown color in fresh

specimens (Lambert, 1945). Prominent supramarginal plates absent in the genus (representing a clear I

distinguishing feature from Astropecten, which is the only other common, five-armed seastar co-occurring II

in most soft substrate areas off California; Figures 6.2 and 6.3). Pedicellariae also absent (Lissner, 1980).

Marginal arm spines distinct (Figure 6.2). Arms rectangular in cross-section; podia pointed, without suckers I

il

II

4Pteraster tessefatus arctuatus, representing a "pin cusMon" slar of the Family Pterasteridae, is the only species in the _ll

study area that has a nidamental (brood) chamber; very short and thick arms; and a fleshy, light brown to tan (in

fresh specimens) epidermis. in

5Henricia leviuscuta annectens is distinguished from 1-1.teviuscula by having pseudopaxillae with fewer than 20 spinelets, U

while the latter species has 20 or more spinelets. Henricia aspera is distinguished by having papular areas that

are larger than the pseudopaxillae, while in H. teviuscuta the papular areas are smaller than the pseudopaxillae. II

Henricia pofyacantha is distinguished from all these species by having flexible as compared to rigid arms. | Additionally, from fresh specimens, H. teviuscula is bright orange to, rarely, purple in color while the other

species are white.

Myxodermaptatyacanthum and M. saccutatum, representing the two local congeners, are both light tan in color (in fresh !

6

specimens) with distinctive, reticulated patterns associated with papnlae on the aboral surface. A distinguishing m

feature is that the terminal arm plate is enlarged (ovoid) in M. saccutatum but is tapering (not ovoid) in M.

platyacanthum. _l

102 I
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I (Lambert, 1945). Arms autotomize easily when an individual is handled or disturbed, and specimens often

fragment during preservation. An important distinguishing feature betweenL foliata and the sometimes col occurring a darker field", L. armata as compared is that theto afomore rmer mosaic consistently patternhasof"different only scattered, colors ligh in thet-clatter olored(D.individ Cadienualinpaxillae SCAMIT, on

1994).

I burrow Bi intoology. substratum The species to capture is foundpreyon soft (Lawrence, substrates1987). (shell,Luidia mud, and foIiolata sand) and commonly individuals is observed are knownintoa

"humped" feeding posture, requiring flexion of the rays, and is an active predator on soft substrate organisms

I including entire, intact molluscs, heart urchinsrustaceans, have beenadnoted otherwithin echinoderms. the bodyFeeding cavity ofisthisby species ingestion(Lissner, of the pers. prey obs.). item, The and

maximum bivalve that can be swallowed typically cannot exceed the mouth diameter (e.g., 40 mm; Sloan

I and probably Robinson, i an indication 1983). Theof escape the voracious responsenature elicitedof this fromseastar the sea(Margolin, cucumber,1976). Parastichopus californicus,

Remarks. Fell (1963) redefined this species into the Genus Petalaster, but most researchers (e.g.,

I Maluf, 1988; SCAMIT, 1994) recognize the original description of Luidia, as followed for this atlas. Distribution. The species ranges from southeast Alaska to the Galapagos Islands, at depths from 0-

476 m (Maluf, 1988).

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Figure 6.2. Aboral view of arm from Luidia foliolata (Station BRA-22), emphasizing lack of prominent

i supramarginalplates, but distinct arm spines.

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Family Astropectinidae I

Astropeeten verrilli (Fisher, 1906) i

Figure 6.3

Astropecten californicus Fisher, 1906. I

Astropecten verrilli: de Loriol, 1917.

Material Examined. California: Santa Mafia Basin, Sta. BRA-01 (USNM), 69-73.5 m (one I

specimen, 80 mm ann length); Sta. BSR-21 (SBMNH), 29 m (one specimen, 85 mm arm length); Sta. R-3,

Rep. 3 (SBMNH), 409 m (one specimen, 29 mm arm length).

Description. This species typically five-armed, with light tan to grayish color in fresh specimens. I

Aboral surface covered with small paxillae, of which 4-5 equal in length to 2 supramarginal plates ((Figure

6.3; Lissner, 1980). Prominent, spine-beating, supramarginal plates present on upper edges of arms (Figure

6.3; Feder, 1980). Podia occur in double rows in ambulacra, and lack suckers, consistent with occurrence of I

this species on soft substrates. Distinctions between juvenile A. verrilli and co-occurring A. armatus can be

made based on the "relatively longer and more robust lateral spines, and the smaller arm length to disc

diameter ratio" for the latter species (D. Cadien in SCAMIT, 1994). I

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Figure 6.3. Aboral view of arm from Astropecten verriUi (Station BRA-01), showing prominent, spinous, i

supramarginal plates andrelatively small paxillae.

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! Biology. The species is found subtidally on soft substrates such as sand and mud (Feder, 1980).

Species in the genus are active predators on a wide variety of organisms, including snails, clams, crustaceans,

I other into theechinoderms, soft substrateandaround dead fish. a prey Feeding item byandthese thenseastars swallowing is typically the organism accomplished whole;bytheforcing stomach theirisarms not

everted.

I Pedro, California, Distribution.ranging Astropecten from Pt. Arena verrillisouth is thetoonly Central Astropecten America atspecies depthsknown from approximately to occur north2-488 of Sanm

(Maluf, 1988).

I Order Valvatida

I Family Goniasteridae

Hippasteria spinosa Verill, 1909

I Figures 6.4A and B

I Material Examined. California: Santa Maria Basin, Sta. BRA-20 (USNM), 90-130.5 m (one specimen, 70 mm arm length).

Description. Typically five-armed somewhat stout, with orange to vermillion body in fresh specimens

I (-Lambert, 1945), fading to off white upon preservation. Aboral surface with extensive, thin, sharply-pointed spines, and double row of thicker spines on prominent marginal plates (Figure 6.4A; Lissner, 1980). Oral

surface lacks long spines (Figure 6.4B). All plates surrounded by granules producing stellate appearance

I (Lambert, 1945). Paxillae on oral surface circular to oval in pattern, commonly with central, sessile, clam- like pedicellaria.

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I Figure 6.4. A, on Aboral surfaceview and margina of arm fi'om l spines. Hippas B, Oral teriaview spinoofsaarm(Stafrom tionBRHippas A-20),teriaindicating spinosaexte (StantisionveBRA-20), spination

depictinglack of prominent surface spines and paxillae surrounding clam-like pedicellariae.

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1

Biology. Hippasteria spinosa is typically found on sandand mud, and occasionally rocks (Fisher, n

1930). The species is an active predator that is capable of excavating soft substrates up to at least 7.5 cm

deep to capture prey (Mauzey et al., 1968). In Puget Sound the species is known to feed primarily on the I

seapen, Ptilosarcus gurneyi, which it consumes by first everting itsstomach over the apical tip and progressing I

down the length (Mauzey et al., 1968). Less common prey items include polychaetes and anemones.

Distribution. This species occurs from Alaska (Kodiak) to southern California (Fisher, 1930), _l

ranging from at least 49-1170 m depth (Maluf, 1988: SAIC and MLML, 1992). m

Order Spinulosida I

Family Pterasteridae l

Pteraster tesselatus arcuatus Oves, 1888) •

Figures 6.5A and B I

_w

Material Examined. California: Santa Maria Basin, Sta. BRA-21 (USNM), 75-90 m (one specimen,

60 mm arm length). I

J

Deseription. This species a five-armed seastar with stubby arms and central, aboral osculum (Figure

6.5A). Color in fresh specimens usually tan, but ranges from cream to yellow to dull gray-purple, with

orange or yellow podia (Lambert, 1945). Pedicellariae absent (Lawrence, 1987), with spines separated by i

membranous fans (Lambert, 1945). Unique nidamental (brood) chamber present (Lissner, 1980). Oral surface w

with spinous ambulacral grooves and prominent web-like mouth plates (Figure 6.5B).

Biology. The species (as P. tessalatus) typically inhabits broken to solid rock substrate (Mauzey et B

aI., 1968), but also is noted from sand and mud (Maluf, 1988). Feeding is accomplished by everting the

stomach over sessile prey such as bryozoans, hydroids, and tunicates, although sponges may be the preferred all

diet (Mauzey et al., 1968). Individuals are capable of producing copious amounts of mucus when handled or II

disturbed. This mucus is toxic to some snails, hermit crabs, and sea cucumbers. The presence of a supradorsal

I

A B

1

Figure 6,5. A, Aboral view of Pteraster tesselatusarcuatus (StationBRA-21), emphasizing stubby body form,

smooth surface, and central osculum. B, Oral view of Pteraster tesselatusarcuatus (Station BRA21), showing spinous ambulacral grooves and Prominentmouthplates. 1

II

106 I
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I membrane capable of expansion and contraction serves to regulate water flow and thus oxygen uptake

(Lambert, 1945). The species is unique in the asteroids in that there is an absence of typical larval body plan

i features, be selectionsuggesting for increased directdevelopment development efficiency from embryos in brooded to juveoffspring niles (McEdwards, such as these. 1994). There appears to

Distribution. The species (as P. tesselatus) is found from the Bering Sea (Fisher, 1930) to at least

I Santa 1988).Monica Bay (Catalina Island), and broadly over the northeast Pacific at depths from 0-435 m (Maluf,

I Family Poraniidae

I Poraniopsis inflata (Fisher, 1906)

Figures 6.6A and B

i Material Examined. California: Santa Maria Basin, Sta. BRA-21 (USNM), 75-90, (one specimen,

60 mm arm length); Sta. BRA-20 (SBMNH), 90-130.5 m (one specimen, 14 mm arm length).

I The bodyDesccolor ription. orangePoranio to yellow psis iornflata pinkis(Anderson a five-armedand sShimek, eastar with1993), somewith whatwhite, fa arms sparse, (Figure but eve 6.6A). nly

distributed spines in fresh specimens (Lambert, 1945). Aboral plates covered with thick skin (Lissner,

I 3-4 1980); perpedicellari row; Figure ae sessile 6.6B). (Hyman, 1955). Podia within ambulacra are numerous and small (approximately

Biology. Preferred substrates are sand, mud, and rock (Maluf, 1988). Feeding habits axe incompletely

I know suggested n, butascaptive likely speci prey minclude ens seeme molluscs, d to precup fer spo corals, ngesand(Ansea derson anemones, and Shimek, potentially 1993).asOther verifiedorganisms by the

escape response that this seastar elicits in the anemone Stomphia didemon. Poraniopsis inflata has a large

I cardiac stomach, implying a likely capacity for extrusion (Lambert, 1945).

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i Figure 6.6. A,Aboral view of Poraniopsisinflata (StationBRA-21), indicating body form andrelatively sparse

and evenly distributed surface spines. B, Oral view of Poraniopsis inflata (Station BRA-21),

i emphasizingnumerous,small podiaper ambulacralrow.

I 107
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Distribution. The species ranges from British Columbia (Hopkins and Crozier, 1966) to South n

America (Maluf, 1988) at depths from 48-1094 m. A subspecies, P. inflataflexilis, is found in deeper water

(600-1080 m) within the depth range (Fisher, 1930). 1

Family Echinasteridae I

Henricia leviuscula anneetens (Fisher, 1910)

Figure 6.7 I

Material Examined. California: Santa Maria Basin, Sta. BRA-01 (USNM), 69-73.5 m (one m

specimen, 40 mm arm length); Sta. BRA-14 (SBMNH), 96-105 m (one specimen, 40 mm arm length), l

m

Description. Henricia leviuscula exhibits wide variety of size, color, and spine shape, leading to

recognition by Fisher (1911) of several subspecies (Feder, 1980), including H. Ieviuscula annectens. This m

subspecies subtidal only, colored white to off-white in fresh specimens, as compared to more familiar, II

orange to purplish H. leviuscula that common in many intertidal and subtidal areas of California. Species

typically with five arms, but four- or six-armed specimens sometimes found. General body form comprises I

small central disc and proportionately long, sharply tapered arms (Figure 6.7). The aboral surface appears U

superficially smooth and without prominent spines (Sutton, 1975), although on close inspection (hand lens)

pseudopaxillae bearing less than 20 spinelets extensive over the surface, forming a "corral" around papulae I

(Lissner, 1980). These papulae produce a small-scale,'_ck-marked" appearancein preserved specimens (Figure II

6.7). Three rows of plates present along lower side of each arm (Lambert, 1945). Pedicellariae absent (Sutton,

1975). I

I

Figure 6.7. Aboralview of armfromHenricialeviuscula annectens (StationBRA-01),depicting"pock-marked" I

appearanceofpapulaesurroundedbypseudopaxillae.

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I Biology. Little is known about the biology of this subspecies, although the body form and occurrence

in hard substrate areas suggests similar habits as H. leviuscula. Henricia leviuscula typically occurs under

I or on the protected sides of rocks (Morris et al., 1980). Typical substrate types are sand, mud, rock, shale, and shelly debris (Maluf, 1988). Two methods of feeding are observed: suspension feeding and eversion of

the stomach for direct feeding on sessile organisms, including bryozoans and sponges. The size of the

i everted stomach Distribution. is limited The tspecies o the size(asofHt.heleviuscula individual's ) ranges mouthfrom(Jangoux, the Aleutian 1982).Islands (Alaska) to lower

Baja California, at depths from the intertidal to 400 m (Maluf, 1988).

I Order Foripulatida

I Family Zoroasteridae

Myxoderma platyacanthum H.L Clark, 1913

I Figure 6.8

I mm armMalength); terial Sta. Exami 100ned(SBMNH), . Californi443 a: Santa m (three Mariaspecimen Basin,s,Sta. 25,09227,(USNM), and 29 mm444armm (one lengths); specimen, Sta. R-7, 31

Rep. 2 (SBMNH), 565 m (one fragmented specimen, approximately 32 mm arm length).

I Description. Five-armed species, consistently light-tan in color in fresh specimens. Relatively long

arms compared to disk diameter, tapering to small, pointed, terminal arm plate (Figure 6.8). Aboral surface

with small, reticulate plates, producing a somewhat retangular pattern, with I-2 papulae per area (Figure

I 6.8). Plates and spines not covered by membrane (Lissner, 1980).

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Figure 6.8. Aboralview of Myxodermaplatyacanthum (Station R-7, Rep. 2), showing body form and reticulate

I pattern of surface plates.

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Biology. Very little is known about the ecology of this species. Myxoderma platyacanthum is I

primarily found on green mud substrates, sometimes occurring in very high densities in trawl samples

(SAIC and MLML, 1992). Its occurrence in some oxygen minimum zones may be facilitated by the relatively II

high density of papulae (functioning in oxygen exchange) on the aboral surface. I

Distribution. Myxoderma platyacanthum ranges from Pt. Piedras Blanc.as,California to San Cristobal

Bay, Baja California (Fisher, 1930), from depths of 256-768 m (Maluf, 1988). I

Family Asteridae I

Rathbunaster californicus Fisher, 1906

6.9A and B I

Figures

Material Examined. California: Santa Mafia Basin, Sta. BRA-26 (USNM), 108-111 m (one li

specimen, fragmented, approximately 15-arms, 170 mm arm length). l

Description. Rathbunaster californicus a many-armed (e.g., 12-17) species, with arms

proportionately very long and easily detached. Tendency for arm detachment and much longer arm to disc II

ratio convenient factors helping to distinguish this species from another large, many-armed seastar, l

Pycnopodia helianthoides, co-occurring in the southern California region (D. Cadien in SCAMIT, 1994).

Body color typically light orange to light salmon in fresh specimens. Stalked, "pompom-like" pedicellariae II

are common on the aboral surface (Figure 6.9B), but also easily detached and may be missing on many arm I

areas of preserved specimens. Papillae long and vermiform (Lawrence, 1987). One inframarginal spine

present (Lissner, 1980). Podia in ambulacra numerous, occurring in rows of up to four across (Figure 6.9B). •

Biology. The species is usually associated with soft-botttoms of mud and sand, or shale (Maluf, I

1988). Little is known of its ecology, although individuals have been observed moving relatively quickly

across the bottom, suggesting an active predator (Lissner, pers. obs.). Potential prey that are common in this n

species habitat include polychaetes, molluscs, anemones, seapens, and other echinoderms (Lissner, pers. l

obs.).

Distribution. The species ranges from British Columbia (Lawrence, 1987) to San Diego, California, i

and possibly tOCabo San Lucas, Baja California (Maluf, 1988), at depths from 99-768 m. I

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_-5_re 6.9. A, Aboral view of Rathbunaster californicus (Station BRA-26), indicating extensive, stalked

pedicellariae on surface.B, Oral view ofRathbunaster californicus (StationBRA-26), emphasizing n

rows of numerous podia and large width of ambulacrum comparedto overall armwidth. IB

110 I
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I Literature Cited

I Anderson,12:499-503. R. and R. Shimek. 1993. A note on the feeding habits of some uncommon sea stars. Zoo Biology

i Barnes, R. 1980. Invertebrate Zoology. 4th Edition. Philadelphia, PA., Saunders. 1089 pp. Donnay, G. and D. Pawson. 1969. X-ray diffraction studies of echinoderm plates. Science 166:1147-1150.

i Feder, H. 1980. Asteroidea: The Seastars. Pp 116-135, Chapter 8,In: R. Morris, D. Abbott, and E. Haderlie (eds.): Intertidal Invertebrates of California. Stanford University Press. Stanford, CA.

Fell, H. 1963. The phylogeny of sea-stars. Philosophical Transactions of the Royal Society of London, B,

I 246:381-435.

Fisher, W. 1911. Asteroidea of the North Pacific and adjacent waters. Part 1. Phanerozonia and Spinulosa.

Bulletin of the U.S. National Museum 76:1-419.

i Fisher, W. 1928. Asteroidea of the North Pacific and adjacent waters. Part 2. Forcipulata (part). Bulletin of

the U.S. National Museum 76:1-81.

Ill

I Fisher, W. 1930. Asteroidea of the North Pacific and adjacent waters. Part 3. Forcipulata (concluded).

Bulletin of the U.S. National Museum 76:1-356.

I Gotshall,CA. E. and L. Laurent. 1979. Pacific Coast Subtidal Marine Invertebrates. Sea Challengers. Los Osos,

Hartman, O. 1955. Qualitative survey of the benthos of San Pedro Basin, Southern California. Allan Hancock

I Pacific Expeditions 19:409-411.

Hartman, O. and J.L. Barnard. 1958. The benthic fauna of the deep basins off southern California. Allan

I Hancock Pacific Expeditions 22:56-58.

Hopkins, T. and G. Crozier. 1966. Observations on the asteroid echinoderm fauna occurring in the shallow

water of southern California (intertidal to 60 meters). Bulletin of the Southern California Academy

I of Sciences 65:129-145.

Hyman, L. 1955. The Invertebrates: Echinodermata. McGraw-Hill Book Company. New York, NY. 763 pp.

I Jangoux, M. 1982. Digestive Systems: Asteroidea. Pages 235-272, In: M. Jangoux and J. Lawrence (eds.)

Echinoderm Nutrition. A.A. Balkema, Rotterdam, Netherlands.

I Lissner, ofA.theandsouthern J. Dorsey.Califor 1986.niaDeep-water continentalbiological borderland.assemblages Bulletin ofofthea har Southern d-bottomCalifornia bank-ridgeof Scociences mplex

85:87-101.

I of the benthic invertebrate collections: Echinodermata. Institution of Luke, S. 1982. Catalog Scripps

Oceanography Reference Series 82-5:2-24, 44-54.

I Lambert,Museum, P. 1945. Victoria, The Seastars Canada. of B153 ritishpp.Columbia. Handbook No. 39. The British Columbia Provincial

Maluf, L. 1988. Composition and Distribution of Central Eastern Pacific Echinoderms. Technical Report

I Number 2, Natural Museum of Los Angeles Los Angeles, CA. 242 History County, PP.

Margolin, A. 1976. Swimming of the sea cumberParastichopus californicusin response to sea-stars. Ophelia

I 15:105-114.

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II

Mauzey, K., C. Birkeland, and P. Dayton. 1968. Feeding behavior of asteroids and escape responses of their i

prey in the Puget Sound region. Ecology 49:603-619.

McEdwards, L. 1994. Evolution of pelagic direct development in the staff'lshPteraster tesselatus (Asteroidea: 1

Valvatida). Biological Journal of the Linnean Society 54:299-327.

i

Muscat, A. 1980. Echinodermata. Pages 263-273, In: D. Straughan and R. Klink (eds.) A taxonomic listing

of common marine invertebrate species from southern California. Technical Reports of the Allan |

Hancock Foundation 3.

Pequegnat, W. 1964. The epifauna of a California siltstone reef. Ecology 45:272-283. I

SAIC and MLML (Moss Landing Marine Laboratories). 1992. Trawl and Remotely Operated Vehicle,

Ocean Studies Report for Detailed Physical and Biological Oceanographic Studies for an Ocean

Site Designation Effort Under the Marine Protection, Research and Sanctuaries Act of 1972 I

(MPRSA). Final Report submitted under Contract No. 68-C8-0062 to U.S. Environmental Protection

Agency, Region IX, San Francisco, CA. i

II SCAMIT (Southern California Association of Marine Invertebrate Taxonomists). 1994. Proceedings of

August 1994 Meeting, Volume 13 (4):7-8. iI

Sloan, N. and S. Robinson. 1983. Winter feeding by asteroids on a subtidal sandbed in British Columbia, i

Canada. Ophelia 22:125-142.

Sutton, J.E. 1975. Class Asteroidea. Pages 623-627, In: R. Smith and J. Carlton (eds.) Light's Manual: I

Intertidal Invertebrates of the Central California Coast.University of California Press, Berkeley, i

CA.

i

Verrill, A. 1914. Monograph of the shallow-water starfishes of the North Pacific coast from the Arctic D

Ocean to California. Smithsonian Institution, Harriman Alaska Series 14:1-408.

Word, J. and D. Charwat. 1975. Invertebrates of Southern California Waters. I. Select groups of annelids, D

arthropods, echinoderms, and molluscs. Southern California Coastal Water Research Project. El B

Segundo, CA.

am

Ziesenhenne, F. 1941. Some notes on the distribution records of little-known Southern Californian i

echinoderms. Bulletin of the Southern California Academy of Sciences 40:117-120.

I

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I 7. CLASSOPHIUROIDEA

by

I Gordon Hendler 1

I

Introduction

I

Ophiuroids are the most agile and active echinoderms, the most diverse, and often the most abundant.

The Class Ophiureidea, with approximately 2,000 species, 250 genera, and 25 families extant, ranges from

I arctic to antarctic waters and from intertidal to abyssal and hadal habitats. Ophiuroids are almost exclusively

marine; few species are successful in brackish waters.

i In the Cenlral Eastern Pacific Region between Pt. Conception, California and Peru, nearly 200 species of ophiuroids have been reported (Maluf, 1988). Representatives of the Southeast Pacific fauna and

the less diverse Northeast Pacific fauna (Austin, 1985) occur in the southern Central California region

i treated inThethisAtlasaxonomic is basedAtonlas,thebuMinerals t only 19Management species are coService vered in(MMS) this report. Phase I and Phase II surveys of the

California outer continental shelf. The surprisingly few species obtained in the surveys probably reflects

I limitations and Pt. Estero, of therepresenting sampling program. localities Col onlylec70tionmiles s wereapart. restriCoverage cted to 50was -930further m depths, constrained betweenbyPt.theConception sampling

gear employed: small box cores deployed on soft sediments and a modest series of submersible and ROV

i samples from hard substrates. In a study of ophiuroids on the continental slope off Central California, Summers (1993) demonstrated that box cores undersample ophiuroid species diversity, missing approximately

half of the taxa recovered by trawls in the same area.

I ophiuroidsDespite of theitsPacific limitations, coasttheofsurvey the United permitsStates. a claThe rificatio resolution n of the oftaxotaxonomic nomic status matters of somprovides e widespread fresh

insight into the ecology of the Eastern Pacific benthos. The synonymy of Amphioplus hexacanthus with

I Dougaloplus boundaries andamphacan composition thus, ofisthea prime "'Amphioexample. pIus hexacanthus As a consequence community" of uniti (sensu ng the Barnard two nandominal Ziesenhenne, taxa, the

1961), a major element on the Southern California continental shelf, can and should be reevaluated.

I In this chapter the following species are treated:

Asteronyx longifissus Dougaloplus amphacanthus

i Amphichondrius Gorgonocephalus granulatus eucnemis Amphiurid Ophiacanthaae diplasia sp. juvenile

Amphiodia (Amphispina) digitata Ophiopholis baked

I Amphiodia (Amphispina) urtica Ophiuroconis bispinosa

Amphioplus (Amphioplus) strongyloplax Ophiothrix (Ophiothrix) spiculata

i AAmphipholis mphipholis puge squamata tana Oph Ophiura iosphalma leptoctenia jolliense

Amphiura arcystata Ophiura luetkenii

I A. diomedeae

I t Invertebrate Zoology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007.

i 113
!

Thorough descriptions and summaries of biological information are provided for most of these I

species for the first time. A lectotype is designated forAmphiura granulata LCitkenand Mortensen, and the

species is placed in Amphichondrius Nielsen. The photographic figures are intended to improve on the 1

inadequate illustrations previously available. The synonymies furnished are not necessarily exhaustive; I

they cover major shifts in nomenclature, making the taxonomic literature more accessible. The fists of

material examined include: locality, collecting station, depth, number of specimens, repository (or collection); I

names assigned to specimens in museum collections and by survey taxonomists, if incorrect, are given in II

brackets.

History I

Ophiuroids were among the creatures collected anddescribed by pre-Linnean naturalists. Linnaeus I

was aware of several species, all of which he placed in the genus Asterias. By the early 19th century the

considerable diversity of ophiuroids became more apparent. New genera were described, a difference between i

asteroids and ophiuroids was acknowledged, and two groups of ophiuroids were distinguished: simple armed

forms and those with branching arms. The early studies were synthesized and augmented by Miiller and

Troschel, and by Forbes, and in the latter half of the 19th century the taxonomic treatments of ophiuroids •

accelerated as specimens from international expeditions became available. During that period, major

contributors included Liitken, Ljungman, Lyman, and Verrilt. Taxonomists of the early 20th century who

studied the wealth of new material from major oceanographic explorations left an enduring mark on ophiuroid n

systematics; among the most prolific were Koehler, Drderlein, Matsumoto, and Mortensen, and later H. L. II

Clark and A. M. Clark. Brief historical reviews of these developments appear in Fell (I960) and Hyman

(1955). Surprisingly I little effort was ever directed toward the study of the Eastern Pacific ophiuroid fauna.

Early workers including Ltitken, Ljungman, Lyman, Verrill and others, helped to delineate the shallowwater fauna. Major contributions on the offshore fauna by Liitken and Mortensen (1899) and H. L. Clark i

(1911, 1917, 1923), were based on explorations of the fisheries research vessel Albatross. Supplemental II

studies made by H. L. Clark (1901, 1902, 1910, 1913, 1915), MeClendon (1909), and May (1924) were

incorporated in Nielsen' s (1932) overview of the fauna between Southern California and British Columbia. 1

Later publications by Ziesenhenne (1937, 1940) also augmented knowledge of region's fauna.

A contemporary, worldwide, monographic treatment of the Ophiuroidea is unavailable, Lyman's

(I 882) Challenger report being long out of date. There are no monographic works on the ophiuroids of the _l

Eastern Pacific. Fortunately, Fell's (1960) synoptic key to the genera of ophiuroids and H. L. Clark's (1915)

catalog of Recent species are apphcable to the region.

I

Systematics

! Early taxonomic works assigned ophiumid species to genera and families, and segregated the families

into two or three major divisions. After the turn of the century, the divisions Euryalae and Ophiurae generally

used, the former including animals with branching arms (basket stars), and the latter consisting of •

were

most simple-armed species (brittle stars). The basis for the separation was the nature of the arm-joint

Ill

articulation, a poorly defined characteristic. Then and now there were ambiguously def'med genera and /m

families, and species whose placement was in question. 1

I

114 I
!

I Matsumoto's (1917) iconoclastic monograph proposed a phylogeny of the Ophiuroidea based on

morphology of the genital plate, genital scale, radial shields, and other internal and external ossicles.

i Murakami's (1963) analysis of the oral and dental plates supported Matsumoto's arrangement to a great extent. Their contributions underlie Fell's (1962) synoptic key to the genera of ophiuroids. In turn, his

arrangement was the basis for Spencer and Wright's (1966) widely used classification.

I species extant), Spencer theandPhrynophiurida, Wright recognized and the threeOphiurida. orders ofThe livingPhrynophiurida Ophiuroidea: isthecomposed Oegophiurida of the (with Suborder one

Ophiomyxina (consisting of one family) and the Suborder Euryalina, both groups characterized by a thick,

I dense were formerly integument. classified The Euryalina as Euryalae. includes The four Ophiurida familiesis composed of basket stars of taxaandwith relatedreduced taxa, integument which together and

relatively robust disk and arm plates. Within the Ophiurida, Spencer and Wright recognize the Suborders

I Chilophiurina, Laemophiurina, and Gnathophiurina, the distinctions between the suborders based on internal skeletal morphology.

Smith et al. (1995) have published a cladistic analysis of the class based on the type genera of 26

I Recent They conclude, families,since and only usingparts dataofforthe43most morphological parsimoniouscharacters, cladogram largely seem securely drawn from founded, published that a complete sources.

revision of the higher taxonomy of ophiuroids is still required. However, they feel there is sufficient support

I tofrjustify all othera classificatory taxa. Within scheme the Ophiuridea of two subclasses: they recognize Oegophiuridea the Orders for Euryalida the Ophiocanopidae, with four families and Ophiuridea (equivalent

to the Euryalina mentioned above), and the Ophiuridea. The latter is equivalent to the Ophiurida, except that

I itOphiurina includes the constitutes SubordertheOphiomyxina, remainder ofexcluding the SubclasstheOphiuridea, subfamily Ophiobyrsinae and includes the asincertae infraorders sedisHemieuryalina . The Suborder,,

Chilophiurina, Gnathophiurina, Ophiodermatina, and Ophiolepidina. It is likely that a number of these

I recommendations considering additional willtaxa be adopted and a revised in thesetfuture. of characters. However, a more comprehensive analysis is needed

I Morphology

I Despite a deceptive similarity in form, ophiuroids show a considerable range of size and body plans. At one extreme are commensal species several millimeters in diameter, and at the other are basketstars

with intricately branched arms spanning a meter. The general anatomy of ophiuroids is covered in Hyman

I (1955) and Spencer and Wright (1966), and their microscopic anatomy is reviewed by Byrne (1994). Information on ophiuroid reproduction and development is summarized by Hendler (1991).

Major anatomical structures of ophiuroids are depicted in Figure 7.1. The disk and arms are well

I demarcated, and protected by series of integument-covered ossicles called shields, plates, scales, spines, and granules. Minute, finger-like, fluid-filled tube feet emerge from each ann joint. Some species employ

the tube feet for locomotion, but most move by flexing and extending the arms to push the disk ahead;

i several species can swim. The adhesive properties of the tube feet enable some ophiuroids to scale smooth vertical surfaces and many to catch and manipulate particles of food.

Within the disk are a capacious stomach, a series of gonads, and sac-shaped bursae. The latter

I structures, which are invaginations of the body wall functioning in respiration and reproduction, through

open

bursal slits beside the base of the arms. The stomach is a blind sac; an anus is absent. The stomach and

gonads are situated within a fluid-filled perivisceral coelom. Ophiuroids, like other echinoderms, have

I additional coelomic structures including the hemal and the water-vascular systems.

!

i 115
In many species, pairs of radial shields are evident on the dorsal surface of the disk near the base of •

the arm. Near its distal tip, the radial shield articulates with a genital plate which is attached to a part of the

arm within the disk, and a genital scale which supports the outer adradial edge of the bursal slit. Conspicuous ll

structures below the disk include five oral shields, at least one of which (incorporating the madreporite) i

typically is enlarged and perforate. Beside each oral shield is a pair of adoral shields, homologous with

lateral arm plates. II

According to many accounts, ophiuroids move their arms only in a horizontal plane. However, the I_

arms are composed of a series of moveable joints, and in most species are capable of a degree of vertical

bending or coiling. At the core of the arm are the vertebrae, ossicles resembling bones of the human spine, I

which are connected in series by muscles and connective tissues. Modified arm ossicles are joined in a ring U

beneath the disk, forming the five triangular jaws that frame a centrally placed mouth. Attached to the jaws

are ossicles called oral papillae and/or dental papillae, and teeth. The ring of jaws, oral shields, adoral l1

shields, and associated ossicles is referred to as the oral frame. II

The arm joints beneath and beyond the edge of the disk have protective lateral arm plates bearing

arm spines; the joints are capped above and below by dorsal and ventral arm plates. The arm spines may be I

well-developed or inconspicuous, and they and the arm hooks and tentacle scales serve in feeding and II

defense. At the tip of the arm is a cylindrical ossicle, the terminal plate, through which the distal-most tube

foot protrudes. New arm joints arise at the inner edge of the terminal plate. Therefore, the youngest joints ll

are those nearest the arm tip. As the disk grows, new plates arise between older ones, and the disk overgrows II

the oldest, most proximal arm joints as it enlarges.

The reproductive organs (testes and ovaries) of ophiuroids consist of clusters of gamete-lined tubules I

thattypically are housed within the disk; they occur inside the arms in only two families. Spawning ophiuroids II

assume a "push-up" posture, raising their disks off the substrate while releasing gametes from gonoduct

openings near the edge of the bursal slits. Fertilization is extemal, n

g

Ophiuroid larvae develop for a few days to over a month before metamorphosing to juvenile

ophiuroids. Some species have large, yolky eggs and abbreviated development. Their non-feeding, barrelshaped vitellaria larvae swim using rings of cilia, and metamorphose in less than a week. In contrast, many 1

species produce hundreds of thousands of tiny eggs, which develop into multi-armed, long-lived larvae. The g

larva is a transluscent, microscopic ophiopluteus, with anintricate crystalline supporting skeleton, and with

sinuous ciliary bands that are used for food gathering and locomotion. I

16

A minority of ophiuroid species are hermaphroditic, either sequentially (transforming from one sex

to another), or simultaneously. There is a correlation between hermaphroditism and mode of development, roll

all hermaphroditic ophiuroids (and a few species with separate sexes) brooding their young. Their embryos 1

are retained in a bursa of the parent where they develop into miniature "crawl-away" young. Relatively few

Ill

ophiuroids are capable of asexual reproduction by a process called fission, whereby autotomy across the iBm

disk produces a pair of half-ophiuroids. The halves heal, regenerate missing structures, grow, and can split !

again.

tll

Ecology and Behavior I

Ophiuroids are members of many benthic communities, from tide pools to the deep sea (Gage and I

Tyler, 1991; Hendleret al., 1995). As Bamard and Ziesenhenne (1961:136) point out, "ophiuroid communities

I

cover a relatively large share of southern California coastal shelf bottoms: roughly 26% of the 916 nautical

square miles (3,125 square km) between Pt. Conception and the Mexican border...in depths of 0-50 fathoms n

(0-91.5 meters)." Their conclusion was substantiated by data from surveys of the shelf, island, and basin

faunas of the southern California borderland (Hartman, 1955, 1966; Hartman and Barnard, 1958, 1960;

Jones 1969). I

116 I
I

I

I dorsalarm plate ventralarm plate

accessorydorsal

I tube foot

arm spme . tentaclescale

i

| arm

I A

first ventral

i disk -- arm plate

-- mouth

primary plates:

I { centralplate radial plate D -- bursal slit

l radialshield

I ,_ ,aw

i__ tooth

oral papilla

q oraltentacle

i ' _ oralshield

' "; iV" adoralshield

'

"" " _ genital papilla

_ nital scale

I

I

Figure 7.1. Major anatomicalfeaturesof anophiuroid:A, thedorsalsurfaceof thedisk; B, thedorsalsurface of

i an arm; C, the ventral surface of an arm;D, the ventral surface of the disk; E, a ventral interradius and the jaw structures that project into the mouth. (AfterHendler et af., 1995.)

I

i 117
!

The significance of ophiuroids in benthic communities is reflected in their abundance and distribution. I

Burrowing amphiurids frequently number in the thousands per square meter, and epifaunal species such as

ophiurids can literally carpet the seafioor. Despite their seemingly refractory skeleton, ophiuroids figure in ti

the diets of commercially important crabs, shrimp, and fish, and they are consumed by sea stars and other II

ophiuroids. Surprisingly, the extent of their ecological role is largely untested, and their interactions with

other organisms are poorly documented. Unfortunately, the natural history of not a single endemic Eastern I

Pacific ophiuroid species has been well and thoroughly described. I

The feeding behaviors of ophiuroids are flexible and remarkably diverse. They can take up dissolved

compounds through their integument, and they convey particulate food to their mouths with tube feet and in i

loops of the arms. Some ophiuroids quickly coil their arms, particularly the slender tips of the arms, and II

capture small active organisms including fishes. The tube feet are capable of sensing extremely dilute

concentrations of chemicals such as amino acids and vitamins, enabling individuals to detect food. Coordinated I

movements of the hundreds of feet under the arm can collect and relay minute particles with great dexterity. l

The characterization of all ophiuroids as detritivores is inaccurate. Macroparticulate feeders may be carnivores

or scavengers with either restricted or relatively indiscriminate diets. Specialized deposit feeders and •

suspension feeders harvest microparticulate material. The subject of ophiuroid feeding is reviewed by Warner II

(1982).

Most species remain concealed during the day, perhaps avoiding diurnal predators. Ophiuroids lack •

eyes, but it is thought that nerves beneath microscopic transparent nodules (expanded peripheral trabeculae

= EPT) in the skeletal plates are photosensitive. Furthermore, the discovery that some ophiuroids can change

color suggests how the amount of light striking the nerves is controlled. Pigment-containing cells shield the

transparent nodules during the day and uncover them at night. The adjustment enables ophiuroids to detect I

crevices in sunlight or moonlight, an ability that is essential for their escape from predators and competitors

in shallow water. The colors of most species are fixed, although pigmentation patterns may transform •

dramatically as individuals grow. Color patterns of an individual are as specific as a f'mgerprint, and in some I!

species intraspecific variations can be striking.

Ophiuroids owe their common name "brittle stars" to a remarkable capability for antotomy, the 1

ability to voluntarily break their arms. This behavior is potentially advantageous when an individual is

attacked by a predator. Some species (probably most Amphiuridae) can also autotomize the disk, and with

it the stomach, gonads, and other tissues. That process is all the more remarkable because these individuals

can completely regenerate the broken arms or regrow the disk and associated viscera, in several weeks or

months. Autotomy in ophiuroids is effected by nervously-controlled mumble collagenous tissue, the same

tissue permitting the arms to flex and soften. Other ophiuroid defenses include distastefulness and

luminescence, speedy escape, and the shadow-response, which is a stop-motion reflex triggered by passing

predators.

Major ecological and morphological differences among groups of ophiuroids are most readily

recognized at the family level. Ophiuroids may show greater fidelity in body plan to the substratum they It

occupy than to phylogenetic affinity or distribution. For example, species in the family Amphiuridae living

in soft sediments have adaptations for burrowing, such as soft disks and long delicate arms, which would

jeopardize their survival in an exposed habitat. Burrowing OphiopsiIa species, in the family Ophiocomidae,

are remarkably similar in appearance to amphiurids. The huh'owing ophiuroids maintain semi-permanent

burrows composed of a mucous coated chamber that surrounds several upraised arms and the disk. The

arms undulate to pump water through the burrow, and they can be extended to the surface to convey waste

material and railings from the burrow, and carry food particles towards the disk. Species in several families,

which are epizoic on soft coral and sponges, have hooked spines used to cling to their host and similar II

structural and behavioral defenses against predation. II

I

118 i
!

I Collection and Preservation

I Ophiuroids occur on, within, and under many benthic substrates in the marine environment, and

some even live on pelagic scyphozoans. Their delicate construction, and the tendency of ophiuroids to

autotomize makes it advantageous to isolate them from samples before more resistant taxa are processed. It

I is sometimes helpful to allow material, such as rubble, to rest in a bucket of water it in order to permit delicate cryptic fauna to emerge, before fragmenting the substrate.

A full and up-to-date account of techniques for the preservation of ophiuroids and other echinoderms

I is in Hendler et al. (1995). When possible, freshly collected ophiuroids should be anesthetized by

presented

placing them in a solution of magnesium chloride that is isosmotic with seawater, or by gradually adding

magnesium sulfate (epsom salts) to the seawater in which ophiuroids are held. Other invertebrate anesthetics

I be effective to a degree, but gradually adding alcohol to seawater is not a reliable means to relax

may

ophiuroids. In contrast, basketstars, which do not respond to magnesium salts, are best preserved by holding

individuals with extended arms in a tray of concentrated alcohol so that they quickly die with the branches

I extended.

Anesthetized specimens, with their arms disentangled, can be briefly fixed in buffered 10% formalin

or placed directly in 70-80% ethanol. In either case, specimens should be washed in distilled water to

I remove formalin and excess salts before their permanent storage in ethanol. Museum specimens of ophiuroids

are sometimes dried because microscopic details are most clearly seen using desiccated material, and because

maintaining alcohol levels requires additional resources. However, it is preferable to maintain specimens in

I long-term storage, partially or completely drying only required

alcohol for them when for examination.

That approach permits the conservation of soft tissue and lessens the potential for damage by fungi and

other pests.

I

Glossary

I Accessory dorsal arm plate. In some ophiuroids, Arm length. A body dimension of art ophiuroid

one or several small, symmetrically measured from the edge of the disk to the tip

• _ arranged plates that are inserted between the of an arm.

W

dorsal arm plate and the lateral arm plate. Arm spines. Spines attached to the lateral arm

I

i Aborapar l. In adirection away from the mouth; the plate.

t of the body opposite the mouth. Asexual reproduction. Reproduction that occurs

Adoral shields. A pair of plates, one of which is without the fusion of male and female

I found at each side of the oral shield, gametes, by splitting of the body into two parts that regenerate. The genetically

Arm. A moveable, jointed ambulacral projection, identical offspring of asexual parents are a

i distal to the disk, that carries a radial branch clone. of the water vascular system and the

nervous system. Autotomy. A defensive process initiated in

response to adverse stimuli, involving the

I Arm joint. One of a series of articulating units

loss of portions of the arlns or disk.

comprising the arm, consisting of an internal

vertebral ossicle, the dorsal, lateral and Brooding. Reproductive mode in which the

I ventral arm plates, and associated structures, embryos parent, andareemerge protectedas tiny, on, incrawl-away , or beneath the

juveniles.

I

I 119
!

I

Bursa. An organ within the disk formed by an EPT (= expanded peripheral trabeculae). I_

inpouching of epidermis (see bursal slit). Microscopic, transparent nodules on the

Bursae function as respiratory structures and surface of skeletal plates. In some II

are associated with the gonoducts. They ophiuroids they are a component of a I

house the developing embryos of brooding photoreceptor system.

species. Fission. Asexual reproduction by splitting of the I

Bursal slit. The opening of a bursa, located on the body into two parts, each of which

ventral interradius of the disk at the base of regenerates into a complete animal. i

the arm. Typically, there is one bursal slit on Genital papilla. Granules or spinules attached to 1

each side of an arm. I

the edge of the bursal slit. The term is also

Calcite. The mineral form of calcium carbonate used (perhaps more appropriately) for the •

that makes up the echinoderm skeleton, fleshy outlets of the gonoducts in ophiuroids 1

dd. See disk diameter, that lack bursae.

Genital plate. A bar-like ossicte connecting the I 

Dental papillae. See papillae, radial shield to the arm and supporting the

Dermis. The stratum of cells beneath the radial edge of the bursal slit.

roll

epidermal covering of the body wall. In

Genital scale. An ossicle connected to the genital l

echinoderms, the skeleton develops in the

dermis and is filled with dermal tissue, plate, which supports the adradial edge of

the bursal slit. mum

Disk. The round or pentagonal central body region Granules. See skeletal elements. I

of an ophiuroid.

Disk diameter. A body dimension of an ophiuroid HookS.articulateMinute, withm°veabossiclesC le'arm rescenticin ossicles that I

measured from the distal edge of a pair of gorgonocephalid ophiuroids (other

radial shields to the disk edge in the ophiuroids may have hook-shaped arm

opposite interradius. Abbreviated dd.

spines attached to the lateral arm plates). I

Distal. In a direction away from the center of the lnfradental papillae. See papillae.

body; for example, towards the tip of the l1

arm. Interradial. (interradii). Referring to I

interambulacral sectors of the body, dorsal

Dorsal. In echinoderms, this term is variously or ventral regions between two arms.

applied, but in ophiuroids it refers to the l

surface of the body that is opposite the Jaw. A moveable triangular structure that extends I

mouth, into the mouth.

m

Dosal arm plate. A plate on the aboral surface of Joint. See arm joint, l

an ophiuroid armjoint. Larva. An early developmental stage that is

Echinulate. Referring to a roughened texture independent of the fertilization membrane l

composed of tiny spines, andthe parent (pl. larvae). Through

metamorphosis it transforms into ajuvenile.

Embryo. An early developmental stage that is m

enclosed in a fertilization membrane or Lateral arm plates. Paired plates covering the l

protected by the body of the parent. It sides of each ann joint and bearing the arm

transforms into a juvenile through spines.

metamorphosis. I

120 I
!

I Lecithotrophy. A mode of reproduction in which Papillae. In ophiuroids, certain skeletal elements

free-swimming larvae develop using of the jaws or disk: dental papillae - a

I nuLecithotrophic trients laid down in the egg. cluster of small, blunt, spinelike structures larvae do not feed on on the dental plate, near the ventral tip of

particulate matter, but they may supplement the jaw. Fence papillae - peg-shaped

i yolk reserves by the uptake of nutrients spinelets fringing the edge of the disk of

dissolved in seawater. Ophiophragmus species. Infradental

papillae - in amphiurid ophiuroids, a pair of

Madreporite. A perforated plate that is connected small, blocklike plates attached at the

I ventral tip of the jaw, below the teeth. Oral

to the water-vascular ring by a so-called

stone canal. In ophiuroids it is a modified papillae - small plates at the edge of the

oral shield, mouth, attached to the edges of the jaw plate

I OFL. See oral frame length, to papillae can

and/or the adoral shield. Oral

be variously shaped, from spinelike to

Ophiopluteus. The free-swimming larval stage of scalelike. Also see genital papillae.

I an ophiuroid. Ophioplutei have appendages that are supported by skeletal rods and bear Planktotrophy. The mode of development of

ciliary bands. During metamorphosis, the free-swimming larvae that feed on

I juvenile on the ventral ophiuroid surfacedevel ofothe ps from larva;atherudiment partic plankton. ulate matter obtained from the

process may involve resorption or loss of Plates. See skeletal elements.

I parts of larval structures. Primary plates. The first-formed plates on the

Oral. In a direction towards the mouth; a part of dorsal side of the disk. In ophiuroids these

i Oral Frtheame. bodyTheon the seriessameof ossicles surface asencircling the mouth, the are adults, the they centrmay al andformfivea rad rosette ial poflatesscales . In

mouth, including the oral, dental, and near the center of the disk, or they may be

I peristlates,omial the oral plates, and the adoral firstshveields, ntral and arm separated developed byscales. numerous secondarily

associated structures. Proximal. Towards the center of the body.

I Radial. In a direction towards the central axis of

Oral flame length. An index of ophiuroid body

size measured from the distal edge of an an arm or ambulacrum; a part of the body

oral shield to the distal edge of the first

I ventral arm plate in the oral slit of the near an arm or ambulacrum. opposite interradius. Abbreviated oil Radial shields. Pairs of plates on the dorsal

surface of the ophiuroid disk, which lie near

I Oral papillae. See papillae, the base of each arm. They are usually Oral shield. A relatively large plate at the distal relatively large and conspicuous, but may be

end of the jaw. At least one of the oral hidden by granules or superficial scales.

i shields is modified as a madreporite. Rheophilic. Literally, "current-loving."

Oral tentacles. Tube feet inside the mouth, Rheophilic organisms prefer habitats with

i arising from the jaws. Also referred to as significant water movement. Rheophobie oral tube feet. organisms avoid currents, and occupy

sheltered positions or are restricted to lowOssiele. A small, usually microscopic skeletal energy habitats.

I element, is sometimes embedded used toinrefer integument; to all skeletal the term Scales. See skeletal elements.

elements.

I

i 121
!

Skeletal elements. Supporting and protective Tentacle scales. Small, moveable spines or scales, •

dermal structures consisting of a calcite associated with ophiuroid tube feet, which

meshwork (stereom) and invested with a are attached to the ventral arm plate and/or •

thin layer of tissue (stroma). Various lateral arm plate. Tentacle scales may cover 1

skeletal elements are distinguished as the tentacle pores and protect the retracted

follows: plates are tabular structures with a tube feet. •

characteristic shape and a fixed position; I

scales are fiat, thin structures that are Trifid. Divided into three parts. Regarding "trifid

overlapping, tesselate, or haphazardly spinule" see skeletal elements. I

arrayed; spines are moveable, articulating Tube feel Fluid-filled, fingerlike extensions of I

structures that are long, slender, and the water vascular system, which protrude

attenuated. Small structures fixed to the through openings in the skeleton or between m

surface of scales or plates include: granules skeletal elements. Specialized tube feet are 1

that are minute and nearly equidimensional; used for locomotion, feeding, burrowing,

spinelets that are enlarged, elongate respiration, and a combination of functions 1

cylindrical or angular granules (this term is (see papillae, tentacles). 1

sometimes applied to small spines);

spinules that have various numbers of Tubercle. See skeletal elements. m..

pointed apical projections (e.g., bifid, trifid, Ventral. In echinoderms, this term is variously I

multifid). Also fixed, and relatively larger applied, but in ophiuroids it is the surface of

than granules, are small structures referred the body that carries the mouth. n

to as stumps, which are usually prickly, and •

tubercles which are smooth and more Ventral arm plate. A plate on the oral surface of B

massive. In addition, tubercles (see below) each arm joint. mm

Can refer to outgrowthsof plates, rather than Vertebra. An internal ossicle within an arm joinL •

to separate elements. Also see papillae, connected by ligament and muscle to the

roll

Spines. See skeletal elements, vertebrae of adjacent joints. It is so named

because of a resemblance to bones in the •

Spinelets. See skeletal elements, human spinal column.

I

Stone canal. A tube, usually reinforced with Vitellaria. A type of free-swimming •

ossicles, leading from the madreporite to the lecithotrophic echinoderm larva. It is barrelwater-vascular ring canal, shaped, has several transverse rings of

Teeth. Small plates or spines attached to the locomotory cilia, and lacks a mouth. I

dental plate on the inner edge of the jaw, a

series extending into the mouth.

m

Tentacle pore. An opening between the ventral I

arm plate and the lateral arm plate, through

which a tube foot projects. Each arm joint •

has two tentacle pores.

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122 I
!

l Terminology and Conventions

I The colors noted in the species accounts pertain to living ophiuroids, unless otherwise noted; pigmentation and color patterns are altered or removed by preservatives. Arm structures are described for

the proximal third of the arm, unless otherwise noted. Towards the tip of the arm the relative dimensions of

I the ossicles change; the arm spines and tentacle papillae diminish in number. The numbers of arm spines and tentacle scales refer to the number on each side of an arm joint on a portion of the arm near the disk. For

example, "this species has 3 arm spines" means that near the disk there are 6 arm spines on each joint, and

I 3 on each lateral arm plate. Similarly, the number of oral papillae cited for a species refers to the number on one side of a jaw.

Disk diameter (abbreviated dd) is measured from the distal edge of a pair of radial shields to a point

I on the opposite edge of the disk. Oral frame length (abbreviated off) is measured from the outer edge of an oral shield to the outer edge of the opposite first ventral arm plate. Arm length is the distance from the disk

edge (the base of an arm) to the tip of the longest arm.

I The abbreviations used are: AHF, Allan Hancock Foundation of the University of Southern California;

CAMP, California Phase II Monitoring Program; LACM, Natural History Museum of Los Angeles County;

SBMNH, Santa Barbara Museum of Natural History; USNM, National Museum of Natural History; ZUUC,

I Zoological Museum, University of Copenhagen.

l Description of Species

i Order Phrynophiurida

Family Asteronychidae

I Asteronyx iongifissus Diiderlein, 1927

Figure 7.2

I Asteronyx longifissus Dtderlein, 1927:58, 59, 65-68, 97, pl. 7, figs.l-3.--A. M. Clark, 1965:68.--Thompson

et al., 1993:30, 38.

I Material Examined. California: Santa Maria Basin, Phase I Sta. BSR-025, 390 m (1, SBMNH

secondary voucher [as Asteronyx loveni]); Phase I Sta. BSR-036, 492 m (1, USNM primary voucher [as

I Aste LACMronyx77-204.1).--Off loveni]).---Off Newport HuntingtBeach, on BeaRc/h,V Velero/V RIV Thomas Sta. AHF G. Th22631-75,533 ompson Sta. BLM m (1, LACM 821,68775-148.1).-- -698 m (1,

Off Dana Point, R/V Velero/V Sta. AHF 22723-75, 611 m (3, LACM 75-226.1).

I Description. Moderately large ophiuroid with disk diameter reaching 31 mm, arm length generally 5-9 times, and up to 13 times diameter of disk. Disk and slender, markedly tapering arms smooth, apparently

naked. Scales in integument only conspicuous in small specimens.

I Disk five lobed, indented interradially, appearing naked but for dense layer of small thin scales covering radial shields. Radial shields thin, riblike; proximal end nearly reaching center of disk, distal end

expanded, hoof-shaped, articulating with similarly shaped projection of genital plate.

I Jaws elongate, ventrally projecting. Oral shields inconspicous, frequently overgrown by adoral

shields. Adoral shields large, curved, bridging first lateral arm plates of adjacent arms.

I

I 123
I

I

I

I

I

I

I

I

I

I

I

I

Figure 7.2. Asteronyx longifissus D6derlein: A, dorsal view; B, disk, ventral view; C, distal portion of an arm, I

ventralview, showingelongate,club-tipped ventral-mostspineand hooked dorsal armspines(LACM

77-204.1; dd = 25.0 ram).

I

Genital slit large, extending from first or second arm joint to edge of disk. Ventral interradius nearly

naked, with Armsscattered, all of equal embedded length scales; in juveniles; minutearms scalesof atuneven edge oflength bursalinslit. adults. Arms covered with thick, i

translucent integument, markedly arched dorsally, flattened ventrally, narrowing abruptly beyond edge of

disk. Ventral Dorsalsurface of arm beneath disk markedly concave and broad. I and ventral arm plates reduced to microscopic, thin, perforated scales, numerous, irregularly

arranged, embedded in integument. Arm vertebrae visible through integument, appearing W-shaped dorsally,

V-shaped ventrally. I

Lateral arm plates situated ventrally on arm. Arm spines 5 or more, hooklike; base bulbous, opaque;

distal portion with multiple, hyaline teeth. First several arm joints lacking spines; on distal arm joints

ventralmost spine elongate, club-tipped. Lateral arm plates and spine-groups forming paw-shaped structures I

on distal arm joints.

124 i
!

I Remarks. The range of the species ovedaps that of Asteronyx loveni Mfiller and Troschel. The

latter species is distinguished by its relatively short bursal slits, which are restricted to the inner half of the

i ventral interradius. Di D/Sdertein (1927) provides a key to Asteronyx species and related taxa. stribution. Point Conception to San Diego, 266-1800 m (Dfderlein, 1927).

Biology. The autecology of A. Iongifissus is probably similar to that described (Fujita and Ohta,

I 1988) for the suspension-feedingA, loveni, which clings to stalked cnidaria. Thompson at al. (1993) indicated that it is one of the most abundant megabenthic invertebrates trawled at mid and lower slope depths (300-

780 m) of the San Pedro and Santa Monica basins in Southern California. They suggested that its population

I density may have increased by an order of magnitude following an E1Nifio event.

I Family Gorgonocephalidae

Gorgonocephalus eucnemis (Miiller and Troschel, 1842)

I Figure 7.3

Astrophyton eucnemis Miiller and Troschel, 1842:123.--Ljungman, 1867:335.--Lyman, 1882:263.

I GorgonocephaIus eucnemis, Lyman, 1880:42.--D6dedein 1900:226, pl. 10,fig. 1-4.--Koehler, 1907: 349.--

1911:31, 33,103-104.--H.L. Clark, 191l:185.---Campbell, 1921a:4.--1921b:49.--May, 1924:270-

i 271.--Boolootian and Leighton, 1966:4, 8, fig. 3.--Austin and Hadfield, 1980:154-155, pl. 51, fig. 10.12.--Irimura (in Imaoka et al.), 1990:72-73, figs. A-D.

Astrophyton Caryi Lyman, 1860:424-425.--1882:264.--Ljungman, 1867:335.--Verdll, 1867:326.

I Gorgonocephalus caryi, Verrill, 1867:253.--Lyman, 1882:264.--D6derlein 1911:31, 104.--H.L. Clark, 1911:5, 9, 17-18, 20-21,287-292.--Matsumoto, 1917: 61, 64, 70, 71,354.----Campbell, 1921b:49.--

D' yakonov, 1954:18-19.--Astrahantseff and Alton, 1965:1414-1415, 1419, 1422.--Kyte, 1969:1729,

I 1730, 1735-1737.--Smirnov, 1882:113, 116. Gorgonocephalus Caryi, Lyman, 1880:42.

II Ast Gorgonocephalus rophyton stimpsonii Verrill, 1869: 12:388.--Lyman, 1882:264. stimpsoni, D6derlein, 1911:104.

Gorgonocephalusjaponicus D6derlein, 1902:321.--1911:30-33, 104, pl. 1, figs. 1-3, pl. 7, figs. 1-2c.

I Gorgonocephalus sagaminus Doflein, 1906:204, fig.

Material Examined. Oregon: Off Newport, R/V Waseca Sta. AHF 1481-42, 84-132 m (1, LACM

I 42- vou3c9.4).---California: her).---Off Santa Catalina Western Santa Id., R/BVarbara VeIero/V Channel, Sta. Phase AHF 10698-65, I Sta. BRA-002, 265-290110-126 m (1,mLACM (1, USNM65-333.1). primary

Description. A large basket star with numerous arm branches. Disk diameter reaching 140 ram,

I extended arms 4-5 times as long; total diameter exceeding 0.5 m with arms outstretched.

Disk massive, conspicuously indented between the radial shields, covered with thick integument.

Radial shields tapering, nearly meeting at center of disk; thick, truncate distally, thin, narrowing proximally.

I Radial shields densely covered with scales bearing roughened similar scales concentrated disk

granules, at

margin; dorsal interradial region of disk nearly naked, scattered with small granules. Expanded hoof-shaped

distal tip of radial shield articulating with similarly shaped projection on genital plate.

I Jaws broad, ventrally directed. Teeth and oral papillae spinelike. Oral frame ossicles obscured by

thick integument.

I

i 125
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Ventral interradius nearly naked, with small scattered granules and scales. Bursal slit beginning •

near first arm branching, restricted to outer half of interradius. Edge of genital scale bearing large granules

or spinelets. Arms complexly branching, coiling; terminal portion of arm with fine, coifing branches. First arm I

branching beneath disk, second branching at or beyond disk edge; 6 joints before first branching, 9 joints to

second branching. Arms higher than wide in cross section; covered with thick integument; edges of lateral n

arm plates forming thick ridges below disk.

Conspicuous dorsal and ventral arm plates lacking; embedded scales and granules covering dorsal

and lateral arm surfaces; microscopic, thin, perforated plates covering ventral surface. Microscopic, flesh- n

covered, crescentic, hyaline arm hooks, borne on embedded ossicles; hooks present after 2nd arm branching, U

becoming more numerous on distal joints; 1-2 hooks on proximal joints; by 6th branching, arms annulated

with bands of hooks on ridges girdling the sides and top of each joint, n

Lateral arm plates positioned sublaterally, bearing up to 5 echinulate spines on 1st and 2nd arm

branches; 4 or fewer spines on distal branches, hooked, usually with 3 hyaline teeth. Articulated tentacle

scales absent; granules embedded in walls of some tube feet. I_

u

Color: Pigmentation ranging from maroon, reddish, brown, tan, beige, orange, salmon, pink, to

almost white; the disk usually darker than the arms. The dorsal surface is more intensely colored than the m

ventral surface. Frequently, coloration is not uniform. According to Patent (1970b), in a collection of 100 l

animals 34 were brown, 45 pink or coral, 18 orange, and 3 yellowish.

Remarks. Extensive collections of Gorgonocephalus from Atlantic and Pacific localities must be []

compared for the taxonomy of Californian basket stars to be resolved. In the present report, G. eucnemis is II

regarded as a senior synonym of G. caryi because none of the characteristics cited in the literature reliably

distinguish the two nominal taxa. []

Lyman (1960: 425) separated the small, unique type-specimen of A. caryi from A. eucnemis by II

"...more regular granulation of the disk, by having blunter tentacle-scales, and by close rows of grains along

the edge of the genital slits." On the basis of extensive northern Pacific collections, H. L. Clark (1911) •

synonymized G. japonicus and G. stimpsonii with G. caryi, but he reserved judgement on the distinction IN

between G. caryi and G. eucnemis pending study of the latter species. Moreover, he suspected that"...agassizii,

eucnemis, and caryi may prove to be three names for a single species. Material from the west coast of ',n

Greenland and the regions north of America would be very desirable in settling the question" (H.L. Clark, II

1911:292). Without further explanation, H. L. Clark (1915) synonymized G. caryi and eucnemis, as did

Downey (1969) and Irimura (in Imaoka et al., 1990). D6derlein (1927), D' yak•nov (1954), and Kyte (1969) []

recognized both nominal taxa. II

GorgonocephaIus diomedeae LiJtken and Mortensen is the only eastern Pacific congener of G.

eucnemis. Described from off Panama at 1270 m, it is distinguished from G. eucnemis by the short spines []

covering its disk. []

Distribution. California to Alaska, Sea of Japan, and Korea Strait to Bering Sea, Okhotsk Sea and

Laptev Sea, Finmark, Spitzbergen, and across the Arctic Circle to Cape Cod and Greenland; at depths from i

15-1850 m. U

Biology. The species is rheophilic, clinging to various cnidaria and sponges in high current areas mira

(May, 1924; Astrahantseff and Alton, 1965; Austin and Hadfield, 1980). Photographs taken in sit-u, off E

Point Conception, California (Lissner and Benech, 1993:figs. 4.2, 4.10) show G. eucnemis in typical array

W

with some arm branches gripping the substratum and other branches upraised. The extended arms are used

to suspension feed, presumably in the same manner as other basket stars (Kyte, 1969; Patent, 1970b; Austin

and Hadfleld, 1980; Hendter, 1982). The species is not a predator on sea pens as once claimed (MacGinitie, I

1949). Its metabolic rate has proved to be similar to simple-armed, suspension feeding ophiuroids (LaBarbera,

1982). I

126 I
!

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! Figure 7.3. Gorgonocephalus eucnemis(Miiller andTroschel):A, dorsalview (LACM42-39.4; dd= 35.0 mm);

B, ventralview (LACM65-331.1; dd = 56.0 ram).

!

I Patent (1969, 1970a,b, 1976) treats the gonadal morphology, reproductive cycle, and development

of the species. She interpreted the spawning season to last the 6 months of summer and fall in the San Juan

Islands, Washington, but her data suggest that gonadal growth occurring in the summer precedes spawning

I February. egg moderately large (0.22 mm diameter) gives a yolky,

between October and The is and rise to

non-feeding larva of unusual morphology, which metamorphoses in about a week. Early developmental

stages are associated with the alcyonacean Gersemia sp., and the larger juvenile cling to the soft coral and

I' adult basket stars.

Individuals are occasionally snagged by fishermen using hook and line (Campbell 1921a). Given

i that large basket stars can have a relatively long life-span of over 7 years (Wolfe, 1982), curio-hunting scuba divers could pose a threat to the survival of shallow water populations (Austin and Hadfield, 1980).

The populations could be slow to recover, depending on the age at reproduction and the frequency of successful

settlement.

!

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!

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i 127
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Order Ophiurida I

Family Amphiuridae i

Amphichondrius granulatus (Liitken and Mortensen, 1899) new combination

Figure 7.4 I

Amphiura granulata Liitken and Mortensen, 1899:155-156, pl. 10, figs. 1-4.

Amphipholis granulata, H.L. Clark, 1917:437.--Fe11, 1962:13, 21. H

i

Amphipholis puntarenae McClendon, 1909:44, figs. 20-21 (notAmphipholis puntarenae Liitken).

Amphichondrius granulosus Nielsen, 1932:304-6, fig. 21 a,b.--H.L. Clark, 1940:335, 337.--Bamard and •

Ziesenhenne, 1961:143 .--Hartman, 1963:85, 217 .--Boolootian and Leighton, 1966:4, 10, fig. 8.-- i

A. M. Clark, 1970:31.

Amenichondrius granulosus, Fisher (in Parker, 1964:357). i

Amphichondrius unamexici, Caso, 1983:209-212, pls.1-17.

Material Examined. California: Santa Maria Basin, Phase I Sta. BSV-042, 100 m, (2, USNM i

primary voucher); Phase ITSta. R-l, 91 m (2, CAMP voucher; 2 juvenile LACM 89-411.1).--Western

Santa Barbara Channel, Phase I Sta. BSS-094, 96 m (4, SBMNH secondary voucher).--Mexico: Gulf of

California, off Isla Espiritu Santo, R/V Velero III Sta. AHF 634-37, shallow water (1, LACM 37-35.2) -- •

Panama: Gulf of Panama, R/V Albatross Sta. 3355, 333 m, lectotype (1, USNM 19539 [as Amphiura

granulata, syntype]; herein designated lectotype).--North Pacific Ocean, R/V Albatross Sta.?, (3, USNM

19540 [asAmphiuragranulata, syntypes]).---Colombia: Off Isla Gorgona, RIV Velero lllSta. AHF 854-38, •

73-110 m (4, LACM 38-122.4).--Ecuador: Off Cabo San Francisco, R/V Velerolll Sta. AHF 850-38, 27 m l

(1, LACM 38-118.24).

Description. A moderate sized amphiurid ophiuroid. Disk diameter up to 9 mm; a specimen 6.5 i

mm dd has arms approximately 34 mm long.

Disk round, flat, covered with moderately sized imbricating scales. Scales somewhat angular,

regularly arranged, thickened at disk margin. Sharp transition between dorsal and ventral disk surfaces. 'D

Radial shields prominent, somewhat less than one-fouth disk diameter, distal halves in contact, proximal I

halves separated by wedge of one or several scales. Radial shields D-shaped in small specimens, quadrilateral

in large specimens, tapering to a point proximally, to a short straight edge distally, i

I

Jaws broad, oral gaps nearly sealed by oral papillae and teeth. Infradental papillae separated, small,

narrow, rectangular. Middle papilla flattened, ovoidal to subquadrangular. Distal papilla largest, ovoidal to

subquadrangular, with protruding, pointed, proximal adradial edge. Teeth wedge-shaped, with proximal i

edge of imperforate stereom. Oral shields roughly diamond-shaped, proximal end pointed, distal end rounded;

180

paired proximal edges longer and more concave than distal edges. Madreporite larger than oral shields,

rounded, inflated, multiperforate. Adoral shields large, maximum length equals oral shield; proximal lobes _R

i

narrow, in contact proximal to oral shield; radial lobes larger, touching fh'st ventral arm plate; adradial lobe HI

nearly right angled. i

Arms broad near disk, markedly narrowing near tip; in cross-section, dorsal surface gently curved, i

ventral surface flattened. i

Ventral interradius flattened or inflated, appearing wrinkled in living individual, covered with minute, •

angular granules. Genital plate and genital scales bare at edge of bursal slit. Parallel "furrows" on genital n

scale (noted in Nielsen, 1932) actually demarcate edges of overlapping series of scales. Bursal slit narrow,

concealed beneath arm, nearly reaching edge of disk. i

I28 I
|

Dorsal arm plates ovoidal, 2 times wider than tong, in contact at base of arm; at tip of arm separated

by lateral arm plates, fan-shaped; proximal edge slightly concave, or with medial lobe, or convex; distal

I edge convex. Lateral arm plates prominent, with wedge-shaped extension separating successive dorsal arm plates

and ventral arm plates, spine articulation ridge on distal edge. Three arm spines, with thick, hollow base,

I abr length uptlyofnarrowing joint on proximal to blunt tip, joints. slightly Dorsal curving -most distally, spine relatively rounded, longer smooth.at Spines base ofapproximately arm than at tip. equalTwoto

rounded tentacles scales, nearly covering tentacle pore, appear to be aligned with edge of ventral arm plate;

I smaller proximal scale attached to lateral arm plate, larger distal scale attached to ventral arm plate. Ventral arm plate in mouth angle bearing blunt conical protrusion extending into mouth; more

distal arm plates pentagonal, broader than wide, with slightly concave edges; outer edge medially indented,

I thickened. Plates nearly in contact at base of arm, separated distally.

I

Variation: The ventral interradial granules and bare dorsal scales that characterize the genus are

present in juvenile specimens of A. granulatus, only 1.0 - 1.2 mm dd. Individuals of that size have only a

I few disk plates and just three arm joints.

Color: Disk very dark brown or brownish-gray, larger scales surrounded by pale (connective?)

i tissue; radial shields darker than other scales, marginal scales more pale; granules of ventral interradius pale gray or whitish; jaw structures dark gray, madreporite contrasting cream or yellowish. Arms salmon, salmonbrown, or reddish brown; dorsal arm plates with indistinct medial salmon or whitish stripe and pale gray

edges; lateral arm plates brown, arm spines gray or brown, tentacle scales gray. Boolootian and Leighton

I (1966) note arms gray or mottling.

white with blackish

Pigmentation fades in alcohol preserved specimens.

I poor illustration, Remarks.unmistakably Lfitken and Mortensen's characterizes (1899) the unique description disk armament of Amphiuraof thegranulata species., They despitediscussed their rather the

possible placement ofA. granulata in Ophiocnida, a genus that is distinguished by short disk spines. Nielsen

I (1932) correspondence overlookedof their these work specieswhenwaserecting verfiedAmphichondrius by comparison granulosus of Nielsen's, newdefinitive genus illustrations and species. with The

specimens identified by Liitken and Mortensen. Its oral formula and disk granulation exclude the species

, Amphichondrius from Amphiura asis appropriate. currently defined Therefore, (see A. Araphichondrius M. Clark, 1970), granulosus and its Nielsen placementis regarded in Nielsen's as a junior genus

synonym ofAmphiura granulata Lfitken and Mortensen, and a new combination is proposed,Amphichondrius

i granulatusThe(Lfitken and Mortensen). specimen upon which Lfitken and Mortensen based their description, which lacks primary

plates, matches the syntype catalogued as USNM 19539, Albatross Sta. 3355, Gulf of Panama, 182 fathoms

I (later proposelabelled: USNMPanama, 19539 asAzuero the lectotype Peninsula, of Amphiura 7° 12' 20"Ngranulata 80° 55' 00"W, L/itken333andms, Mortensen. 23 Feb 1891). Another Therefore, lot, withI

three specimens catalogued as USNM 19540, Albatross 1891 (later labelled: Type, between Mexico and

I: Galapagos) numbered 8158. may correspond to the second lot examined by Lfitken and Mortensen, which was originally

Ziesenhenne (1940) described a second species in the genus, Amphichondrius laevis, also from

I Eastern Pacific waters. He distinguished it from A. granulatus based on several ambiguous characteristics including the primary plates, lateral arm plates, and oral shields. Reexamination of Ziesenlaenne's material

in the Allan Hancock collection of LACM, including the types, shows that A. laevis is a small species,

I generally capped byless a large than 4mid-marginal mm dd; its disk scaleisapproximately pentagonal withtwice pointed as wide extensions as longjutting (ratherbetween than thrice the arms as inandA.

granulatus). The dorsal arm plates ofA. laevis are narrower and more triangular than those ofA. granulatus.

!

I 129

I A. laevis has minute pointed granules covering the edges of the bursal slits; granules are confined to the

interradii in A. granulatus.

I 1 A third nominal Eastern Pacific congener, Amphichondrius unamexi Caso, is regarded as a synonym

ofA. granulata. A paratype ofA. unamexi (UNAM, Mexico City, Acc. No. 2368), provided by F.A. SolisMarin, was not distinguishable from typical specimens ofA. granulatus.

I and Mortensen, Distributi1899; on. Pt. H.L.Conception, Clark, 1917,California, 1940; Barnard to Mexico, and Ziesenhenne, and Panama,1961; at depths Hartman, of 18-384 1963).m (Liitken

Biology. Barnard and Ziesenhenne (1961) estimated fewer than 0.4 specimenslm 2for 176 stations

I between Pt. Conception and the Mexican border at 9-183 m. A few living individuals that I have examined behaved sluggishly, and their tube feet lacked microscopic papillae. These traits suggest that the species is

a relatively inactive deposit feeder. Specimens are gonochoric, and those as small as 4.5 mm dd have

I recognizable gonads.

] t_ Amphiodia (Amphispina) digitata Nielsen, 1932

1 Figure 7.5

Nielsen, 1932:275,277-279, 12a-c.--A. M. Clark, 1970:25.--Sutton, Amphiodia (Amphispina) digitata fig.

1976:347, 350-352•

l Amphiodia digitata, Barnard and Ziesenhenne, 1961:143.--Parker, 1964:359.--Boolootian and Leighton, 1966:5, 8, fig. 11.--Hartman, 1965:85, 154, 200, 217, 237, 251,285•

Amphiodia barbarae (Lyman), McClendon, 1909:45, figs. 22, 23 (in part).

I Material Examined. California: Santa Mafia Basin, Phase I Sta. BSR-073, 98 m (1, USNM primary

voucher [as Amphiodia occidentalis]); Phase II Sta. R-l, 91 m (1, CAMP voucher [as Amphiodia urtica]);

Phase II Sta. R-1 (1, CAMP voucher).---Off Santa Cruz Id., R/V Velero III Sta. AHF 1005-39, 59-62 m (1,

l LACM 39-109.11).---Off Santa Barbara Id., Velero III Sta. 978-39, m (1, 39-82.7).

R/V AHF 38-51 LACM

--Off San Nicolas Id., R/V Velero lllSta. AHF 1219-40, 27-40 m (2, LACM 40-194.7).--Off La Jolla, Dr.

Th. Mortensen's Pacific Expedition, 165-183 m, syntypes (ca. 60, Zoological Museum, University of

I Copenhagen); 110 m, syntype (1, ZUUC).

Description. A small amphiurid growing to 7 mm dd, with long, slender arms.

I interradius; Diskdiskround, surfacslightl e uneven, y inflraised ated, along with coarse edge ofsradial caling;shields. less thanRadial 10-12shields columns in contact of scales most ofin length, dorsal

separated by 1-2 scales proximally; shields more than twice as long as wide, approximately one-fourth disk

i diameter hyaline points, in length. toothedUppermost scales forming row of continuous ventral disksescales ries between with freeradial edgeshields. expanded into into 2-3 (up to 6)

Jaws moderately wide. Oral gap narrow, oral papillae coveting most of slit. Infradental papillae

I close; madreporite two lateral papillae flattened, with expanded free edge, distal papillae largest. Oral shields pentagonal;

larger, inflated, generally possessing 1pore. Adoral shields in contact, or nearly so, proximal to

oral shield; radial lobe adjoining first lateral ann plate, inserting between first two ventral arm plates.

i Scales covering ventral interradius smaller than those on dorsal disk surface. Bursal slit spanning oral shield, reaching nearly to disk margin; some scales at edge of slit with free edge produced into pointed

projection.

I Arms distal of in cross-section rounded flattened slender, portion arln gradually tapering; dorsally,

ventrally.

I

i 131
i

Lateral arm plates with prominent spine-artculafion ridge on distal edge, bearing 3 spines, sharply •

pointed, of about equal length, 1-1.5 times length of ann joint. Spines, most markedly middle spine, distinctly

wider at base than at tip, curving slightly towards ann tip; dorsal and ventral spines laterally compressed, I

middle spine dorso-ventrally compressed. Two uppermost arm spines on joints below disk with sharp, I

proximally curving, hyaline tip.

Dorsal arm plates subrectangular, somewhat wider distally, generally with slightly concave distal I

margin. Soft tissue between joints not readily visible in dorsal aspect. II

Ventral arm plates pentagonal, nearly in contact, long as wide, widest across proximal edge, distal

edge with medial notch. First ventral arm plate U-shaped, set at distal angle of oral slit. Two tentacle scales i

at right angles, in contact; scale on ventral arm plate markedly larger, with broad free edge.

Color of living individuals: Disk gray to dark grayish brown tinged with purple; scales grayish- I

brown edged with gray or blackish; radial shields brown with yellowish-brown distal edge; ventral interradii I

pale gray. Arms reddish with irregular gray blotches or irregularly banded with gray and pale gray or cream,

m

with indistinct, discontinuous, medial, orange-red stripe; ventral surface gray or brown. Eggs and ovaries

orange-red, or salmon colored. I

orange,

I

Specimens in alcohol fade; Nielsen (1932) described them as whitish-yellow.

Remarks. A. digitata and its sympatric congener, A. urtica, are similar but distinct species. Their II

identification has been clouded by the appearance of seemingly "intermediate" individuals, generally with II

regenerating disks and arms, lacking the characteristics of perfect specimens. In addition, consistent

differences between the juvenile stages of the two species have not been determined. The misidentification •

of specimens made during preliminary taxonomic analysis in the present survey is typical. An examination l

of earlier Allan Hancock Foundation material in LACMNH indicates the two species were confused in

previous surveys of Southern California waters. (Bergen, 1995; pers. obs.). Characteristics distinguishing /I

A. digitata are: I

1) relatively few, large, dorsal disk scales on specimens with primary plates

2) uppermost row of ventral disk scales with some free edges expanded into into 2-3 (up to 6) 'I

prominent, widely diverging, hyaline points. The modified scales form a prominent, nearly continuous I

series between the arms. They are aligned with smaller hyaline-toothed scales that border the bursal slits. A.

urtica has ventral interradii with roughened spiny scales, generally with a single hyaline point, which are i

most evident along the genital slit and at the base of the arms

3) relatively rectangular dorsal arm plates with lateral edges that conceal the tissue-f'llled gaps I

between successive joints. A. urtica has more ovoidal dorsal arm plates, between which wedges of soft i

tissue are apparent. II

4) disk coloration grayish in A. digitata, reddish in A. urtica. _i

Sutton's (1976) contention thatAmphiodia digitata is a junior synonym ofAmphiodia (Amphispina) I

periercta H.L. Clark, 1911 is rejected based on a study of Eastern Pacific Amphiodia (Amphispina) species

(l-Iendler and Wang, in prep.).

Distribution. Point Conception, California to Mexico, at depths 18-280 m (Nielsen, 1932; Barnard I[

and Ziesenhenne, 1961; Hartman, 1963; Parker, 1964).

Biology. Barnard and Ziesenhenne (1961) reported the species from 6% of 176 stations at 9-183 m I

depths between Pt. Conception and the Mexican border, with an overall average density of 7.5 individuals/

m2.Specimens collected in Santa Monica Bay, California in October, 1991 and December, 1992 had eggs

that appeared to be ripe, with maximum diameters of 0.16 mm. This is a somewhat larger gamete than !

typical for species with ophiopluteus larvae (Hendler, 1991), perhaps an indication that the larva has an

abbreviated mode of development.

I

132 i

!

n

Amphiodia(Amphispina) urtica(Lyman, 1860)

Figure 7.6

I

Amphiura urtica Lyman, 1860:195-196.--1865:12, 115, 128-129.--1880:21.--1882:125, 147, 311.

? Amphiura urtica, H.L. Clark, 1901:330 (not A. urtica Lyman). i

Amphipholis urtica, Ljungman, 1867:315. l

Amphiodia urtica, Verrill, 1867:325.--1899:313.--H.L. Clark, 1911:11, 154, fig. 64a-c.--1915:249.--

Campbell 1921b:47.--May, 1924:288-89, fig. l la-e.--Berkeley, 1927:4-5.--Ziesenhenne, •

1937:224.--1940:331,338.--Barnard and Ziesenhenne, 1961:133-151.--Hartman, 1963: 85, 137, n

153, 170, 185, 201,217, 237, 251,285, 305, 334, 349, 365, 382, 400.--Boolootian and Leighton,

1966:5, 8, fig. 12.--Kyte, 1969:1729, 1734-1736, t738.--A. M. Clark, 1970:26.--Austin and i

Hadfield 1980:150-151, pl. 49, fig. 10.5. n

? Ophiophragmus urtica, Fell, 1962:14, 26.

Amphiura barbarae Lyman, 1875:5, 17, pl. 3, figs. 32-34.--McClendon, 1909:45, figs. 22-23 (in part).-- g

H.L. Clark, 1915:245. Campbell, 1921a:3, fig. 4..--1922b:47.--1922:38, pl. 3, figs. a-f (in part?).

Amphiodia Barbarae, Lyman, 1880:21.--Verrill, 1899:313. •

i Amphiura (Section Amphiodia) barbarae, Koehler, 1907:306.

Diamphiodia barbarae, Fell 1962:14, 19. il

Material Examined. California: Santa Mafia Basin Phase I Sta. BSR-023, 195 m (1, SBMNH

secondary voucher); Phase I Sta. BSR-073, 98 m (20, USNM primary voucher; 35, SBMNH secondary

voucher [asAmphiodia occidentalis]); Phase II Sta. R-I, 91 m (78, CAMP voucher); Phase II Sta. R-l, 91 m i

(52+, CAMP voucher [asAmphiodia sp.]).--Westem Santa Barbara Channel, Phase I Sta. BSR-085, 113 m

(92, USNM primary voucher).---Off Pt. Loma Lt., R/V Velero IV Sta. AHF 5200-57, 72 m (numerous,

LACM 57-240.3). I Description. A small amphiurid, reaching 9 mm dd, with arms 12 times as tong. Disk rounded to

pentagonal, inflated, covered with delicate, minute, imbricating scales. Primary plates and scales surrounding

radial shields larger than other disk scales. Radial shields approximately 3 times longer than broad, length n

approximately one-fifth disk diameter, in contact most of length, proximal tips separated by small wedge of R

scales.

Jaws narrow, oral gap wide. Three oral papillae; infradentals blocklike, separated, middle papilla i

smallest, outer papillae largest; both lateral papillae flattened, rounded triangular shape, with point of

m

attachment narrow, free edge expanded. i

Oral shields pentagonal to rhombic, about as long as wide; proximal end pointed, distal end broadly i

rounded. Madreporite almost twice size of oral shield, thick, inflated, with 1 or more pores. Adoral shields

with rounded lobes in contact proximal to oral shield; pointed radial lobes in contact with fLrstlateral arm •

plate, inserting between first two ventral arm plates. Ventral interradius finely scaled; scales generally quite !

small and sparse near oral shield. Bursal slit conspicuous, extending to edge of disk. Scales near genital slit

enlarged, generally beating single, minute, hyaline teeth. Series of spinose scales commonly extending to n

distal comer of radial shield, sometimes sparsely distributed across the lower lateral edge of the disk between

the arms.

U

Arms slender, broader than high; dorsal and ventral surfaces gently rounded. Characteristic gaps []

between arm joints filled by wedge of soft tissue; dorsal edge of the vertebra visible within. i

Dorsal arm plates ovoidal, with convex proximal edge, lateral edges lobed, distal edge with broad

medial covexity; successive dorsal arm plates barely in contact or slightly overlapping, n

134 i
!

Lateral arm plates thin, with distally protruding spine-bearing ridge. Three arm spines of about

equal length, rounded in cross section, with thick base, tapering to acute point; dorsal and ventral spines

slightly several armlaterally joints compressed, beneath diskmiddle with proximally spines slightly curved,dorso-ventrally sharply pointed, compressed. hyaline tip.Dorsal Two tentacle spines onscales, first

thin, ovoidal; scales covering little of tentacle pore; scale on ventral arm plate larger, longer, often bi-lobed.

I medio-distal Ventral edges, arm slightly plates pentagonal separated with by interdigitating concave edges, lateral forming arm 2plates. postero-lateral Proximal lobes ventralandarmindented plates

about as long as wide, distal plates longer than wide. First ventral arm plate U-shaped, set at distal angle of

oral slit.Color: Disk reddish to grayish-brown, primary plates contrastingly dark, radial shields gray with

pale tips. Arms reddish gray, with irregular gray or brownish blotches, reddish medial stripe; dorsal arm

I plates and eggs graypink, with pinkish gray border, ventral arm plates yellowish brown, arm spines pinkish gray. Ovaries testes whitish.

In alcohol disk and arms fade to pale yellowish brown.

I Remarks. The subgenus Amphispina Nielsen is characterized by the presence of scales with spiny

margins among the uppermost row of ventral disk scales. Although mention was omitted from the original

i description, there are commonly spinose scales along the bursal slits of the type species, Amphiodia (Amphispina) digitata. A. urtica has spinose scales on the disk margin, mostly occuring near the arms, and

as noted by A.M. Clark, individuals exhibit varying densities of spinose scales on the ventral interradius and

along the bursal slit. Therefore, I do not concur with A. M. Clark's (1970) exclusion of A. urtica from

I Amphispina, retained the species in the subgenus as originally proposed by Nielsen (1932), together

and have

with A. digitata, A. duplicatus (Koehler), and A. microplax Burfield.

i m, and possibly Distribution. as deepMexico as 708 mand(H.L. Southern Clark, California 1911,1915,to1940; BritishBerkeley, Columbia1927; andNielsen, Alaska, 1932; at depths Ziesenhenne, of 9-198

1937; May, 1924; Barnard and Ziesenhenne, 1961; Hamnan, 1963).

I shelf andBoffshore iology. Thebasins species (Hartman, has proved 1955,to1966; be the Hartman most numerous and Bamard ophiuroid1958, of Southern 1960; Jones, Calif1969). ornia mainland Based on

176 stations at 9-243 m depths between Pt. Conception and the Mexican border, Bamard and Ziesenhenne

i (1961:133) coastal shelves, characterized occurring Ain. urtica 86% ofasall"...the samples mostandabundant havig and a frequency widespreadfor animalll depthsin any of 422phylumanimalson per the

square meter. It forms the dominant in the two most widely distributed communities of the coastal shelf."

i They with Amphiura found that it is most numerous in sandy silts that feel sticky to the touch, and that it frequently occurs arcystata and in a community dominated by Amphioplus hexacanthus (=Dougaloplus

amphacanthus). Bergen (1995) concluded that A. urtica is most abundant in sediments with a median grain

i size of 0.035-0.093 mm (coarse silt to very fine sand) at depths 15-85 m. The species is similarly abundant in Puget Sound, Washington, where it appears to have a life span of at least five years (Lie, 1968).

Juvenile individuals commonly comprise a major component of the population, as they did in the

i collection under study. The many small specimens examined can be identified with certainty only to genus, because they lack the features of taxonomic importance used to characterize the adults. Therefore, many

specimens from the Atlas survey that are probably immature A. urtica are not listed above with other

material examined.

I In southern California, A. urtica is routinely collected in studies assessing the impact of wastewater

pollution on benthic communities. It is considered an indicator species, and is frequently cited in the extensive

i gray coveredliterature of governmental agencies dealing with wastewater management. Some of that literature is in a recent symposium proceedings (see: Diener and Fuller (1995), Dorsey et al. (1995), Scanland

(1995)). It is generally thought that populations of A. urtica are diminished near wastewater outfalls, and

i recover when pollution is abated (Dorsey et al., 1995; Scanland, 1995). However, it has also been suggested that population density may be held in check by predatory fish which congregate near outfalls (Diener and

Fuller, 1995).

i 135
I

II

II

I

i

II

,I

i

l

I

Figutre 7.6. Amphiodia (Amphispina) urtica (Lyman): A, disk, dorsal view; B, disk, ventralview; C, disk edge,

showing spinosescales ondisk margin (LACM 56-66.7; dd =4.9 ram). I

The available literature treats the density and distribution of the species and its response to polluted i

sediment. However, its life history, behavior, and ecology are poorly known. Austin and Hadfield (1980)

summarize the limited information on its biology, estimating densities up to 1,500 individuals/m 2,pointing I

out that it burrows, suspension and deposit feeds, and is capable of arm and disk autotomy and regeneration.

The species is gonochoric. Its eggs are approximately 0.12 mm diameter, the size indicating development as

gravel a feeding, andophiopluteus coarse larva (Hendler, 199l). Kyte (1969) reports the young ofA. urtica occur under loose Isand. In British Columbia, its predators include lemon sole, flathead sole, sand sole, and

the sea star, LuidiafolioIata. Nielsen (1932) notes that individuals are frequently infested with a copepod

which he identified as a CanceriIla species, g

!

i

!

136 i
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I Amphioplus (Amphioplus) strongyloplax (H. L. Clark, 1911)

I Figures 7.7

Amphiodia strongyloplax H. L. Clark, 1911: 10, 164-165, fig. 71a-c.--1915:249.---Campbell, 1921b:47.--

Sutton, 1976:349.

I Amphioplus strongyloplax, Nielsen, 1931:295-296, fig. 18a-b.--Barnard and Ziesenhenne, 1961:143.--

Hartman, 1963:85, 125, 154, 170, 185,217, 251,285,305,334, 350, 365, 382.--Astrahantseff and

i Alton, 1965:1414, 1416, 1420.--Booloofian and Leighton, 1966:5, 9, fig. 16.--Alton, 1972:507, 510.

Unioplus strongyloplax, Fell 1962:16, 25.--Kyte, 1969:1729, 1733, 1735, 1737.

I Amphioplus (Amphioplus) strongyloplax, A.M. Clark, 1970:20, 44, 57, 77, fig. 9b.

Material Examined. Washington, Cape Flattery, RIVAlbatross Sta. 2866, 313 m, holotype (USNM

i 25615 (1+, LACM[asAmphiod 64-242.iaI).--California: strongylopIax]).--Orego Santa Mafia n: Basin, Off ColPhase umbiaI Sta. RiverBSV-042, mouth, R100/V Commando m (1, SBMNHSta.secondary ?, 400 m

voucher); Phase II Sta. R-5, 154 m (1, CAMP voucher [asAmphiolpus macraspis]); Phase II Sta. R-I, 91 m

I (3, Sta.CAMP PJ- 11, voucher); 136 m (2, LACM Phase II86-492.9).--Western Sta. PJ-4, 150 m (1, CAMP Santa Barbara voucherChannel, [as AmphioPhase plusI Sta. hexacan BSS-084, thus]); 394Phase m (1,II

USNM primary voucher).---Off San Miguel Id., R/V Velero IVSta. AHF 23165-75, 96 m (1, LACM 75-

I 409.4 off El).---Off Segundo, SantaR Rosa Id., R/V Velero 111Sta. AHF 993-39,229-238 m (1, LACM 39-97.4). - Los Angeles, /V Velero IV Sta. AHF 3400-55, 400 m (4, LACM 55-110.I).----Off San Pedro, R/V

Velero IH Sta. AHF 1228-41,230-252 m (2, LACM 41-8.3).

I specimenDeofscri5.5ption. mm disk An diameter amphiuridestimated of modertoatehavesize,armsthe50maximum mm longdisk (H. L.diameter Clark, 1911), exceedone ing 810mmmm; diska

diameter has arms over 95 mm long; judging from the fragmented specimens of the present survey, arm

I length probably exceeds 12 times disk diameter. Disk soft, flat, indented interradially. Disk coveting variable, ranging from scales restricted to region

surrounding the radial shields, to completely coating entire disk. Scales irregularly arranged, sometimes

I multi-layered, smallest at center of disk, largest near radial shields, those below disk drastically reduced in size and number. Primary plates discernible on all but regenerating disks. Radial shields diverging and

tapering proximally, nearly in contact distally, with radial edge straight, abradial edge convex; length

I aregenerating pproximatelydisks. 4-5 times width, approximately one-fourth disk diameter, relatively larger or smaller on

Jaws narrow, ventrally projecting, with deep indentation distal to apex; oral gap pronounced, widening

I distally. Generally 5 oral papillae; infradental papillae touching or nearly so, small, blocklike, with protruding proximo-radial comer; buccal scale sometimes equal in size to lateral papilla, high in jaw, triangular or

spinelike; two lateral papillae flattened, triangular to blunt spinelike, outermost usually largest; distal-most

i minute, set between first ventral arm plate and adoral shield. Teeth chisel shaped; proximal tip irregular or

straight, composed of imperforate stereom.

Oral shields variable, subpentagonal, diamond-shaped to irregularly ovoidal; somewhat longer than

I wide; madreporite thicker, larger, more rounded, usually with single perforation. Oral shield overlapping little of adoral shield. Adoral shields L-shaped; proximal lobes of adjacent plates touching or nearly so;

distal lobes larger, inserting between first and second ventral arm plates. Ventral interradii with extremely

I small thin scales andirregular granules separated by integument; larger flat scales on abradial edge of bursal slit. Bursal slit extending from oral shield to disk edge.

Arms long, slender, narrowing near disk, wider at midlength, tapering distally, gently rounded

I dorsally, ventrally.

flat

l 137

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1

i Figure 7.8. Amplu'optus(Amphioplus) macraspis (H.L. Clark): A, disk, dorsal view (USNM 25584, syntype,

Amphiodia macraspis; dd = 11.5 mm); B, disk, ventral view; C, oral frame, showing broad oral

I shields(USNM 25584, syntype,Amphiodia macraspis; dd = 8.5mm).

t Dorsal arm plates thin, little broader than long, ovoidal to hexagonal with short lateral edges, smaller than joint, separated by lateral ann plates and wedges of soft tissue between joints. First few basal plates

reduced in size.

I Lateral arm plates with prominent spine-bearing ridge. Five arm spines, occasionally 6 on basal joints, gradually tapering, acutely pointed, smooth, delicate, hollow with wall of spine much thinner than

internal cavity. Spines beneath disk equal or slightly exceeding length of ventral arm plate. On joints with 5

i spines dorsal-most and ventral-most spines longest, middle spine shortest, all slightly compressed, gradually tapering, pointed. On joints with 4 and 3 spines near middle of arm, spines increasing in length ventrally,

second spine from bottom noticeably broadened at base, dorso-ventrally compressed; spines near arm tip

i very slender, acute.

First ventral arm plate set in oral slit, with distal edge broadened; second ventral arm plate hexagonal.

Those plates on proximal joints slightly longer than broad, with rounded posterolateral lobes, generally with

I plates contact or nearly so; arm plates separated by

distal medial indentation. Several basal in distal ventral

lateral arm plates. Tentacle scales 1-2, minute, embedded in soft tissue, covering little of large tentacle pore.

i Variation: Number of arm spines is size related: at 2.0 mm dd (off 1.8) there are 3 arm spines, at 2.3 mm dd (off 2.0) 4 arm spines, at 7.6 mm dd (off 3.2 mm) 5 arm spines. Likewise, number of tentacle scales

size is related: some joints of small specimens lack tentacle scales; a few joints of large specimens have 2

tentacle scales.

i Color: Alcohol preserved specimen with regenerated disk has pale scales and blotches of yellow

and brown; radial shields gray, yellow, and white; dorsal and ventral arm plates marked with orange to

I yellow-brown some arm spines, pigmentation, oral and adoral forming shields, discontinuous jaws, and radial medial shields. stripe. According Orange-yellow to H. L. pigmentation Clark (1911), present specimenson

dried from alcohol with "disk grayish, arms very light brownish; radial shields and arm plates with orangei brown tint."

' I 139
!

Remarks. H. L. Clark (1911 ) noted the similarity ofA. strongyloplax to Amphioplus (AmphiopIus) i

macraspis (compare Figures 7.7 and 7.8). Comparison of type material indicates A. macraspis (USNM

25584, syntype) differs in having the ventral arm plates broadly in contact, broad oral shields, and relatively Ill

wide dorsal arm plates. A. strongyloplax has been described as resembling Amphioplus hexacanthus. The II

latter is discussed below, and synonymized with Dougaloplus amphacanthus.

Distribution. Mexican border and Southern California to British Columbia; depth range 91-623 m; Ii

a doubtful record from 4 m (H.L. Clark, 1911, 1915; Nielsen, 1932; Bamard and Ziesenhenne, 1961;

Astrahantseff and Alton, 1965; Alton, 1972; Kyte, 1969).

Biology. Bamard and Ziesenhenne (1961) reported this species from 23% of 176 stations taken 1

between 9-243 m depths from Pt. Conception, California to the Mexican border. They calculated an overall m

population density of 4.3 individuals/m 2,but Astrahantseff and Alton (1965) provide a more realistic estimate

of 60 individualslm 2 off the Oregon coast. The species occurs in mud with varying amounts of sand and 1

gravel (Kyte, 1969). It certainly burrows in soft sediment, presumably the reason that Alton (1972) found it U

is taken with infannal sampling gear, never with trawls. A specimen examined in the present survey (Phase

I Sta. PJ-11) was a female with eggs of 0.18 mm diameter, an oocyte size indicative of an ophiopluteus 1

larval form or an abbreviated mode of development (Hendler, 1991). II

Amphipholis pugetana (Lyman, 1860) I

Figure 7.9

sit

Amphiurapugetana Lyman, 1860:193-194.--1865:12, 15,115,125-126.--1880:19.--1882:125, 145, 311.-- !

Koehler, 1907:305.

Amphipholis Pugetana, Verrill, !865:325.--1899:312. i

I

Amphipholis pugetana, Ljungman, 1867:312.--McClendon, 1909:43-44, figs. 12, 13 (in part?).--H.L. Clark,

1910:323, 340-341, pl. 9, fig. 2.--H.L. Clark, 1911:166-67, fig. 73a-c.--1915:242.--Matsumoto,

1917:185,191,358, 365, fig. 51.----Campbell, 1921b:48.--May 1924:289-290, fig.12a-c.--Berkeley, B

1927:5, fig. 12.--Nielsen, 1932:281,288-90.--D'yakonov, 1954:58.--Barnard and Ziesenhenne,

1961:143.--Fe11, 1962:13, 24.--Hartman, 1963: 85, 137, 154, 201,217, 286, 320, 335, 365, 383,

391,401.--Boolootian and Leighton, 1966: 5, 9, fig. 9.--Austin and Hadfield 1980:148-149.-- II

Smimov, 1982:113. J

Amphiopholis pugetana, Bush, 1921:72. I1

Axiognathus pugetana, Kyte, 1969:1728, 1729, 1735-1737. I

Material Examined. Oregon: Off Columbia River mouth, R/V Commando, 900 m (1, LACM 64- m

241.2). - California: Santa Maria Basin, Phase I Sta. BRA-013, 92-100 m (58, USNM primary voucher [as 1

Amphipholis squamata]); Phase I Sta. BRA-016, 91.5-123 m (36, USNM primary voucher [as Amphipholis

w

squamata]); Phase 1ISta. R-4, 92 m (3, CAMP voucher [as Ophiura luetkeni]).--Off Santa Catalina Id., R/V ,...,

Velero/V Sta. AHF 1989-50, 604 m (1, LACM 50-45.2); RN Velero/V Sta. AHF 2425-53, 64-70 m (8, i[

LACM 53-159.1).--Off San Clemente Id., R/V Velero IV Sta. AHF 22943-75, 319 m (9, LACM 75-297.2). u

Description. A small amphiurid, with maximum disk diameter 5 ram; an individual 3.5 mm dd has I

arms 25 mm long. |

Disk round to rounded pentagonal, inflated, with radial shields appearing slightly sunken; disk

margin sharply demarcated by broad ventral scales overlapping elongated dorsal scales. Disk scales i

conspicuous, rounded, imbricating, with opaque, thickened outer edge. Primary plates forming rosette I!

occupying central one-third of disk. Radial shields three times longer than wide, length approximately onefifth diameter of disk; radial edges straight, adradial edges curved; proximal end pointed, distal end short, I

straight. Paired shields in contact, except proximal ends separated by 1-2 small scales. |

,40 |
!

t Jaws broad. Oral gap narrowed by oral papillae. Infradental papillae rounded, blocklike, separated.

Middle papilla rounded, quadrangular, breadth increasing from attachment point to free edge. Distal papilla

i largest, quadrangular, Oral elongate, narrowing distally, overlapping fn'st ventral arm plate. shields diamond to spade-shaped, corners rounded, about as long as wide; proximal portion

elongated and pointed, edges concave; distal corner abutting genital scale. Madreporite larger than oral

I shields, shields; inflated, paired plates beatingin contact. one pore.Ventral Adoralinterradii shields quadilateral, inflated, covered usuallywithlarger small thanimbricating and extending scales. above Genital oral

scales frequently with proximal ends touching or nearly so, short axis aligned dorso-ventrally. Bursal slit

i partly hidden by arms, nearly reaching edge of disk. Arms generally 6 times disk diameter, slightly increasing in width to a distance of approximately 10

joints from disk, then gradually tapering. In cross-section dorsal surface somewhat rounded, ventral surface

I nearly Terminal fiat.plate Jointsgradually with lateral edges diverging gradually, distal segments not sharply knotted in appearance. tapering, with truncate distal tip.

Dorsal arm plates barely in contact, or separated by lateral arm plates; proximal edge convex,

, widely diverging lateral edges, lateral comers rounded, posterior edge slightly convex.

Lateral ann plates prominent, broad, with pronounced spine attachment ridge. Lateral plates forming

i tight wedge between dorsal arm plates, meeting conspicuously between ventral arm plates. Arm spines slender, tapering gradually, smooth, lacking microscopic grooves. Spines equalling or exceeding length of

arm joint, gradually tapering to blunt tip, dorsal and ventral spines slightly laterally compressed and curved

distally, middle spine slightly dorso-ventrally compressed. Spines frequently 4 in number for one to several

I disk. When 4 in in when 3 in number middle joints beyond number, increasing length ventrally; spine

generally largest and longest. Beginning 5 to 15 joints from edge of disk, on widest portion of arm, middle

arm spine markedly enlarged, length at least equalling width of the dorsal arm plate on that joint. Enlarged

t spines 1.5 times length of joint, somewhat 8-pin with bulbous base and inflated Tentacle scales

shaped, tip.

paired, covering tentacle pore; scale on lateral arm plate with pointed tip; scale on ventral arm plate similar

or larger, elongate or teardrop-shaped.

I First ventral arm plate flat, pentagonal, overlapped by adoral shield and distal oral papillae; proximal

few ventral arm plates in contact, equilateral, with lateral edges narrowing adjacent to tentacle pore, width

across distal lobes no greater than across proximal lobes; distal plates longer than wide, regularly pentagonal,

i separated by lateral arm plates.

Color: Disk gray or orange-red and gray, darker than arms. Arms dorsally blotched with orange and

I reddish-orange, arms orange colored, with greenish pale brown, and gray, or gray. or pale Austinbrown. and ArmHadfield spines pale, oral frame and ventral surface of (1980) describe the species as "gray or

banded gray and white." Nielsen (1932) notes alcohol preserved specimens are "colorless" with the disk

I reddish inRemarks. a few specimens. In the region covered by this report only two Amphipholis species occur, A. pugetana

and the cospmopolitan A. squamata. D'yakonov (1954:58) regarded A. pugetana as "probably identical

I with unconvincing, the ubiquitous as are the A. squamata accounts "of most findingauthors "no tangible cited in thedifferences partial synonymy between above. the two." My reexamination His data are

of specimens from this and previous coastal surveys represented in the LACMNH indicated that many

i existing H.L. identifications Clark and records for the two species are incorrect. (1911: 167) claimed that "characteristic mouth parts" easily distinguish A. pugetana -

the onlyAmphipholis species represented in the Albatross collection. He provided the first illustration of the

I species, separatedAshowing , pugetana spade-shaped fromAmphipholis oral platesspecies and the of themiddle tropical armEastern spine longer Pacific than by itsothers. numberNielsen of arm (1932) spines

and rectangular distal mouth papilla. He distinguished it from Amphipholis squamata by the presence of

I arm spines as long as 1.5 arm joints (noting that spine length equal to that of the joint in A. squamata), and

I 141
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Figure 7.9. Amphipholispugetana (Lyman): A, disk, dorsal view; B, disk, ventral view (LACM 75-297.2; dd = I

4.t mm).

i

pointed out the middle arm spine exceeds the dorsal and ventral spines on some part of arm, and in large

specimens usually becomes thickened at the proximal and distal ends on 10-15 arm joints near middle of

arm. Furthermore, the arm length is 7-8 times the disk diameter (compared to 4 times in A. squamata), and m

the oral shield is narrower than tong (while inA. squamata the length equals width). Nielsen recognized that i

A. squamata broods its young internally, unlike A. pugetana. Later authors have not improved on the

distinctions drawn by Nielsen, and we still are unable to distinguish small and sexually immature specimens wl

of the two species. m

Distribution. Peru (record highly doubtful) and Southern California to Washington, British i

Columbia, and Alaska, at depths from 15-604 m (H. L. Clark, 1910, 1911, 1915; Berkeley, 1927; this

report), and possibly to 1,620 m (Hartman, 1963). Published records suggest it is amphi-Pacific (Matsumoto, aim

1917; Smirnov, 1982; Irimura et al. 1995). See distribution for A. squamata below. •

Biology. Barnard and Ziesenhenne (1961) recognized this species at 1.7% of 176 stations taken

between 9-243 m depths from Pt. Conception, California to the Mexican border. They calculated an overall ti

population density of 2.2 individuals/m 2. However, my examination of samples still at LACMNH from |

their survey indicated that specimens ofA. pugetana andA. squamata were frequently confused. Thompson

and Jones (1987) reported both species from the lower slopes of Southern California offshore basins, with m

densities of A. pugetana up m 45 individualslm 2. |

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142 I
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I Amphipholis squamata (Delle Chiaje, 1828)

Figure 7.I0

I

Asterias squae_ta Delle Chiaje, 1828:74, 77, 79, pl. 34, figs. 1-4.

i Amphiura Ophiolepis squamata squamata, , Liitken Mtiller 1859:226-227.--Lyman, and Troschel, 1842:92. 1865: t 2, 14, 115, 121-123.-- 1875:5, 16.--Lyman,

1880:19.--1882:125, 136, 291,296, 309-310, 314, 321-322, 324.--Ives, 1889: 177.--Koehler,

t Amph 1907:305. ipholis squamata, Verrill, 1899:312.--H.L. Clark, 1910:323, 340-341 .----Clark,1915:242.--1923:154-

155.--1940:331,337.--Mortensen, 1927:221, fig. 125.--Nielsen, 1932:281,290.--Ziesenhenne,

i 1937: 1961:143.--Fe11, 223-224.--A.H. 1962: 13, Clark, 25.--Hartman, 1939:2.--D'1963: yakonov, 85, 137,1954:57.-- 154, 185,B200, arnard218, 237, and 251,285,305, Ziesenhenne,

334, 349, 366, 382, 401.--Boolootian and Leighton, 1966: 5, 9, fig. 10.--A. M. Clark, 1970: 7, 30-

I 31, fig. lg.--Austin and Hadfield 1980:148-149, pl. 49, fig. 10.3. Axiognathus squamata, Thomas, 1966:831-832.--Kyte, 1969: 1729, 1733, 1735-1737.

I Material Examined. California: Santa Maria Basin, Phase IISta. R-4, 92 m (11, CAMP voucher); (1 juvenile ?, CAMP voucher [as Amphiodia sp.]); Phase 1/Sta. R-4, 92 m (6, CAMP voucher [as Ophiura

luetkeni]).--Los Angeles, off Portuguese Bend, R/V Velero 1II Sta. AHF-39, intertidal (31, LACM 39-

I Laguna 117.13).---Off Beach, Newp intertidal ort B(14, each,LACMR/V 57-278.1). Velero III Sta. AHF 1225-41, intertidal (4, LACM 41-5.12). - Off

Description. In many respects the description A. pugetana applies toA. squamata. The structure of

l the arm spines is the most reliable differential character inadult specimens. Other features distinguishing A.

squamata are the following:

Dorsal disk scales smooth, flat, edges not thickened.

I Distal oral papillae with flaring, rounded outer edges meeting to seal oral gap.

Oral shields more pentagonal than diamond-shaped; distal edge broad or lobate. Oral shield larger

I than shieldadoral overlapping shield, greatest and separating width ofproximal oral shielend ds exofceegenital ding greatest scales. width of adoral shield. Distal end of oral

Arms short, up to 4 times disk diameter. Distal-most arm segments appearing knotted due to proximal

I constriction of lateral arm plates. Basal dorsal arm plates of equal length and width.

I Later Ventral al armarmplates plates with pentagonal, expanded, widthrounacross ded, spine distal alobes ttachment generally ridge.exceeding that across proximal

lobes.

i Terminal arm plate with bulbous base, abruptly narrowing distally.

• Arm spine length equal or less than length of arm joint. Spines with broad base, abruptly tapering

near tip, somewhat compressed, surface echinutate with microscopic grooves. Tentacle scale on ventral arm

I plate generally smaller than scale on lateral arm plate.

Remarks. Since its discovery, A. squamata has been assigned at least 25 different scientific

binominals; a partial synonymy is provided by Fell (1946). A.M. Clark (1970) points out that the name

l Ophiura Leach, 1815, has However, her to the International

elegans probably priority. following application

Commission on Zoological Nomenclature the name squamata was placed on the Official List of Specific

Names in Zoology (Opinion 1560, 1989). l

i 143
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Figure 7.10. • Amphiphotis squamata (Delle Chiaje): A, disk, dorsal view (LACM 41-5 12;dd =3.6 mm);B, disk, I

ventral view (LACM 41-5.12; dd = 3.3 mm).

!

It should be noted that the species, as currently recognized, may actually comprise more than one

taxon (Hendler et al., 1995). See remarks under A. pugetana, m

Distribution. Reputedly cosmopolitan, reported for all but extreme polar regions, and at depths

from the intertidal to 1,330 m (Hendler et al., 1995). Preliminary examination of material at LACMNH m

suggests that in Southern California this species typically occurs in littoral habitats; A. pugetana may be II

relatively more numerous offshore.

Biology. Aspects of the species' biology summarized in Austin and Hadfield (1980) and Hendler et m_

al. (1995), include its specializations for viviparous reproduction, hermaphroditism, and luminescence and il

fluorescence. A. squamata is a browser and detritus feeder (Kyte, 1969). Despite its small size, it is the host

of a number of unusual parasites. Nielsen (1932) notes that elsewhere the species is parasitized by copepods II

in the genus CanceriUa, but none of 500 Eastern Pacific specimens in his collection were infested. |

The importance of this species in California waters is evidenced in the survey reports of Hartman

(1955, 1966), Hartman and Barnard (1958, 1960), and Jones (1969), but some occurrences of the species m

noted in these and similar studies probably are based on records of misidentified A. pugetana. Barnard and

Ziesenhenne (1961) recognizedA, squamata at 25% of 176 stations taken between 9-243 m depths from Pt.

Conception, California to the Mexican border. They calculated an overall population density of 44 individuals/ m

m2.My examination of samples still at LACMNH from that survey indicate that identification ofA. pugetana !1

and A. squamata were frequently confused, casting doubt on the survey data. Thompson and Jones (1987)

report both species from the lower slopes of Southern California offshore basins, with densities ofA. squamata I

up to 70 individuals/m 2. I!

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i.

i Amphiura arcystata H.L. Clark, 1911

Figure 7.11

I Amphiura acrystata (lapsus forarcystata) H.L.Clark, 1911:10, 145-148, fig. 58a-i.--Matsumoto, 1917:194,

196, 203, 359.--Downey, 1969:20.

i Amphiura arcystata, Clark, 1915:224.--1940:331, 336.--May, 1924:283.--Nielsen, 1932:264.--

Ziesenhenne, 1937:223.--D'yakonov, 1954:64-65.--Barnard and Ziesenhenne, 1961:140, 143.--

i Ham-nan, 1963: 85, 125, 154, 185,218, 251,285, 320, 335, 349, 382.--A. M. Clark, 1965:44-45.-- Boolootian and Leighton, 1966: 4, 8, fig. 7.--A.M. Clark, 1970:10-11.

Amphiura (Amphiura) arcystata, Irimura (in Imaoka et al.), 1991:142, text fig.

I Hemilepis arcystata, Fell, 1962:10, 19.

Material Examined. California: Santa Maria Basin, Phase I Sta. BSV-042, 100 m (1, USNM primary

I Amphiura voucher); Phase acrystaI Sta. BSS-064, 59 m (3 juvenile, SBMNH secondary voucher [as Amphiuridae sp. A,

ta]); Phase II Sta. R-l, 91 m (3 juvenile, CAMP voucher).--Off Santa Cruz Id., R/V

Velero III Sta. AHF 996-39, 64-82 m (26, LACM 39-100.7).--Off Santa Barbara Id., R/V Velero III Sta.

I AHF 975-39, De 46-49 m (60, LACM 39-79.11). scription. A large amphinrid, attaining over 16 mm dd with arms 10 to 20 times the disk diameter.

A 7 mm dd individual has arms 80 mm long, a 10 mm dd individual has arms 150 mm long, a 15 mm dd

i individual has arms over 195 mm in length. Disk lobate, indented interradially, covering ranging from full coating of small, imbricating scales

to nearly naked (see Clark, 1911: fig. 58). Disk central and radial regions generally covered with soft tissue

I and relatively few minute scales, largest scales surrounding radial shields. Radial shields slender, blunt

tipped, distinctly curved, diverging proximally; shields shaped like paired parentheses (), not in contact.

Jaws projecting ventrally, framing tissue-covered cavity proximal to oral shield. Oral gap

i conspicuous, widening distally. Infradentals blocklike, in in mouth.

large, generally contact, extending high

Buccal scale high on jaw, small, flattened, pointed or spinelike; distal papilla large, bluntly pointed, erect,

m/ projecting ventrally, originating near proximal tip of adoral shield. Tooth chisel-shaped with broad convex

,_1 or straight free edge of imperforate stereom.

Oral shields diamond-shaped to spade-shaped, sides slightly concave, comers lobate, lateral lobes

i largest. Madreporite twice as large as oral shields, inflated, with pores in lateral series. Adoral shields with 3 rounded comers, proximal lobes separated, radial lobes abutting f'trst ventral ann plates. Ventral interradii

fleshy, scales small, scattered; conspicuous imbricating scales bordering adradial edge of bursal slit. Bursal

slits capacious.

I Arms constricted near disk, widening, then tapering distally; in cross section broader than tall,

dorsal surface very slightly rounded, ventral surface nearly flat.

I edges short, Dorsal sometimes arm platesangular. twice asSeveral wide basal as lonplates g, generall reduced y overlap in size, ping,surrounded ovoidal tobysuinbrectangular, tegument. In lateral dors

aspect, wedge of soft tissue and part of lateral arm plate visible between successive dorsal arm plates. In

I alcohol plates Lateralappear arm plates thin, translucent, revealing vertebrae beneath. slender, widely separated; successive plates connected by soft tissue. Five to 8

arm spines near disk, smooth, thick at mid-shaft, gradually tapering toward hollow base and slender rounded

i tip. Spines increasing markedly in length from dorsal to ventral, 1-2 joints in length, dorsal and ventral spines compressed laterally, middle spines dorso-ventrally compressed. Two tentacle scales, thin, flat, ovoidal;

scales on lateral arm plate and on ventral arm plate at right angles, incompletely covering large tentacle

I pore.

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i

Figure 7.11. Amphiura arcystata H.L. Clark: A, disk, dorsal view (LACM 39-79.11; dd = 6.4 ram); B, disk, I

ventral view (LACM 39-100.7; dd = 10.1 ram);C, disk, dorsal view; D, disk, ventral view (LACM

39-79.11; dd = 6.4mm). Note variability in disk scalation, i

Ventral arm plates thick, opaque; first ventral ann plate in oral slit small and slender, several adjacent i

plates longer than wide, other proximal plates wider than long, hexagonal or octagonal, overlapping.

Color: Disk gray to violet gray or brownish, in small specimens primary plates brownish-black; I

radial shields brown or brownish-black with white edge or distal tip, scales surrounding radial shields tan,

brown, or brownish black with white edge, ventral interradius gray, oral frame gray or tan and whitish,

whitish scales border bursal slits, reddish brown stomach and gonads visible through thin body wall. Dorsal I surface of arms gray, brown, or tan, with irregular bands of pale tan, cream or whitish; arm spines tan,

brown, or gray, ventral surface of arms salmon colored, more cream colored distally, connective tissue on

arms whitish to violet. Eggs pale pink. i

Specimens in alcohol fade to pale gray, brown, and yellowish, with disk darker than arms and with

dark radial shields, lateral arm plates, and arm spine bases. Densely pigmented stomach visible through

naked portions of disk. Purplish muscle masses visible between joints on dorsal surface of arm. I

146 I
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i Distribution. Mexico and Southern California to Monterey, California at 15-844 m, and in Japan at

108-848 m depths (H.L. Clark, 1911, 1915, 1940; May, 1924: Nielsen, 1932; Ziesenhenne, 1937; Hartman,

I 1963; A.RemM. aClark, rks. Juvenile 1965; Irimur and asmall , in Imaoka individuals et al.,1.5-2.5 1991).mm dd are unlike the adults in appearance. They

have prominent primary plates, the disk completely covered with thin scales, radial shields relatively larger

I than array,inandmature theyinlack dividua tentacle ls, fan-shaped scales. H.doL.rsalClark arm (1911:147) plates, elongate observed ventralthatarmsmall plates,specimens modifiedhave oral papillae the disk

almost completely scaled and the largest have scales only around the radial shields, but scale cover is not

i wholly remarkable, relatedaffording to age; "diversit an unusually y in theinteresting amount ofcasescaling of resorption on the diskof calcareous and interbrachial matter."spaces A.M.beClark low is(1965, most

1970) discussed taxonomic implications of the morphological variability shown by this species.

I be The correct spelling of the species name is arcystata, although Downey's (1969) comments might interpreted to suggest otherwise. In the original description (H. L. Clark, 1911), acrystata was a lapsus

forarcystata, and the error was noted and corrected by the author in a later publication (H. L. Clark, 1915).

I Biology. Barnard and Ziesenhenne (1961) found this species at 22% of 176 stations taken between 9-243 m depths from Pt. Conception, California to the Mexican border. They calculated a mean population

density of 2.8 individuals/m 2,but densities are certainly higher at some localities where the species occurs.

I They found that A. arcystata is typically a "subdominant" in the benthic community characterized by Amphiodia urtica. It is a burrowing ophiuroid, and the very long arms of adult specimens suggest that the

disk may be situated at least several centimeters below the surface of the sediment. I have found that the

I proximal tube feet are smooth and the distal tube feet are highly papillose, as is often the case in suspension

feeding ophiuroids.

.! Amphiura diomedeae Lfitken and Mortensen, 1899

Figure 7.12

i Amphiura Diomedeae Liitken and Mortensen, 1899:151-152, pl. 12, figs. 1-7.

Amphiura diomedeae, H.L. Clark, 1911:10,15,17,18, 20,140-141.--I915:225.--1913:214.--1917:436-

I 437.---Campbell, 1921b:47.--Boone, 1933:105-106, pl. 61.--Koehler, 1922:4, 158-159, pl. 67,

figs. 7-10.--Fell, 1962:21.--H. L. Clark, 1939:56-57.--Hartman, 1963:86, 383, 401.

i Amphiura Amphiura (Section diomediaeAmphiodia , A.M. Clark, ) diomedeae 1970:39-44, , Koehler, 48, fig.1907:306. 8n,o.

Amphiura koreae, Matsumoto, 1917:151-152, 174, 195, 198, 356 (in part ?, norA. koreae Duncan, 1879).--

i Astrahantseff and Alton, 1965:1414, 1416, 1420, 1422 (notA. koreae Duncan ?).

Material Examined. Oregon: Off Columbia River mouth, R/V Commando, 460 m (1, LACM 64-

i Amphioplus 240.1). - Califsp.ornia: A]).----Off SantaSanta MafiaBarbara, Basin, Phase R/V Velero/V I Sta. BSS-062, Sta. AHF5823036-55, m (1, 400 USNMm (1,primary LACM v55-19.3).-- oucher, [as

Off Santa Catalina Id., R/V VeIero/I/Sta. AHF 2413-53, 375 m (11, LACM 53-112.4).--Mexico: Off

i Albatross Mazatlan, Sta. R/V3394, Albatross 935 m, Sta.syntypes 3429, 1,68(3,0USNM m, syntype 19533); (1, USNM R/V Albatross 19535).--Panama: Sta. 3393, 1,866 Gulf m, of Panama, syntypes R(10, /V

USNM 19532); off Azuero Peninsula, R/V Albatross Sta. 3392, 2,323 m (3, USNM 19531).--Ecuador: Off

I/ Cojimies, R/V Albatross Sta. 3398, 2,877 m, syntype (1, USNM 19534).--Japan: Honshu Id., off Port

_, _ Arafi, R/V Albatross Sta. 3699, 732-1,328 m (12, USNM 26630); Sagami Bay, R/V Albatross Sta. 5087,

[ 1,123 m (2, USNM 26618); Suruga Gulf, R/V Albatross Sta. 5057, 494 m (3, USNM 27114).

!

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il

Description. An amphiurid of moderate size, attaining a disk diameter of 15 ram. A specimen 7 l_

mm disk diameter has arms over 50 mm long; one 13 mm disk diameter has arms 135 mm long.

Disk rounded, 5-lobed, indented interradially, covered with relatively few, distinct, angular, i

imbricating scales. Primary plates discernible; primaries and several scales between paired radial shields U

are largest on the disk. Radial shields prominent, 1/4 disk diameter, 2-3 times longer than wide, broadest

distally, with straight, thickened distal edge, abruptly narrowing proximally; shields in contact distally or •

separated by single small scale, separated by wedge of scales proximally.

Jaws with sides concave and flaring, narrowing ventrally, diverging dorsally. Oral gap broad, large

diastema separating proximal and distal oral papillae, broad distal end of oral slit filled by large first ventral m

arm plate, second oral tentacle pore conspicuous in ventral aspect. Oral papillae may vary in number and I

appearance among jaws of an individual. Infradental papillae flattened, often with proximal point; pairs

separated, in contact, or displaced by small apical oral papilla. Buccal scales high in jaw, ranging from i

small, triangular to large, crescentic. Distal oral papilla spinelike, or flattened and bluntly pointed, may be i

single, paired, or absent on different jaws. Tentacle scale of second oral papilla adpressed, flat, operculate,

set between adoral shield and first ventral arm plate. Teeth usually with constriction behind free edge. n

m

Oral shields pentagonal to rhombic, comers rounded, with central depression. Adoral shields large,

equal in length to oral shields, proximal lobes blunt, radial lobe sharply pointed. Adoral shields broadly i

separated by first ventral arm plate. Oral and adoral shields thin; madreporite inflated, multiperforate, i

i=

Arms gradually tapering, beginning at edge of disk; in cross section rounded above and flat below.

Proximal dorsal arm plates broadly fan-shaped; proximal end pointed, edges straight; distal end •

convex. Dorsal arm plates narrowly separated by lateral arm plates and wedge of connective tissue. g

Lateral arm plate with prominent, thick, spine-bearing ridge. Three arm spines with thick base,

gradually tapering to acute point with slight distal bend, smooth, delicate, hollow. Dorsal-most spine shortest,

most slender; middle spine largest, somewhat torso-ventrally compressed, slightly longer than the arm

joint.

Ventral arm plates pentagonal, edges concave, disto-lateral comers large, lobate; broader than long R

on proximal joints, becoming more elongate on distal joints. Two tentacle scales rounded or bluntly pointed, m

covering most of tentacle pore on basal joints.

m

Color: As noted by H. L. Clark (1939), preserved specimens range from entirely ivory white to i

having grayish disks and yellowish white arms. Liitken and Mortensen (1899) described specimens in alcohol

u

as having a bluish disk and white or pale brown arms, some with a reddish at the outer end of the radial

shields. I

Remarks. It is only because of the intraspecific variability of the oral formula in A. diomedeae,

characterized by A. M. Clark (1970), that I have retained this species in the genus Amphiura. As shown by l

Hendler (1988), the homologies between the oral papillae of amphiurids suggest that distinctions between

nominal Amphiura and Amphioplus species with 4 pairs of oral papillae can be baseless. Once sufficient

material is available to clarify the homologies of its oral papillae, the inclusion ofA. diomedeae inAmphioplus i

may be warranted. •

m

H.L. Clark (1939), based on published descriptions, did not accept Matsumoto's (1917) synonymy

of Amphiura diomedeae IAitken and Mortensen with A. koreae Duncan. A. M. Clark (1970) examined m

relevant type material, noting that most specimens of A. koreae (sensu Matsumoto) are referrable to A. I

diomedeae Liitken and Mortensen. In addition, she found that the holotype of A. koreae Duncan has oral

structures unlike an Amphiura species. /

A. M. Clark (1965, 1970) suggested thatA, diomedeae may be synonymous withAmphilepis diastata II

Murakami and withAmphiura caulleryi Koehler, raising additional questions regarding its taxonomic status.

Fortunately, Irimura's account (in Imaoka et al. 1991) of Amphiura (Amphiura) koreae, based on new •

Japanese collections, shows that additional material could be made available to evaluate the(se) species. !1

|
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I Figure 7.12. Amphiura diomedeae Ltitken and Mortensen: A, disk, dorsal view; B, disk, ventral view; C, oral

frame (USNM 19532,syntype; dd = 15.1 mm); D, oral frame (USNM 19532,syntype; dd = 11.0

I nun). Note variability of Amphioplus-like oralpapillae.

I Distribution. Since its discovery, the species has been reported from off: Panama and Ecuador,

m 935-2,877 m (Liitken and Mortensen, 1899), Japan 71-1,369 m (H.L. Clark, t911), California, 869-1,205 m

i (H.L. Clark, 1913, 1915), Tahiti, 1,325 m, Peru, 1,895 m (H.L. Clark, 1917), Philippine Islands, 368-567 m

(Koehler, 1922), Costa Rica, 183 m (Boone, 1933), the Maldives, 797 m and South Arabian Coast, 1,046 m

(H. L. Clark, 1939). Additional records now exist for California, Oregon, and Washington, 450-3,030 m

1 (Hartman, 1963; Astrahantseff and Alton, 1965 [as A. koreae], Carey, 1990). As discussed above, some of

! U these reports may pertain to A. koreae Duncan.

Biology. Summers (1993) found that A. diomedeae and Amphiura carchara H. L. Clark are the

i dominant between 200 and 3,000 off the central California continental In her ophiuroids at depths m slope.

study area, A. diomedeae composed 26% of the ophiuroids sampled. Densities of individuals, when they

occurred in box cores, ranged from 0.2-0.42/0.1 m2;similar densities (asA. koreae) have been found offthe

i Oregon coast (Alton, 1972).

The extensive bathymerdc range of the species exposes it to a considerable range of environmental

variation. Bottom temperatures recorded at collecting sites range from 1.6-20.3°C (Liitken and Mortensen,

I 1899; H. L. Clark, 1911, 1913, 1917).

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i 149
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ii

Dougaloplus amphacanthus (McClendon, 1909) •

Figures 7.13 and 7.14

i

Ophiocnida amphacantha McClendon, 1909:46-47, figs. 24, 25.--H.L. Clark, 1915:250.---Campbell,

1921b:48.

Amphiacantha amphacantha, Matsumoto, 1917:178.--Nielsen, 1932:296-297.--Barnard and Ziesenhenne, I

196I:143.--Hartman, 1963:85,217, 237, 285, 305,349, 382.--Boolootian and Leighton, 1966:5,

10, fig. 15. ii

Dougaloplus amphacantha, A. M. Clark, 1970:33-34, 79. I

Amphioplus hexacanthus H.L. Clark, 1911:10, 170.--Matsumoto, 1917:170, 172, 358.--Barnard and

Ziesenhenne, 1961:135-151.--Fe11, 1962: 22.--Hartman, 1963:85, 153, 217, 285.--Parker,

1964:359.--Boolootian and Leighton, 1966: 5, 10, fig. 17.--Sutton, 1976:349. II

Amphilimna pentacantha H.L. Clark, 1911:10, 172-173, fig. 77a-c.--1915:259.---Campbell, 1921b:48.--

May, 1924:281-282. i

Amphiacantha pentacantha, Matsumoto 1917:178.

Material Examined. California: Point Reyes, R/V Albatross Sta. 3165, 91 m, syntypes (4, USNM I

25583 [as Amphioplus hexacanthus]).--Monterey Bay, off Point Pinos, R/V Albatross Sta. 3131, 88 m,

In

paratypes (36, USNM 27141 [as AmphiIimna pentacantha]).--Santa Maria Basin, Phase I Sta. BSV-042,

100 m (1+, USNM primary voucher); Phase I Sta. BSR-073, 98 m (1, SBMNH secondary voucher); Phase i_

II Sta. PJ-1,145 m (5, CAMP voucher); Phase I Sta. BSS-064, 59 m (5, SBMNH secondary voucher [as m

Amphioplus hexacanthus]); Phase I Sta. BSR-021,49 m (2, USNM primary voucher).--Los Angeles, offEl

Segundo, R/V Velero IV Sta. AHF 5960-58, 144 m (4, LACM 58-140.7).--Mexico: Gulf of California, off

Isla las Animas, R/V Velero IV Sta. AHF 11837-67, 1,100-1,326 m (48, LACM 67-302.1). il

Description. This moderately large amphiurid was originally described on the basis of a specimen I

7.5 mm dd with arms 10 times that length. Disk soft, lobed, indented radially and/or interradially, covered 1

with numerous small, overlapping scales bearing evenly spaced spines. Spines acute or with tip slightly

expanded. Disk spines short, slender, tapered, smooth. Radial shields slender, tapering and diverging 1

proximally, distal tips in contact. _

Jaws narrow with apex jutting ventrally, oral gap teardrop-shaped. Teeth with straight, narrow tip,

edge composed of imperforate stereom. Five oral papillae; infradental papillae separated, blocklike, smoothly ill

rounded; buccal scale near ventral edge of jaw, laterally flattened, pointed; two distal papillae in contact, I

pointed, spinelike; oral tentacle scale rounded, flat, abutting f'trst ventral arm plate and adoral shield. Oral

shields equidimensional or longer than wide, with four lobes or ovoid with narrow distal lobe, overlapping m

central part of adoral shield. Madreporite more irregular in outline than oral shield, larger, thicker, with 1

single pore. Adoral shields with small proximal lobes touching or nearly so above oral shield, adoral lobes

forming acute angle between first and second ventral arm plates. II

Ventral interradius with scale and spine cover similar to dorsal disk, cover reduced near oral shield. I

Bursal slits extending from oral shield nearly to disk edge; prominent series of overlapping scales reinforcing

adradial edge of slit. I

Arms long, slender, somewhat wider at midlength, narrowing proximally and distally; in cross

section more rounded dorsally than ventrally, wider than high. Conspicuous band of soft tissue between arm

joints. I

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I Dorsal arm plates ovoidal, wider than tong, barely in contact, lateral edges sometimes truncate;

several plates near disk rounded, diamond-shaped.

I narrow. Lateral Up arm plates narrow, with prominent spine-bearing ridge. Arm spines hollow, internal channel to 6 arm spines on proximal joints, upper and lowermost spines longest, pointed; middle spines

with 1-2 microscopic teeth at tip; on distal joints with 3-5 spines, upper spine shortest, lowest 1-2 spines

I longest, hyaline teeth secondat tifrom p. In adults, bottomupper flattened arm spines with broad laterallybase, flattened, thin lateral toothededgearmmicroscopically spines dorso-venlTaUy roughened, flattened, 1-2

ventral spine nearly round. Two (or one) tentacle scales, small, thin, leaving most of tentacle pore uncovered;

i scale on lateral arm plate larger. Ventral arm plates barely in contact or slightly separated by lateral arm plates; first ventral arm

plate forming knob at outer edge of oral slit; plates below disk longer than wide, distal plates nearly equilateral,

I octagonal to pentagonal, with rounded proximal edge, medial distal indentation. Variation: Specimens 4 mm disk diameter or less are not truly juvenile since they have recognizable

gonads, but they differ from larger individuals in having a spinelike buccal scale, distal oral papillae with

I rounded tips, four arm spines, numerous joints beating only 1 tentacle scale, ventral arm plates in contact, and reduced disk spination.

i Color: Inalcohol disk gray; radial shields, jaws, adoral and oral shields, and lateral arm plates often with patches of yellow pigment. Arms faded reddish-pink, lateral arm plates with orange pigmentation,

central portions of ventral arms plates reddish. Various authors report thatpreserved material is yellowish

overall, or has the disk gray or greenish and the arms yellowish, or yellowish white (McClendon, 1909;

I May, 1924; Nielsen, 1932). specimens are pale gray.

Dried

Remarks. As indicated in the synonymy above, this species was placed in Ophiocnida by McClendon

I (1909), and transferred to Amphiacantha by Matsumoto (1917). Nielsen (1932), proposed A. pentacantha (H. L. Clark) to be a junior synonym, a point that I verified by examination of H. L. Clark's type material.

Since the generic name Amphiacantha was preoccupied, A. M. Clark (1970) provided the replacement

i name Dougaloplus, and revised the genus. She suggested that Amphiacantha derjugini D'yakonov is a

synonym of D. amphacanthus, but possible differences in the arm spine shape and tentacle scale number

open that conclusion to question.

, I have synonynftzedAmphioplus hexacanthus with D. amphacanthus, based on a comparison of the

II syntypesof A. hexacanthus (Figure 7.14) with the original description and specimens recognized as D.

amphacanthus, McClendon's original specimens of the latter presently being unavailable. I surmise that

Hdorsal .L. Cldisks ark wamong ould hthe ave 6recfragme ognizedntarythespecimens corresponden of Ac.e hexacan betweenthustheattwhiso disposal. nominal species My onlyhareservation d there beenin

this matter is the absence of any mention of toothed arm spines by McClendon (1909) and Nielsen (1932),

i Clark a characteristic that is distinctive in this species. However, the oversight is less surprising given that H.L. recognized toothed spines inA. hexacanthus, but ignored them in his description ofA. pentacantha, as

did May (1924); toothed spines are present in the type material of both nominal species (Hendler, prev.

I--' unpub')'I)istribution. Gulf of California (doubtful locality?), San Diego to Pt. Reyes, Califomia, and Uraga

Strait, Japan (doubtful locality, see below), at 62-1,326 m depths (McClendon, 1909; May, 1924; H. L.

I C(consisting lark, 1911,of19arm15; Nielse fragments n, 1932; only,Hartman, which I 1963, have not thisexamined) report). Onreportedly e ofH. L. Clark originated 's typefromlots ofUraga A. hexacanthus Strait, off

Tokyo, Japan. However, the nominal speciesA, hexacanthus, has not been rediscovered from Japan (lrimura,

i et al. 1995). On that basis, I consider an amphi-Pacific distribution doubtful.

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Figure 7.13. Dougaloplus amphacanthus (McClendon): A, disk, dorsal view; B, disk, ventral view; C, oral frame; 1

D, arm, dorsal view; E, arm, ventral view; F, arm spines, showing glassy teeth (LACM 58-140.7; dd

= 7.4 mm). .I

implicationBiologwithy. regard The sytononymy the offshoref Amphioplus benthic ecology hexacanthus of SouthernwithCalifornia. D. amphacanthus DougaIoplushasamphacanthus an important I

would appear to be the third most abundant ophiuroid on the Southern California coastal shelf, its distribution

more extensive Barnard and its population densities greater than previously recognized. I and Ziesenhenne recorded A. hexacanthus at 30% of 176 stations taken at 9-243 m depths

from Pt. Conception, California to the Mexican border, estimating a mean population density of 13.1

Amphiacantha individuals/m ) 2, with amphacanthus densities higher was found at localities at 15%where of theindividuals stations, atarelower present. densities In thethansamereported study, Dfor. (asA. i

hexacanthus. Presumably, the original data could be reexamined to accurately determine the distribution of

D. amphacantha. I

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I Figure 7.14. Amphioplus hexacanthus H. L. Clark: A, entire, dorsal view, disk missing; B, entire, ventral view; C,

oral frame; D, arm, dorsal view, showing elongate, flattened, middle ann spines; E, arm spines,

I showing glassy teeth (USNM 25583, syntype; off = 2.5 mm).

Barnard and Ziesenhenne (1961) described theAmphioplus hexacanthus community (more correctly the Dougaloplus amphacanthus community) as one of three major benthic communities on the coastal shelf

between Pt. Conception and Mexico, notingAmphiodia urtica as a community subdominant. The association

I was found to occur in silty-sand sediments in which silt and sand are found in nearly equal proportion, most prominently on the California shelf off San Pedro, Santa Monica, and Oceanside in 13-55 m depths, and

several laundered miles to the north in deeper waters, 55-91 m, on the Monterey Bay shelf. Hartman (1955,

I 1966), Hartman and Barnard (1958 1960), and Jones (1969) provide additional evidence for the numerical abundance of the species.

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I

Amphiuridae sp. juvenile g

Figure 7.15

I

Among the many immature specimens of Amphiuridae in this collection, one was sufficiently

distinctive to warrant a detailed characterization. mm

Material Examined. California: Western Santa Barbara Channel, Phase I Sta. BSS-087, 762 m (1, 1

USNM primary voucher [as Amphiodia psara]).

Description. The specimen has a disk diameter only 3.3 mm (ofi 1.4 mm), with arms broken near I

the disk. It

Disk rounded pentagonal, flat; cover consisting of primary plates, large interradial and marginal

scales, separated by smaller, thin, overlapping scales. Radial shields half-circle shaped, twice as long as _l

wide, in contact throughout. m

Infradental papillae elongated, with distal ends pointed, diverging. Buccal scale flattened, n

subtriangular, situated high in jaw. Two lateral papillae, fiat, free edge expanded. !

Oral shields subpentagonal, about as long as wide; madreporite larger, inflated, bearing one blunt

spinelet. Adoral shields nearly equal in size to oral shield, meeting broadly above oral shield; radial comer am

pointed, inserting between first and second ventral arm plates. 1

Ventral interradius scale-covered, scaling reduced in density near oral shield. Genital slit extending

from oral shield to disk edge, scales bordering edges of slit.

Dorsal II arm plates nearly half-circle shaped with proximo-lateral edges straight, distal edge somewhat

protruding, plate about twice as wide as long; successive plates in contact.

Lateral arm plates broad, with strong, distally directed spine-bearing ridge. Three arm spines, smooth, I

robust, with broad base, gradually tapering, tip quite blunt; dorsal and ventral spines laterally compressed; Ill

middle spine dorso-ventrally compressed. Tentacle scales absent.

_lm

Ventral arm plates subpentagonal, as long as wide, posterolateral comers lobate, distal edge concave; I

successive plates narrowly separated by lateral ann plates.

Remarks. This specimen belongs to one of the amphiurid taxa with an adult oral formula consisting m

of 4 or more oral papillae. It does not appear to be a juvenile growth stage of a species described in the 1

present report.

The possibility that it is an Amphiodia species can be ruled out. Amphiodia individuals of this size t

lack a buccal scale and show species-specific morphological features; for example juvenile Amphiodia It

psara H. L. Clark possess a long shank on the oral shield and have the arm spines laterally compressed. It is

not Amphioplus strongyloplax or Dougaloplus amphacantha, both of which at a similar disk diameter have

a differing and distinctive spine morphology. It does not appear to represent the Eastern PacificAmphiopIus II

species,A, da/eus (Lyman) orA.philohelminthius Ziesenhenne, and does not appear to be ajuvenileAmphiura

diomedeae or a more typical Amphiura species. Furthermore it lacks the disk spines, granulation, oral I_

papilla shape, and other characteristics of Eastern Pacific Ophiocnida and Amphichondrius species. I!

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I Figure 7.15. Amphiuridae sp.juvenile: A, disk, view; B, disk, (USNM, primary

dorsal ventral view voucher; dd

= 3.3mm).

I

Family Ophiacanthidae

I Ophiacantha d_lasia H. L. Clark, 1911

i Figure 7.16

Ophiacantha diplasia H. L. Clark, 1911:10, 209-211, fig. 97a-c.--1915:198.--May 1924:272-273, fig.

3a,b.--Hartman, 1963:85.--Astrahantseff and Alton, 1965:1414-1415, 1420.--Boolootian and

I Leighton, 1966:7, 8, fig. 30.

Ophiophthalmus diplasia, Kyte, 1982:508.

I Material Examined. California: Santa Maria Basin, Phase I Sta.BRA-016, 91.5-123 m (3, SBMNH

secondary voucher); Phase I Sta. BRA-027, 96-126 m (1 juvenile cf. O. diplasia, USNM primary voucher

i [as Ophiophthalmus normani]).--Westem Santa Barbara Channel, Phase I Sta. BRA-002, 110-126 m (6, USNM primary voucher).--Monterey Bay, off Pt. Pinos, R/V Velero 111Sta. AHF 890-38, 90-99 m (1,

LACM 38-158.2).--Off Santa Barbara, R/V Velero/V Sta. AHF 6888-60, 113 m (6, LACM 60-276.1).--

Off Santa Rosa Id., R/V VelerolI1Sta. AHF 1397-41,132-141 m (5, LACM 41-177.2).---Off Santa Catalina

I Id., R/V Velero/V Sta. AHF 1995-50, 732 m (1, LACM 50-100.2).---Off San Clemente Id., II1

R/V Velero

Sta. AHF 1023-39, 101-201 m (1, LACM 39-127.13).--Los Angeles, off Redondo Beach, Burch 40122, 90

m (3, LACM 40-341.1).

I Description. An ophiacanthid species of moderate size. Largest specimens attain 25 mm dd; a

specimen 23 mm dd has arms approximately 150 mm long.

i granulesDisomewhat sk roundedlonger pentagonal, and largerflat. towards Densel disky margin. coveredRadial with close shieldsly subtriangular, spaced, rough,small, elongate approximately granules;

one-tenth diameter of disk in length; broad distal edge relatively bare of granules.

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Jaws broad, with apex jutting ventrally; oral gap wide, not sealed by oral papillae. Terminal oral •

papilla generally single, sometimes paired; 5-7 lateral oral papillae, long, spatulate, with bluntly rounded

tip; proximal papillae laterally compressed, distal papillae dorso-ventrally compressed. Outer oral tentacle t

large, emerging at edge of oral slit, associated tentacle scale triangular, originating on first ventral arm II

plate. Teeth chisel-shaped, free edge unevenly rounded. Oral shields wider than long, with two large lateral

lobes, narrow distal shaft; madrepofite larger, more irregular in shape. Adoral shields three-lobed; proximal •

lobe extending above oral shield, paired shields nearly in contact. U

Bursal slit widest proximally, extending from shaft of oral shield to edge of disk, adradial edge

bordered by naked scales. Periphery of ventral interradius covered by V-shaped field of granules or spinelets, l

thin imbricating scales covering proximal region. tl

Arms gradually tapering, shape obscured by long arm spines; in cross section broader than tall,

dorsal surface somewhat more curved than ventral surface.

n

First ventral arm plate U-shaped, others subpentagonal, broader than long on proximal joints,

becoming equidimensional, then longer than wide and fan-shaped near arm tip; plates beneath disk depressed,

those beyond disk edge with slight medial keel or thickened distal edge; successive plates barely in contact I

or separated by lateral arm plates. i

Lateral arm plates wide, with prominent spine-bearing ridge, sharply pointed lobes intruding between

dorsal and ventral arm plates; successive plates separated by narrow band of soft tissue. Maximum of 7 arm •

spines, decreasing in size ventrally, two uppermost spines dramatically longer, more acutely pointed than

two ventral-most spines, longest spine equal in length to 4-7 arm joints. Spines erect, hollow, compressed,

gradually tapered, tips truncate or bluntly rounded, lateral edges bearing microscopic echinulation. Longest 1

spines on first two armjoints directed toward center of disk. Two tentacle scales elongate, flat; inner scale

w

longer, narrower, more acutely pointed than outer scale. I

Dorsal arm plates diamond-shaped rhombic to fan-shaped, medial portion elevated; proximal plates U

wider than long, in contact; distal plates longer than wide, separated by lateral arm plates.

Variations. Large specimens may have granules between or on some proximal dorsal arm plates. •

Specimens less than 6 mm disk diameter have all the dorsal arm plates separated. There are scattered

spinelets on dorsal disk surface of individuals 1.4 - 2.4 mm dd. A juvenile specimen in this collection 1.2

mm dd has: disk covered with transparent scales bearing widely spaced irregular stumps; jaws with apical •

and only 3 lateral oral papillae; second oral tentacle pore opening outside the mouth angle; arm spines 1

slender, smooth; ventral arm plates with slender waist, successive plates in contact. A juvenile Ophiacantha

normani of similar size has: disk bearing conspicuous central primary plate and sharp spinelets; arm spines I

thickened at base; ventral arm plates not in contact. l

Color: Living specimens with disk irregularly mottled brown and gray. Arms reddish; arm spines

light brown with conspicuous white basal ligament, t

It In alcohol, mottled gray, white and brown; darker pigmentation retained by disk, ventral arm spines,

and dorsal arm surface. Arms generally paler than disk.

Kyte (1982) assigned this species to Ophiophthalmus Matsumoto, but that generic name 1

Remarks.

is an invalid junior synonym (see Paterson, 1985). A new generic assignment is probably warranted for O.

.U

diplasia and other ophiacanthids such as Ophiacantha ("Ophiophthalmus") normani Lyman, but they can

be considered as Ophiacantha species pending a revision of the entire genus, n

Distribution. Southern California to Oregon, at depths 7t-1,330 m (May, 1924; H. L. Clark, 1911,

1915; Hartman and Barnard, 1958; Astrahantseff and Alton, 1965). m

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I Figure 7.16. Ophiacanthadiplasia H.L.Clark:A, disk,dorsal view;B, disk, ventralview (USNM,primaryvoucher; dd = 14.8mm).

I

Biology. This species usually occurs on hard substrates, associated with rock structures and sessile

I portray invertebrates. it as anPhotographic abundant element recordsof a(Lissner diverse and hard-bottom Benech, 1993:fig. community. 4.3)Individuals from off Pt.taConception, ke exposed orCalifornia partially

sheltered positions, with their arms slightly upraised. This posture is often shown by suspension feeders,

i preserved and like many ophiacanthids, O. diplasia is probably capable of suspension feeding. Its tube feet, in alcohol

specimens, are large, blunt-tipped, and highly papillose. Tube foot papillae are typical, but not

necessarily exclusively characteristic, of suspension feeding ophiuroids.

I Family Ophiactidae

i Ophiopholis baked MeClendon, 1909

Figure 7.17

I Ophiopholis bakeri McClendon, 1909:41, figs. 26-27.--H.L. Clark, 1911:10, 115-116, 121-123, 128, fig.

46a-c.--1915:267.---Campbell, 1921b:48.--May, 1924:279-280, fig. 7a-c.--Berkeley, 1927:4.--

I Nielsen, 1932:260-262, fig. 5.--Ziesenhenne, 1937:224.--Bamard and Zieserthenne, 1961:143.-- Hartman, 1963:85, 201, 218, 366, 383, 401.--Astrahantseff and Alton, 1965:1414, 1420.--

Boolootian and Leighton, 1966:7-8, fig. 28.--Kyte, 1969:1729, 1732, 1735, 1738.

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Material Examined. California: Santa Maria Basin, Phase ISta. BRA-016, 91.5-123 m (58, SBMNH g

secondary voucher); Phase ISta. BRA-016, 91.5-123 m (1, juvenile O. bakeri? [asAmphipholis squamata]);

Phase I Sta. BRA-025, 64.5-72 m (1 juvenile, SBMNH secondary voucher [as Ophiothrix spiculata]).-- t

Western Santa Barbara Channel, Phase I Sta. BRA-002, 110-126 m (24, USNM primary voucher).--Los II

Angeles, off Redondo Beach, Burch Sta. 40122, 90 m (16, LACM 40-341.2).--Mexico: Pacific Coast, Baja

California, off tslas San Benito, R/V Velero III Sta. AHF 1010-39, 130-174 m (numerous, LACM 39- II

114.17). U

Description. An ophiactid species of moderate size. Individuals attain 10 mm dd with arms 65 mm

in length. A specimen 7 mm dd has arms 32 mm long. The lateral and ventral arm plates and the major 1

ossicles of the oral frame appear rough due to numerous microscopic bumps (EPT). It

Disk round to pentagonal, interradiat regions somewhat outpouched, densely covered with pointed

granules and spinules borne on thin, rounded, imbricating scales and radial shields. Spinules longest, most

numerous in central and lateral areas of disk, generally reduced to sharply pointed granules on radial shields. It

Spinules echinulate or multifid, generally with 2-5 points, up to 5 times longer than wide. Primary plates

evident in small and sparsely armored specimens. Radial shields separated except distally, twice as long as i

wide, approximately one-fourth diameter of disk. II

Jaws narrow, oral gap broad. Oral papillae 4-6 on each side of jaw, flattened to spinelike. Free edge

of ventral teeth with several points or knobs, sometimes fragmented; dorsal teeth with straight tip. i

Oral shields lozenge-shaped, at least twice as wide as long, lateral lobes wing shaped. Madreporite

twice as large as oral shield, more rounded in outline. Adoral shields nearly equal in size to oral shield, L- !1

shaped, distal tip touching second ventral arm plate, proximal tips extending above oral shield, in contact or i

nearly so.

i

Ventral interradial region covered with small, multifid spinules bearing 1-2 points, pointed granules •

near oral shield. Bursal slit narrowing distally, extending to disk margin; naked scales on edge of bursal slit. !1

Arms tapering. Terminal portion appearing beaded due to protruding lateral arm plates.

Dorsal arm plates inflated, rounded on proximal joints, becoming ovoidal and elongate distally, i

teardrop-shaped on terminal segments. Supplementary scales bordering free edges of dorsal arm plates not i

fully in contact, granular to slender and flat in shape, with one or two sharp points.

Lateral arm plates broad, with prominent spine-bearing ridge; successive plates separated by band i

of connective tissue. Up to 7 ann spines, tapering, solid, generally 2 and less than 3 arm joints in length with

i=

expanded, hoof-shaped base. Longest spines occurring dorsally, near disk, occasionally topmost spine of

reduced size. Longer spines echinulate, blunt tipped; smaller ventral spines with echinulate edge and tip; i

lowermost spines on distal half of arm with prominent hyaline teeth, 2 teeth on terminal segments. Hooked g

spines arising on proximal arm segments in juvenile individuals. m

Ventral arm plates octagonal, length equal to width; proximal edge convex, distal edge markedly i

concave. First ventral arm plate small, enclosed by adoral shields. Ventral surface of basal plates depressed,

forming medial trough. Successive plates separated by connective tissue. Single tentacle scale (occasionally i

2), flat, elongated, with expanded, blunt distal tip, partially covering large tentacle pore.

Variation: A 1.2 mm dd specimen of O. baked from the present survey had: central plate perforate

and prominent but less than 1/3 the disk diameter, radial and central plates of almost equal size; teeth with •

1central and 2 lateral cusps; basal arm plates separated, some possessing spiny stumps; arm spines neither B

palmate nor webbed, short thorns on proximal arm spines, hooked arm spines with 2 hyaline teeth. It was

incorrectly identified as Ophiothrix spiculata by survey taxonomists. Individuals of the latter species less /

than 2 mm dd can be distinguished as they have: disk pentagonal with interradial outpouchings, central plate |

covering 113the disk diameter, convex, hyaline, with minute perforations restricted to edge of plate; radial

plate small; dorsal arm plates lacking spiny stumps, basal plates in contact; proximal arm spines palmate, i

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I Figure 7.17. 7.9 Ophioph mm).olisbaked McClendon: A, disk, dorsal view; B, disk, ventral view (LACM 40-341.2; dd=

I

bearing long thorns webbed with gossamer integument, other spines with small thorns; hooked arm spines

I with 2 hyaline Color: teeth. In living specimen, disk reddish brown with gray central region; arms reddish brown above,

yellowish-white with brown spots below, arm spines brown, tube feet yellow; conspicuous connective tissue

I between occasionalventral light armband;plathetesarmspines . Ziesenhenne are white (1937:or224) gray."noted that the "arms, orange in life, are pink with an

In alcohol, whitish to pink or reddish, sometimes variegated with white or brown.

I Remarks. H. L. Clark (1911:116) provided a key to 5 species and 3 varieties of Ophiopholis: mirabilis, brachyactis, longispina, baked, aculeata (and toaculeata varieties japonica, typica, and kennerly_).

Ophiopholis baker/, longispina, and aculeata typica and kennerlyi occur in California waters.

I Full specific rank originally accorded O.japonica was revived by Irimura (in Imaoka et al., 1990).

The infraspecific status originally accorded var. kennerlyi by H. L. Clark (1911) has been supported by later

authors (May, 1924; Nielsen, 1932).

I Distribution. Baja California, Mexico, California, Oregon, Washington, British Columbia; 18-902

m (McClendon, 1909; H. L. Clark, 1911; Berkeley, 1927; Nielsen, 1932; Ziesenhenne, 1937; Hartman,

i 1963; Astrahantseff Biology. It isandlikelyAlton, that1965; O. baker/ Kyte, , like1969). its congener O. aculeata, is a rheophilic suspension feeder,

probably epizoic or occupying positions elevated above the sediment in hard bottom habitats (see LaBarbera,

I the 1978, base1982). and Iholdfasts have foundof living it in a Macrocystis dead kelp holdfast . It is commonly recovered from recovered Santa fromMonicasiltyBay, sand,andandit likely sand, occupies rock and

shell substrates (Kyte, 1969). Its affinity for hard substrate and rock outcroppings may account for the low

i numbers taken in the survey reported by Bamard and Ziesenhenne (1961).

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Family Ophiodermatidae I

Ophiuroconis bispinosa Ziesenhenne, 1937 1

J

Figure 7.18

Ophiuroconis bispinosa Ziesenhenne, 1937:228-230, fig. 2a-c.--Barnard and Ziesenhenne, 1961:143. I

Material Examined. California: Santa Maria Basin, Phase II Sta. R-8, 90 m (2, CAMP voucher).--

Off Santa Barbara Id., R/V Velero II1 Sta. AHF 978-39, 38-51 m (18, LACM 39-82.8).--Los Angeles, I[

Santa Monica Bay, R/V La Met Sta. C-3, 60 m (4, LACM 92-53.2).--Mexico: Islas Revillagigedo, off Isla

Socorro, R/V Velero Ili Sta. AHF 924-39, 31-84 m (11, LACM 39-27.5).--Ecuador: Islas Galapagos, off

I

Isla Espafiola, R/V Velero IH Sta. AHF 814-38, 37-73 m (5, LACM 38-80.9).

Description. A small, fragile species. A typical specimen is 3 mm dd with arms 17 mm long.

Disk circular to rounded pentagonal, soft, flat, loosely attached to arms. Entire disk with cover of In

f'me, imbricating scales overlain with rounded granules, some granules at edge of disk elongate. Radial II

shields approximately one-fifth disk diameter, broadly in contact.

Jaws widely separated by broad oral gaps. Oral shields small, longer than wide with sharply pointed I

triangular proximal lobe on oral frame, round distal lobe on ventral interradius. Adorals shields narrow, II

curved, 4 times longer than wide, pointed proximally, in broad contact proximal to oral shield, outer end

abutting first lateral arm plate. Apical and most proximal oral papilla spinelike, pointed; 3-4 distal papillae 1

on jaw flat, rounded, largest one outermost; small papilla at comer of oral slit on fu-st ventral arm plate. II

Granules on oral ft'arne, including few on oral shields, adoral shields, oral plate, proximal arm

segments; outpouched ventral interradii with denser granule cover. I

II

Arm long, slender, distal half tapering.

Dorsal arm plates quadrangular, widest distally, with sharp medial ridge; wider than long at base of 1

arm, becoming much longer than wide near arm tip; successive plates overlapping except at arm tip. |

Lateral arm plates broad, with sharp midlateral ridge, conspicuously etched with parallel microscopic

lines. Plates broadly in contact beneath arm joint, separated above. Two arm spines at distal margin of I

lateral ann plate, smooth, slender, gradually tapering to rounded point; upper arm spine larger, longer than U

arm joint at base of arm. Accessory arm spine minute, comb-shaped, set above the larger spines (Hendler,

prev. unpub.). Single tentacle scale, longer than wide, scoop-shaped, ventral surface convex; relative length •

increasing distally, exceeding length of arm joint near arm tip. |

First ventral arm plate within oral slit small, with medial furrow; second and third ventral arm

plates pentagonal; others more rectangular with curved edges, rounded comers, nearly twice as wide as Ill

long, widely separated by lateral arm plates. U

Color: In living individuals, disk blotched with black-brown, brown, gray, greenish gray; jaw pale

greenish brown tipped with deep brown, granules pale gray. Arms with dark and light bands of tan to pale I

brown, greenish, dark patches on lateral arm plates near base of spines; ventral arm plates dark, tinged with II

khaki; microscopic dark brown specks uniformly scattered on arm plates. Tube feet translucent, oral tentacles

salmon color. Ovaries pink. II

II

Preserved specimens retain considerable pigmentation; as described by Ziesenhenne (1937:230),

"dried from alcohol...disk cream color with irregular light brown markings over the radial shields. Arms

light tan with bands of brown or black, two to four segments wide (usually two), alternating with a light I_

colored area; near disk the light bands axe four segments wide, but distally they axe only half as wide. Lower l1

surface white, with some under-arm plates (usually in pairs) faintly dusky." I

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i Figure 7.18. Ophiuroconis bispinosa Ziesenhenne: A, disk, dorsal view; B, disk, ventral view (LACM 92-53.2;

dd = 2.5 mm).

I

Remarks. In a forthcoming publication on this species (Hendler, in prep.), its familial assignment

will be revised, and the generic and specific names will be changed.

Distribution. Costa Rica, Galapagos, Mexico including Gulf of California, and southern California;

i 5-143 mBi(Ziesenhe o nne, 1937; this report). logy. Bamard and Ziesenhenne (1961) recorded O. bispinosus at 3% of 176 stations taken between

9-243 m depths from Pt. Conception, California to the Mexican border. Individuals from Santa Monica Bay,

I California, epifaunal. collected using a grab sampler, are capable of rapid movement, and the species appears to be

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Family Ophiotrichidae I

Ophiothrix (Ophiothrix)spiculata Le Conte, 1851 I

U

Figure 7.19

Ophiothrix spiculata Le Conte, 1851:318-319.--Liitken, 1856:22.--1859:151 .--Lyman, 1865:167.--1882: /

214, 216, 226, 312, 325.--Ljungman, 1867:331 .--Verrill, 1867: 320, 332-333, 342.--Lyman, 1865: It

13, 153, 167-169.--1875:5.--1880:35.--1882:226.--Ives, 1889:178.--Ltitken and Mortensen,

1899:181.--H.L. Clark, 1902:525-526.--1910:342.--1911:10, 262-263, fig. 129a-c.--1913: 218.--

1915:278-279.--1917: 439.--1923:157.--1940:340.--Koehler, 1907:338.--McClendon, 1909:50- I

51, figs. 38, 39.---Campbell, 1921a:3, fig.7.--1921b:48.--1922:38-39, pl. t, figs. a-i.--May,

1924:274-276, fig. 5a-c.--Ziesenhenne, 1937:225.--Caso, 1961:201-204, fig. 85.--Barnard and i

Ziesenhenne, 1961:133, 143.--Hartman, 1963: 85, 154, 305.--Boolootian and Leighton, 1966:6,

8, fig. 26.--Austin and Hadfield, 1980:151, pl. 49, fig. 10.6.

Ophiothrix (Ophiothrix) spiculata, A.M. Clark, 1966:647. i

Ophiothrix dumosa Lyman, 1860:252-254.--1865:13, 153, 169-171.--Lyman, 1880:35.--Ljungman,

1867:331.--Ven'ill, 1867:270, 326, 329.--Lyman, 1882:226.--Ives, 1889:178.--H.L. Clark, IL

1915:270, pl. 12, fig. 6.--May, t924:274-276, fig.5a-c. I

? Young ophiurid Nielsen, 1932:342-3, figs. 41-42.

Material Examined. California: Santa Maria Basin, Phase I Sta. BRA-025, 64.5-72 m (2 juvenile, I

SBMNH secondary voucher).--Westem Santa Barbara Channel, Phase I Sta. BRC-001,73.5-78 m (2 juvenile,

USNM primary voucher).---Off Santa Barbara Id., R/V Velero III Sta. AHF 975-39, 46-49 m (20, LACM mm

39-79.10).--Mexico: Gulf of California, off Bahia Concepcion, R/V Velero III Sta. AHF 683-37, 22 m (16, i

LACM 37-85.1).

Description. An ophiotrichid of moderate size, growing to 18 mm dd with arms 5-8 times as long. i

A 14 mm dd specimen has 62 mm arms, and one 7 mm dd has 53 mm arms. II

Disk pentagonal with interradial outpouchings. Disk fully covered, radial shields sparsely covered

with rough spinelets and multifid spinules. Radial shields tapering proximally to a point, distally to a short •

straight edge, separated by series of narrow scales, length approximately one-fourth to one-fifth disk diameter.

Jaws with conspicuous central cavity proximal to oral shield. Oral gap quite broad. Oral papillae

absent. Dental papilla cluster at ventral apex of jaw, with central papillae shortest; chisel-shaped teeth high •

in oral cavity. Oral shields wider than long, with medial points on proximal edge, lateral sides lobate;

madreporite larger, more irregular in outline. Adoral shields small, subtriangular, in contact, conf'med to

inner side of oral shield. Ventral interradius covered with spines and minute scales. Large genital scale Jill

evident at distal end of bursal slit. II

Arms tapering to filiform tip; arm joints with narrow waist due to proximally constricted lateral arm

plates. I'

Dorsal arm plates saddle-shaped, raised medially, proximo-lateral edges concave, narrow proximal

end depressed, disto-lateral edges convex, broad distal end upraised. m

Lateral arm plates in contact, with very prominent, flaring spine-bearing ridge. Up to 8 thorny arm :1

spines with thickened, round base, sides of shaft compressed; topmost spine often short, second or third the

longest, 4.5 times length of joint, spines decreasing in length ventrally. Beyond base of arm, most evidently lID

near tip, ventral-most spine with 3 hyaline teeth; terminal tooth largest, curved. Tentacle scale single, small, l

edge bearing sharp tooth.

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I Figure 7.19. Ophiothrix (Ophiothrix)spiculataLe Conte: A, disk, dorsal view; B, disk, ventral view (LACM 37-

85.1; dd = 6.6 ram).

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i Basal ventral arm plates longer than wide, depression at center creating medial ventral ann furrow; proximal plates wider than long, broadest near distal edge, with thickened distal and proximal edges; distal

plates longer than wide. Connective tissue filling gaps between successive plates.

I Variation: See Ophiopholis bakeri above, for description of juvenile. Color: Living individuals with variable color patterns. Generally greenish brown or brown to greenish

yellow above, sometimes with patches or spots of white, gray, orange, or garnet on the disk; the the arms

I swhitish potted, oral blotched region,orotherwise regularly banded disk andwith armsred, pigmented orange, pink, as dorsally, brown,themaroon, arms often or gray. bandedVentral (see McClendon, surface with

1909; May, 1924; Ziesenhenne, 1937, for extensive color notes). Tube feet grayish with microscopic brown

I specks. Ovaries and eggs orange-salmon or orange-peach colored. Preserved specimens fade and change color. The "characteristic blue...coloring of the species" noted

by H. L. Clark results from the degradation of green pigments, a phenomenon seen in other alcohol preserved

I ophiotrichids (Hendier et al., 1995).

Distribution. Northern Chile and Galapagos Islands to Moss Beach, California; intertidal to 211 m

(Liitken and Mortensen, 1899; McClendon, 1909; May, 1924; H. L. Clark, 1911, 1913, 1915, 1917, 1923,

I 1940; Nielsen, 1932; Caso, 1961; Hartman, 1963; Castitlo Alarc6n, 1968; Austin and Hadfield, 1980).

Remarks. The synonymy of Ophiothrix dumosa with O. spiculata has withstood repeated

I examinations (Ives, 1899; May, 1924; Neilsen, 1932). H.L. Clark (1902, 1915, 1917) eventually retracted his proposal that specimens ofOphiothrix galapagensis Liitken and Mortensen are the young ofO. spiculata.

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i 163
m

II

The only sympatric congener of O. spiculata in California waters is Ophiothrix rudis Lyman, and g

Nielsen (1932) provides a detailed discussion of distinctions between the species.

Some features of juvenile Ophiothrix spiculata are evident in Nielsen's illustration of a "Young am

ophiurid from the Gulf of Panama" (Nielsen, 1932: figs. 41, 42), especially the large, smooth central plate

and palmate basal arm spines. However, the differences from O. spiculata in the shape of major plates

suggest that other features are inaccurately rendered, or another species of Ophiothrix is depicted. I

U

Biology. Austin and Hadfield (1980) summarized the natural history of O. spiculata. The species

occurs in shallow water under rocks, in rock crevices, in algae and algal holdfasts and subtidally on hard m

bottom such as shaley pebbles (Barnard and Ziesenhenne, t961). Individuals use the hooked arm spines to i_

secure themselves to the substrate and probably to each other. SmalI O. spicuIata are often seen in shallowwater tide pools. They are found amongst algae, clinging to sponges, in mussel beds, and within interstices

of Macrocystis kelp holdfasts. Scuba divers at the Channel Islands and other islands off Southern California I

and Mexico report great numbers of this species, sometimes 80 individuals/0.1 m2,carpeting the substrate n

in areas of strong currents. These aggregations exhibit the "gregariousness" behaviors reported for Ophiothrix

fragilis in European waters (Hendler, 1991). I

Individuals suspension feed using the mucus coated arm spines and tube feet to strain plankters and

particulate material from the water. They are also capable of gleaning food from the intertidal surface f'tlm

and capturing macroscopic items. O. spiculata lacks the ciliary tracts postulated by MacGinitie (1949). The B

species falls prey to fish including rock wrasse, pile perch, and sand bass, as well as the sea star Astrometis

m

sertulifera (Xantus). I have found the egg diameter to be 0.11 mm, consistent with the production of an

ophiopluteus larval form. •

m

Family Ophiuridae N

OphiosphalmajoUiense (McClendon, 1909) new combination

n

Figure 7.20 U

Ophiomusiumjolliensis McClendon, 1909:36, figs. 2, 3.--H. L. Clark, 1911:9, 20, 111-113, fig. 4Ia-c.-- •

Campbell, 1921b:46.--Astrahantseff and Alton, 1965:1414, 1421.--Thompson et al., 1993:30, 38. U

OphiomusiumjolIiense, H.L. Clark, 1915:330.-- 1917:450.-- 1923"154.--Matsumoto, 1917:285, 291,359.--

Hartman, 1963: 85, 170, 185, 261,305, 320, 383, 401.--Boolootian and Leighton, 1966:6, 10,fig. 21. _

II OphiomuseumjoUiensis, Thompson and Jones, 1987:5.

Material Examined. California: Santa Maria Basin, Phase I Sta. BSS-062, 582 m (3, USNM primary •

voucher; 1 LACM 83-232.1 [as Ophiomusiumjolliensis]).--Western Santa Barbara Channel, Phase I Sta. U

BSS-078, 762 m (1, LACM 83-230.1 [as Ophiomasms sp. A]).--Off Santa Catalina Id., R/V Velero III Sta.

AHF t425-41,512 m (8, LACM 41-205.2).----OffRedondo Beach, R/V VeIero/V Sta. AHF 2788-54, 17 m 1

(1, LACM 54-104.2).--Off Newport Beach, R/V Velero IV Sta. AHF 7352-61,420-521 m (3, LACM 61- l

168.1).

Description. An Ophiomusium species of moderate size. The largest specimens have disk diameters

of 20, exceptionally 30 mm. An individual 15 mm dd has arms over 62 mm long, though relatively shorter U

arms have frequently been reported. Large ossicles appear rough due to numerous microscopic bumps

(EPT). I

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I Disk rounded pentagonal, central region depressed, margins somewhat raised. Central half of disk

covered by thickened, rounded and angular tesselate scales. Primary plates present, slightly larger than

i other scales. Largest scale, about half size of radial shield, at disk edge midway between arms. Radial shields narrowing proximally to acute point, with broad, rounded lobe near arm base; shields approximately

twice as long as wide, length one-fourth diameter of disk. Series of narrow, angular scales between paired

i radial shield Jawss broad. continuing Oral solinttos narrow, base ofnaerarl m,yove searledlapping by oralfirpapillae. st dorsal Oral arm papi platell.ae 6-7 on a side, in continuous

series; lateral papillae subquadrilateral, flattened; apical papilla pointed. Teeth spinelike, sharply pointed,

I dorso-ventrally proximal point, flattened. prominent Oral shaftlike shieldsdistal small, extension nearly fused twice toasinterradius. long as wide, Adoral spearhead-shaped shields as largewithor larger sharp

than oral shield, broad distal edge abutting first ventral arm plate; paired shields with acutely pointed tips

I prominent, meeting proximal to oral shield. Ventral interradii with scales similar to dorsal disk surface. Bursal slit

spanning oral shield nearly to edge of disk; proximal end bifurcate, short branches diverging at

postero-lateral edge of oral shield; slit fringed with continuous series of minute, fiat, quadrilateral scales on

i radial edArms ge andgradually on outer adradial edge. tapering, distal portion filiform; in cross section dorsal surface with shallow medial

ridge, ventral surface flattened, with medial indentation; sharp notched indentations between arm joints.

i Bandasebursal of armslit. withFirst small, threelateral jointsfields beneathof microsc disk broad, opic, bearing angular large granultentacle es between pores; distal distal edgesarmofjoinradial ts lacking shield

apparent pores.

I Dorsal arm plates except firstdiamond-shaped, slightly longer than wide; first several plates touching or nearly in contact, remainder considerably smaller, widely separated by lateral arm plates.

Lateral arm plates large, broad, flat, lacking prominent spine-bearing ridge, covering most of dorsal

I and ventral surface of meeting below and above on first few joints.

arm, except

Ventral arm plate at edge of oral slit minute, situated between distal-most oral papillae, adoral

i shields, and first lateral arm plates. Remaining ventral arm plates with inner edges joining at sharp proximal point, lateral edges narrowing beside tentacle pore, distal edges lobate. Lobes prominent on first three

joints, thereafter decreasing in size, plates becoming diamond-shaped, wider than long. Ventral arm plates

absent near arm tip. Single tentacle scale on first three joints, hinged to lateral arm plate, flat, rounded,

I operculate, sealing tentacle pore; sometimes additional minute scale present on arm plate. Ann

ventral

spines small, approximately one-fifth length of arm joint, affixed to distal edge of lateral arm plate near

ventral ann surface; 1 spine on fourth joint, 2 on next several joints, 3 spines by tenth joint. Ventral-most

I spine dorsal hook-shaped with directed, hyaline, terminal teeth.

tapering, pointed; spines proximally

Variation: The juvenile specimens (1.6-4.3 mm dd) are quite unlike the adults in appearance: dorsal

i surface of disk covered by thickened primary plates, radial shields, and one or two marginal scales; ventral

surface covered by the oral shield and several interradial scales; genital slit papillae absent in specimens

less than 2.8 mm dd. Oral papillae in smallest specimens consist of: apical papilla, 2 small proximal papillae,

i elongate distal papilla; number of papillae increasing during ontogeny. Successive dorsal and ventral arm

plates widely separated by lateral arm plates. Tentacle scales flattened.

Color: The description in Boolootian and Leighton (1966:10) as "dark brown aborally, tan or yellow

I orally gray; ," may have been based on living material. Preserved specimens pale brown, yellowish, to cream or juvenile specimens translucent white.

Remarks. H.L. Clark (1941) proposed the new genus name Ophiosphalma for species with 3 pairs

I of is onproximal that basis tentacle alone pores, that thisrestricting species the is referred gnus Ophiomusium to Ophiosphalmato .specie Thscorrect with onlynominative 2 basal tentacle singularpores. endingIt

for the specific epithet is the neuter form '_iolliense."

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Figure 7.20. Ophiosphalmajolliense (McClendon):A, disk, dorsal view;B, disk, ventralview (LACM41-205.2; Io

dd = 12.0mm). Ophiosphalma variabile (Ltitken and Mortensen): C, disk, dorsal view; D, disk,

ventral view (USNM 19502,syntype, Ophiomusium variabile; dd = 13.9 nun). I

Ophiomusium (= Ophiosphalma) variabile Liitken and Mortensen (1899) is similar to O.joUiense I_

in overall appearance and details such as the distinctive shape of the oral shield (Figure 7.20). Examination

of type specimens of O. variabile (USNM 19502) revealed that it differs in having: marked interradial

indentation of the disk; large radial shields greater than twice as long as broad, widest at mid-length, in 1

contact distally; edges of lateral ann plates straight or concave in dorsal view; conspicuous gap between II

distal-most oral papillae. H. L. Clark (1911) delineated the similarities and diagnostic differences of O.

jolliense and its congeners O. cancellatum Lyman, O. armigerum Lyman, and O. eburneum Lyman. i

Juvenile O. joUiense and the juveniles of an undetermined Ophiurine species both, were

mistakenly identified by survey taxonomists as an Ophiomastus species. The identity of the juvenile

Ophiosphalma was confirmed by comparison with a growth series of O.jolliense. Individuals of the juvenile _l

Ophiurinae species, 1.5 mm dd or smaller, were found at only one locality, Phase I Sta. BSS-069, 927 m

II

(specimens: 1, USNM primary voucher; 1, SBMNH secondary voucher, 1 LACM 83-231.I). Unlike the

juvenile O. joUiense they have four, rather than three evident tentacle pores, and a rudimentary arm comb. I

II Distribution. Cabo San Lucas, Mexico to Oregon, and Japan, generally from 155-1,463 m

(McClendon, 1909; H. L. Clark, 191 t; Hartman, 1963; Astrahantseff and Alton, 1965; Irimuraetal., 1995).

The LACM specimen from 17m reported herein is a juvenile. Records for occurrences off the Galapagos II

I

(H. L. Clark, 1917) and at depths to 4,300 m (Carey, 1990) require confmnation.

166 I
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I Biology. Thompson and Jones (1987) provide underwater photographs of large numbers of

ophiuroids, identified as O. jolliense, inhabiting soft bottom at 698 m, on the lower slope of the San Pedro

i Ba occsuinr off in theSouopthen, ern oCalif thersoinrnia. deprIness diviiodns, ualsorhperched ave the odnistasmall, l partharofdarms structuuprrais es. ed and sinuously bent; some

Thompson et al. (1993) determined that this species is among the most abundant megabenthic

I inverteb They suggested rates trawled its population at the middensity and lower mayslohave pe depincthsreased (300-78 by 0anm)order of Sanof magnitude Pedro and Santa following Monica an E1Basins. Nifio

event.

I Ophiura leptoetenia H. L. Clark, 1911

! I Figure 7.21

Ophiura leptoctenia H.L. Clark, 1911:4, 9, 15, 18-19, 51-55, 58-59, 60, fig. 12a-c.--1913:208-209.--

I 1915:321.-- 1924:291, fig. 1923: 13a-c.--Nietsen, 153.--Matsumoto, 1932:315, 1917:268, 317-321,27figs. 0, 277, 27b, 29a, 354.---Campbell, b, 30a-c.--D'yakonov, 1921b:47.--M 1954:105- ay,

106.--Hartman, 1963: 86, 350, 383, 391,401.--Astrahantseff and Alton, 1965: 1414, 1422.--

I Kyte, 1969:1729, 1731, 1735, 1739.--A. M. Clark, 1965:68.

Material Examined. Oregon: Off Columbia River mouth, R/V Commando, 460 m (3, LACM 64-

i 24Off0.2).---California: Santa Mafia Basin, Phase I Sta. BSS-063, 930 m (1, SBMNH secondary voucher).-- Santa Catalina Id., R/V Velero/V Sta_AHF 9168-63, 1,262-1,298 m (13, LACM 63-203.1).

Description. A small ophiurid, most individuals 8 mm disk diameter or smaller, although H. L.

I Clark (1911) reports a specimen 11 mm dd with arms 30 mm long. Disk round, flat, covered with thin, irregularly shaped, imbricating scales; primary plates larger

than all other scales. Radial shields large, longer than wide, length approximately one-fourth dd, separated

I oaccording r in contactto Nielsen distally. (1932). The disk sometimes bearing scattered small, slender spinelets, numbering up to 25

Oral shields longer than wide with pointed proximal edge and broad semicircular distal edge. Adoral

I shields small, narrowed proximally to oral shield, with long radial extension joining first lateral arm plate. Oral papillae 5 or more, narrow, sharp, spinelike near apex of jaw, broader at jaw base, sometimes with

serrate free edge. Teeth narrow, pointed. Venl_alinterradial area covered with fine scales. Bursal slit extending

I from oral shield to dorsal side of disk; genital scales large; series of minute spines along each scale continuing dorsally, increasing in size, forming arm comb. Arm comb of approximately 12 slender spines, middle ones

longest; opposed by smaller V-shaped secondary comb at base of arm.

I arm plates fan-shaped proximal edge concave, edge convex; some

Proximal dorsal with distal basal

plates wider than long, others longer than wide, becoming triangular toward tip of arm; successive plates in

contact, except near arm tip. ,!i Lateral arm plates wide, broadening ventrally, meeting below arm. Three slender, sharp ann spines;

uppermost longest, length equaling or exceeding joint on first several joints. Tentacle pores large; 9 scales

i surrounding oral pore, 5 onjaw, 4 arising from first ventral arm plate; on first arm joint 1-2 scales on lateral arm plate, 2-3 on ventral arm plate, within several joints tentacle scales reduced to only 1 on lateral arm

plates.

I First ventral arm plate within jaw, bilobed, with medial furrow. Second ventral arm plate triangular, remainder wider than long, with sharp projection from proximal edge; successive plates separated a distance

exceeding their length by lateral arm plates. In large specimens, ventral surface of basal arm segments

i depressed, forming a medial furrow.

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Figure 7.21. Ophiurateptoctenia H. L. Clark: A, disk, dorsal view; B, disk, ventral view (LACM 64-240.2; dd =

6.1 mm). I

|

Variation: Juveniles have: disk covered by primary plates, radial shields, and few additional scales; •

fewer oral papillae than adults; pear-shaped oral shield with narrow triangular lobe merging with oral frame,

rounded distal lobe occupying most of ventral interradius; dorsal arm plates fan-shaped, separated. An arm •

comb first appears in individuals 1-2 mm dd, the inner comb arises later in ontogenesis. On one jaw of one

specimen from the present survey the adoral shield bears a spine, but this condition may be aberrant.

Color: Living individuals reddish brown or gray and khaki above, pale gray below. Eggs orange- i

salmon and pink. Preserved specimens fade to white, yellowish, or gray.

Remarks. Nielsen (1932) specified features that may differ between typical individuals and those

from British Columbia. H. L. Clark (1911) compared and contrasted O. leptoctenia with congeners from •

northern waters, including O. quadrispina H.L. Clark, O. ljungmani (Lyman), O. kinbergi Ljungman, O.

I

sarsii Ltitken, and O. luetkenii (Lyman). i

Distribution. Channel Islands to northern California, Oregon, British Columbia to western Aleutian

Islands and Bering Sea (to 55° 35'N); also, Sea of Japan to Sakhalin Island. H. L. Clark (1911) noted the

peculiar gap between Sakhalin and the Aleutians, depite the Albatross sampling stations in intervening tim

waters. Bathymetric range 122-3,239, with poorly documented reports for 9-16 m (H. L. Clark, 1911, 1913, i

1915; May, 1924; Nielsen, 1932; D'yakonov, 1954; Hartman, 1963; A. M. Clark, 1965; Astrahantseff and

u

Alton, 1965; Kyte, 1969). I

Biology. H. L. Clark characterized O. leptoctenia as one of the most common of the North Pacific I

ophiuroid species, with over 2,000 specimens in the collection he examined. Ophiura luetkenii was a close

runner-up with 658 specimens. On the offshore basin slopes of Southern California, densities ofO. leptoctenia m

reach 44 individualslm 2(Thompson and Jones 1987). On the Central California continental slope, this species I

and Ophiacantha normani Lyman are the dominant ophiuroid species. At 1000-2000 m depths, O. leptoctenia

comprises 50% of the ophiuroids sampled. Its density peaks near 1,500 m where there are in excess of 30 I

individualsl0.1 m2 (Summers, 1993). •

IIR

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I The upper bathymetric limit of the species may coincide with the lower edge of the oxygen minimum

zone off Central California (Summers, 1993). H. L. Clark pointed out the remarkably wide bathymetric

i rheange failedof Oto. note leptoctenia that their , anddepthprovofidedoccurrence information may oben related bottom totemperature temperature,at with samplipopulations ng localities. tracki Honwgever, cold

water masses and ranging from shallow depths in norther localities to deeper water in southern localities.

I specimensSokolova from 2,900-3,140 (1959) characterized m consistedtheprimarily species ofasflocculent omnivorousdetritus, and carnivorous, radiolarian and but diatom gut contents remains,of

and Cumacea; of the latter there was generally only one per ophiuroid, some remaining in good condition.

i The eggs of this species are approximately 0.2 mm diameter, a size consistent with an abbreviated mode of development or an ophiopluteus larval form.

I Ophiura luetkenii CLyman, 1860)

Figure 7.22

I Ophioglypha Liitkenii Lyman, 1860:197-198.--1865:10, 40, 47-48.

Ophioglypha Liitkeni, Lyman, 1875:4.--1880:7.--Koehler, 1907:294, pl. 10, figs. 9-10.

I Ophioglypha liitkeni, Lyman, 1882:35, 76, 310, 313.--Ives, 1889:176.--H.L. Clark, 1901:330.

Ophiura Liitkenii, Verrill, 1867:325.

I Ophiura liitkeni, McClendon, 1909:37-38, figs. 32, 33.--H.L. Clark, 1911:5, 11, 19, 44-47, 55.--1915, 322.--Ziesenherme, 1937:210, 230.--D'yakonov, 1954:102, fig. 38.--Matsumoto, 1917: 267.--

Hartman, 1963: 85,218, 305, 335,350, 365,382.--Kyte, 1969:1729, 1731, 1735, 1739.

I Iiitkini, 1921b:47. Ophiura Campbell,

Ophiura liitkenii, May, 1924:291-294, fig. la-b.--Berkeley, 1927:5.--H.L Clark, 1940:331,343-344.

I Ophiura Liitkeni, Nielsen, 1932:315, 316-18, figs. 27a, 28a,b. Ophiura lutkeni, Barnard and Ziesenhenne, 1961:143.--Boolootian and Leighton, 1966:6, 9, fig. 20.--

Alton, 1972:506-507.---Scanland, 1995:107.

I Material Examined. California: Off Farallon Id., R/V Velero III Sta. AHF 887-38, 68 m (23,

LACM 38-155.5). - Santa Mafia Basin: Phase I Sta. BRC-013, 88.5-100.5 m (1, USNM primary voucher [as

I Phase OphiuraI Sta. lutkenO BSV-042, ); Phase100ImSta. (1, BSS-042 USNM primary , 100 mvoucher (1, SBMNH [as Ophiura secondary leptoctenia voucher ]); Phase [as Ophiura II Sta. R-4, lutke92m])m;

(1, CAMP voucher [as Ophiura luetkeni]). - Off San Clemente Id., R/V VeIero Ill Sta. AHF 1012-39, 101-

I 126 m (nDuesmerous, c LACM 39-116.20). ription. This is an ophiurid of moderate size; specimens as large as 27 mm dd have been

reported. An individual 18 mmdd has arms 90 mm long; one 13 mm dd has 85 mm arms.

I Disk pentagonal, flattened, covered by flat, imbricating scales; primary plates prominent; large radial, interradial, and mid-marginal scales surrounded by small, rounded scales. Radial shields subpentagonal,

somewhat longer than wide, length approximately one-fifth diameter of disk. Paired radial shields separated

I proximally by wedge of scales, distally by series of scales aligned with dorsal arm plates. Arm combs of flat, nearly square scales in tight contact; up to 10 scales in the comb aligned with series of smaller scales

fringing the radial shield and adradial edge of bursal slit.

I Jaws narrowing ventrally, to form shelf within the oral oral expanding a gap. Apical papillae spinelike,

arranged in cluster resembling dental papillae in large individuals; 5 flattened outer oral papillae on jaw

opposed by 3 pointed papillae on In'st ventral arm plate. Teeth slender, with bluntly pointed tip. Oral shield

I large, partially covering ventral interradius, about as wide as long, rounded pentagonal with deeply notched

i 169
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lateral edges. Adoral shields narrow, approximately 112size of oral shield, with proximal edges broadly in •

contact. Ventral interradii covered with thin, imbricating scales. Bursal slit spanning middle of oral shield

to edge of disk, extending above arm base. IlK

Arms tapering markedly toward tip; rounded triangular in cross section. !1

Dorsal arm plates overlapping, fan-shaped to subquadrate in dorsal view, with medial keel. i

arm plates large, covering much of lower arm surface, somewhat figure-8 shaped in lateral Lateral

aspect with dorsal lobe bearing spines on the distal edge, ventral lobe intruding between ventral arm plates.

n

Three arm spines, tapering abruptly to sharp tip, laterally compressed; upper spine distinctly largest and i

longest at base of arm, length equaling 1.5 joints, relative size decreases distally; distal spines of equal size. 1

!111

First ventral arm plate subtriangular with medial furrow; remaining ventral arm plates beneath disk

more than twice as wide as long, proximal edge with protruding medial point, lateral edges narrow, rounded; gum

plates beyond disk edge smaller, thinner; basal plates inflated in large individuals. Tentacle scale on lateral |

arm plate single, small, bluntly pointed; opposing scale on ventral arm plate present in large individuals.

Variation: Nielsen (1932) discussed the ontogenesis of the oral papillae and arm comb in this species. I

The ann comb is absent in individuals less than 2.5 mm dd, making it difficult to distinguish juveniles of II

this species from congeners such as Ophiura sarsii. Specimens of this size and smaller also differ from

adults in having: reduced number of dorsal disk scales, arm spines, and tentacle scales; comb scales with •

free end pointed, dorsal arm plates overlapping only at base of arm. II

Color: Living individuals were described as "salmon red with cinnamon" (May, 1924). Individuals

from Santa Monica Bay, California are greenish-brown and reddish, mottled with black, pink, and white,

the darker pigments sometimes forming a medial stripe on the arm; the lower surface is pale gray or whitish;

the ovaries are salmon colored. The pigmentation pattern varies considerably from one individual to another.

Color is retained for a time in preserved specimens, which have an intense gray dorsal surface, with black •

and white markings, and the ventral surface grayish white. Alcohol preserved material can fade to white.

Distribution. Cedros Island, Mexico, California, Oregon, Washington, British Columbia and Alaska

to Bering Sea, at 9-1,265 m (Lyman, 1882; McClendon, 1909; H. L. Clark, 1911, 1940; May, 1924; Berkeley, n

1927; Nielsen, 1932; Hartman, 1963; Kyte, 1969).

Remarks. H. L. Clark (1911) provides a key distinguishing O. luetkenii from the congeners, O. i

sarsii and O. kinbergi, with which it might be confused.

The epithet "luetkenii" is adopted because the specific name, originally published with a diacritic

mark, is regarded as an incorrect original spelling to be corrected. In such a case, the ICZN indicates "...when Ha

in a name published before 1985 and based upon a German word, the umlaut sign is deleted from a vowel, n

the letter 'e' is to be inserted after that vowel (if there is arty doubt that the name is based upon a German

word, it is to be so treated)" (ICZN 32di[2]). Furthermore, the -ii genitive ending of the original name is m

retained

as required (ICZN 33d). l

Biology. This species occurs in great abundance in suitable habitats. For the southern end of its

range in the Gulf of California and adjacent waters, Ziesenhenne (1937: 230) reported "bushelsful" of O. I

luetkenii taken in a single trawl haul. It has been characterized as "the most abundant deep-sea species in 1

Monterey Bay" (May, 1924:294). However, its distribution may be patchy; Bamard and Ziesenherme (1961)

report it at 15% of 176 stations sampled between Pt. Conception, California and the Mexican border. It m

inhabits sand habitats with varying amounts of silt or mud, but also is found on shell or gravel bottoms |

(Kyte, 1969).

O. luetkenii is epifaunal, as are most of its congeners. A bottom photograph from off Pt. Conception Ill

(Lissner and Benech, 1993:fig. 4.7), shows a number of grayish-colored individuals resting on the surface II

of the sediment, with the ends of their arms upraised. MacGinitie (1949) characterized the species as a

detritus feeder, but it is unlikley to use cilary tracts to transport food in the manner he suggested. According i

170 I
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Il

I

I

I

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Figure 7.22. Ophiura luetkenii (Lyman): A, disk, dorsal view; B, disk, ventral view; C, disk edge, showing blunt,

i close-set scales of the arm comb (LACM 38-155.5; dd= 13.9 mm).

i to Kyte (1969) it feeds on material from the surface of the sediment including protozoans, small animals, and detritus. Scanland (1995) reported a barred sand bass that gorged on O. Iuetkenii off the Orange County

California ocean ouffall, and Dover sole off Oregon, as well as copper rockfish off British Columbia (Murie,

i 1995) also prey on the species.

I

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Acknowledgements I

This study was funded by the Pacific Outer Continental Shelf Region of the Minerals Management Ill

Service, U. S. Department of the Interior, Washington, D. C. J. Blake and A. Lissner offered the MMS

collection for study and arranged for its deposition at LACMNH.

D. Pawson and C. Ahearn (NMNH), R. Woolacott (MCZ), M. Jensen (Zoological Museum, I

Copenhagen), and F.A. Solis-Marin (UNAM, Mexico City) made available specimens in their care. Danish

and German translation was provided by K. Friedmann, Russian by T. Ross, and Japanese by K. White. J.

Dearborn and D. Pawson reviewed a draft of the manuscript. I

Opportunities to collect and study living ophiuroids were provided by personnel of Hyperion Water

Treatment Plant (Department of Sanitation, City of Los Angeles) and Southern California Control Water

Research Project (SCCWRP). SCCWRP also provided partial support for taxomomic study of Amphiodia B

species. J. Watanabe (Hopkins Marine Station) furnished riving material from Monterey Bay. I am grateful u

to all those acknowledged, for their assistance with this chapter. I

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Echinoderm Nutrition. Pp. 169-181. A. A. Balkema, Rotterdam.

I Wolfe, T.J. 1982. Habitats, feeding, and growth of the basketstarAstrophyton muricatum. In: J. M. Lawrence

(ed.), Echinoderms: Proceedings of the International Conference, Tampa Bay. Pp. 299-304. A. A.

Balkema, Rotterdam.

I Ziesenhenne, F. 1937. The Templeton Crocker Expedition. X. Echinoderms from the west coast of Lower

California, the Gulf of California and Clarion Island. Zoologica 22:209-239.

I Ziesenhenne, Expeditions F. 1940. New ophiurans of the Allan Hancock Pacific Expeditions. Allan Hancock Pacific 8:9-59.

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I 8. CLASS ECHINOIDEA

by

I Andrew Lissner _and Debra Hart 2

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l Introduction

Echinoids, commonly known as sea urchins, hearturchins, andsanddollars, are a class of spherical

i or flattened echinoderms containing approximately 900 living species (Figures 8.1 and 8.2). General references utilized for this section are Hyman (1955), Barnes (1980), and Durham et al. (1980). Echinoids occur in all

i oceans and can tolerate salinities ranging from 57 percent to 110 percent seawater. Echinoids may be

i herbivorous or omnivorous. Most regular urchins feed on macroalgae. Irregular echinoids such as heart urchins ingest sediment, digesting the organic material, while sand dollars typically gather particulates

from the substrate or water column, moving food items to the mouth via tube feet.

t I animal's The testname (FigureEchinoidea, 8.1). Spinesmeaning may be"like few ainhedgeh numberog,"anddescribes elongate, theto numerous many spinesandarshort, rangedandarofunction und the

in defense, burrowing, and locomotion. The longer spines are called primaries and the smaller spines are

I secondaries Stalked or miliaries pedicella(Figure ria are 8.1). typical of all echinoids, generally consisting of three articulating jaws.

They can occur over the general body surface (Figure 8.3) and are used for defense, prey capture, and

I cleaning.Echinoids are dioecious, although some examples of hermaphroditism are known (Boolootian and

Moore, 1956; personal observation). Fertilization and larval development are primarily external; however,

I broodi of sandngdollar. is utilized by some cold-water sea urchins and heart urchins, and is present in at least one species

I Regular Urchins

I In the regular or radial urchins, each test plate has only one primary spine (Figure 8.1). The oral

surface is located on the flattened, ventral side, while the aboral surface is moderately to highly arched.

Spines on the oral and aboral surfaces typically are shorter than those on the sides. The oral surface has five

I ambulacral areas containing five double rows of podia that extend in a pentaradiate pattern from the oral to the apical region (Figure 8.1). Podia usually are equipped with terminal disks and are extendable beyond the

spines. Sea urchins use both spines and podia for locomotion.

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1Science Applications International Corporation, 10260 Campus Point Drive, San Diego, CA, 92121

I' 2Science Applications International Corporation, 1710 Goodridge Drive, McLean, VA, 22102

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test I

anal pIate

I

periproct I

primary I

spine

!

I

primary I

spine I

secondary spine

ambulacral spine I

Figure 8.1. Typical aboral morphology of regular echinoid (after Word and Charwat, 1975). I

The mouth is located in the center of the oral surface. The urchin's chewing apparatus, termed •

Aristotle's lantern, is composed of an elaborate system of muscles surrounding 40 calcified ossicles. Five of

these ossicles form hard teeth used to graze food. A membranous area, the peristome, encircles the mouth tim

forming a thickened rim (Figure 8.3). At the edge of the peristome are five pairs of bushy gills. The aboral •

surface also has a membranous central area, the periproct. The periproct may have one to many embedded

endoskeletal plates and also contains the anus. The axis of symmetry extends from the center of the periproct m

to the mouth, and body parts are arranged pentamerousty around the axis. I

182 I
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i anterior ambulacrum

I petaloid

!

internal fasciole

1

i peripetalous fasciole

| L

anal fasciole

|

Figure 8.2. Typic_ abor_ morphology of irregular echinoid(after Word andCharwat, 1975).

!

I The endoskeletal test typically is formed of tightly fired, rigid calcareous plates. However, some

diademids and echinothuriids may have overlapping plates which give their tests a more flexible or leathery

I appearance.

i Irregular Echinoids

In the irregular echinoids such as heart urchins and sand dollars, the periproct and anus are shifted

I fr(Figure om the8.2). centerThisof migration the aboral establishes surface to aanposteri axisoofr area bilateral between symmetry ambulathat cra,isthmore e posterior effectiveinteram for burrowing bulacrum

by these species.

I Heart urchins have a more or less ovoid shape with flattened oral and convex aboral surfaces. The peristome and mouth have migrated anteriorly, and the periproct and anus posteriorly, while the center of

the aboral surface remains at the center of the dorsal surface. Some heart urchins have spines grouped in

I narrow bands (fascioles) on the upper body surface. These smaller spines are associated with numerous cilia that, when beating in unison, create currents that assist in feeding, respiration, and cleaning of the surface.

Podia are degenerate in these species except for areas around the oral and aboral surfaces.

I Sand dollars have flattened bodies with circular circumferences. In these species, the oral and aboral

surfaces are centrally located, with the periproct ventral and situated in the posterior interambulacrum.

I

I 183
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Madreporic

plate

\ Ocular plale I

\ | Genital pore

Madreporite\ I _

\ II _ _ /I Am Genital plate

\ _.__i"!I:,___ _ou_ I

..: ;. _, ._.. :-. _to_: " "" I

__ _.__I'":" _°_,...L """'" C@'"i_ _!_fi_-_ L?'.ii'_ r__ I

... .,.

•.:., ._ ..:, ,.-:

' __' _ ,_: !i__'_'_ I

o,,o / __ ,_®_r_ _._0._i. .-..-!_ "_,,,

Oral tube \

foot Tube,oo, I

Figure 8.3. Oral (upper) and aboral (lower) views of regular echinoid (after Pierce and Maugel, 1987). I

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i Taxonomic History

I The distinctive morphology, color, and relatively large size of most California echinoids have made them the subject of numerous scholarly and popular presentations on their taxonomy and distinguishing

characteristics. Important taxonomic monographs for echinoids worldwide include Mortensen (1928-1951),

I Durham by Grant and and Me Hertlein lville (1957), (1938), and ClarkDurham (1948),etSwan al. (1966), (1952),withJensen Pacific(1974), CoastandtaxDurham a addressed et al.m(1980). ore specific Another ally

classic reference that summarizes key distinctions among Orders and Families of echinoids, as well as

I considerable species-specific information on West Coast taxa is Hyman (1955). A very useful, popular reference, including color plates of shallow subtidal species, is Gotshall and Laurent (1979). Other relevant

summaries of species distributions and their natural history are presented in Ziesenhenne (1941), Hartman

l (1955), (1986). and Important Hartmansynopses and Bamard of taxonomic (1958), Peq collections uegnat (1964) of echinoids , Turnerandet al distributions, . (1968), andincluding Lissner and thoseDorsey from

southern California, are provided in Muscat (1980), Luke (1982), and Maluf (1988).

!

Morphology

I Key featuresthat areuseful in identifying echinoids include the body shape, size and type of priinary

spines, location and shape of petaloids for heart urchins, and, in most fresh and preserved specimens, body

l and include spineanterior color. ambulacrum, Important chara petaloids, cters thatfasciole, must beandrecognizable ambitus (Figures to use 8.1theandtaxonomic 8.2). key in this chapter

The taxonomic key for the echinoids was developed in part with, and the identifications were made

I with the aid of, the monographs of Mortensen (1928-1951). Many specimens also were compared with the echinoid collection of the Allan Hancock Foundation, University of Southern California.

I Glossary

I furrow, prominent most irregular echinoids, containing

Anterior 81nbulacrulll, Antero-dorsal in

tube feet and produced by adepression in the skeletal plates.

I Faseiole. Clusters of spines, usually on the aboral surface, that often are darker colored (e.g., brown or black) in several heart urchin species and which are arranged in characteristic patterns such as ovals,

kidney bean-shaped (reniform), or margin contours.

I most irrePet guallaroids. echinoids, Petal-likpoe,steambul rior toacral the fanurterior rows, amusually bulacrfum. our in number, located on the dorsal surface of

I Methods of Laboratory Examination

I features Idsuchentifiascacolor tion ofanechi d gneneral oids wasshape, accomplis and ahsedtereomi usingcrtheoscounpeaidtoed eyverify e to dist spinineguoirshangro terior ss morpholo ambulacgical rum/

petaloid type and location in small individuals (tess than approximately 10 mm). Microscopic spicules,

I whypochlorite hich can be(bleach), examineared usometimes nder a coused mpouasndtaxomincomic roscopefatures; by dissolving however, athispiece was onot f tube necessary foot into identify sodium

the present species. The first distinction was based on whether the test was circular or ovoid (irregular) in

outline, thereby separating the regular urchins from the heart urchins, respectively. The regular urchins then

i 185
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could be distinguished based on color, which is conservative in preserved specimens and unique (pink or •

whiteish) for the species in the study area. For the heart urchins, the light green to brown color (also

conservative in preservation) indicated that the specimens belonged to either Brisaster or Brissopsis in the i

study area; other genera are either whitish or purple. The arrangement and shape of the anterior ambulacrum l

and petaloids then distinguished the species.

List of Species i

Class Echinoidea I

Order Temnopleuroida

Family Toxopneustidae I

Lytechinus pictus (Verrill, 1867)

Order Echinoida I

Family Strongylocentrotidae

Allocentrotusfragilis (Jackson, 1912) i

Order Spatangoida

Family Schizasteridae I

Brisaster latifrons (A. Agassiz, 1898)

Family Brissidae I

Brissopsis pacifica (A. Agassiz, 1898)

Key to the Species of Echinoidea Collected from the Santa Maria Basin i

and Western Santa Barbara Channel m

(Note: other species may occur in the region but were not collected in the present study) !

1A. Test oval to heart-shaped (irregular urchins) .................................................................................. 3 i

lB. Test globose, circular in outline (regular urchins) .......................................................................... 2 []

2A. Color of test and spines pink; spines distributed evenly over surface of test; spines thin and delicate; I

(note: spine color visible in juveniles greater than 2-3 diameter) ................... Allocentrotus fragiIis

i

2B. Color of test and spines on aboral surface dirty white to dark "blotched" and white to tan on oral •

surface; "skirt" of primary spines around circumference of test; primary spines robust ................ !1

............................................................................................................................... Lytechinus pictus

3A. Anterior ambulacrum deeply notched; subanal fasciole not evident; posterior petaloids typically I

less than half the length of anterior petaloids; color of test and spines light brown to brownish-green

in fresh specimens ................................................................................................ Brisaster latifrons I

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3B. Anterior ambulacrum only slightly notched; prominent, brown to black, reniform subanal fasciole;

posterior and anterior petaloids almost equal in length; color of test and spines light green to brown i

in fresh specimens ............................................................................................... Brissopsis pacifica •

g

186 in
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I Descriptions of Species

Order Tenmopleuridae

i Family Toxopneustidae

I Lytechinuspictus (Verrill, 1867)

Figure 8.4

I Psammechinus pictus Verrill, 1867.

Lytechinus pictus: Clark, 1912.

Material Examined. California: Santa Maria Basin, Sta. BRA-001 (USNM), approximately 70 m (one specimen, 23 mm test diameter); Sta. 002 (SBMNH), 200 m (one specimen, 12 mm test diameter); Sta.

PJ-1 Rep. 1 (note: secondary label in specimen jar lists station as RI-1, presumably in error since it does not

i match a known station code), 145 m (one specimen, 23 mm test diameter).

Description. Test domed aborally and flattened orally, with diameter to approximately 40 ram;

primary spines to 25 mm in length (Tyler, 1944; Durham et al., 1980), commonly observed as a "skirt" of

I around ambitus tubercles of interambulacra small and spines (Lissner, 1980). Primary imperforate (Figure

8.4); pore pairs of ambulacra in 3oblique rows. Test color ranges from off-white to yellowish, particularly

• on oral surface and sides of test, but including grayish to purple "blotches" on aboral surface, extending to

I irregular banding on some spines (Durham et al., 1980; Lissner, 1980).

Biology. Lytechinus pictus is one of the most common megabenthic invertebrates collected on the

I California shelf (Thompson et al., 1987a), occurring in densities up to 20 individuals m2,but noted up to 192 individuals rrd. Populations may appear in vast "herds", usually on sandy bottoms (Dean et aI., 1984) or in

eelgrass beds (e.g., in bays off Baja California). The range of substrate types includes rocks, shells, sand, and

mud (Maluf, 1988). Their diet appears to be omnivorous (Thompson et al., 1987a), but they are best known from

I field studies (e.g., Dean et aI., 1984) that documented extensive grazing effects on juvenile kelp, the extent of

which appears to be mediated by predation from the seastar Pat/r/a m/m_ta (Schroeter et al., 1983).

ofL. pictus Remar . Unpublished ks. Maluf comparisons (1988: page 1(Lissner, 44) notespers. thatobs.) Lytechinus of spicules anaminesus tubeisfeet conalso sidered couldtonot be adistinguish synonym

any differences between these two morphological types, and similarly treat them as synonomous in this

i chapter. __ These conclusio __ _ns are supported further by labspec oraitesoryarerescults ompletely which cross-fert indicatedilethe(Camero two "symp n, a1984). tric"

_i _ Distribution. The species occurs from Monterey,

I CinalifdeoprthniafrotomLathePlainte ta rIsland, tidal toBa3j00a Calif m. Tohrniisais, ranging a much

+ broader depth range than noted in earlier references

I ___ _ (e.g., Gotshall and Laurent, 1979), and likely reflects

the distributions of both L. pictus and L. anamesus, as

combined by the synonomization of these species

i -- _, _ _.. _ (noted in Maluf, 1988; see remarks).

' ' i "

I __ _ Figulre 8.4. Al:_waI view of ulKlisturb_ test alK1spiileS of

Lytechinus pictus (upper) and cleaned test

I (lower)(afterWordandCharwat,1975).

i 187
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Order Echinoida I

Family Strongylocenta'otidae i

Allocentrotus fragilis Jackson, 1912

Figure 8.5 I

Allocentrotus fragilis Jackson, 1912.--Mortensen, 1942, 1943.

Strongylocentrotusfragilis: Clark 1912. I

Material Examined. California: Santa Maria Basin, Sta. 095 (USNM), 198 m (one specimen, 80

mm test diameter, labelled/spelled incorrectly as A. fragile); Sta. R-6, Rep. 1,410 m (one fragmented I

specimen, oral half only, approximately 50 mm test diameter).

Description. Color of test and spines pink, although irregular dark and white spine-banding also

evident, particularly on primary spines; spines thin and delicate (Figure 8.5); test somewhat flattened dorso- /

ventrally. No genital papillae in males or females (Lawrence, 1987). Color retained quite well in dried and

preserved specimens, and evident in juveniles of at least 2-3 mm test diameter. Allocentrotusfragilis is the

only pink sea urchin collected on the shelf and slope of the Pacific Coast. I

I

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!

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I

Figure 8.5. Aboralview of undisturbed test and spines (upper) and cleaned test (lower) of Allocentrotusfragifis

(after Wordand Chatwat, 1975). I

I

188 I
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i Biology. Allocentrotusfragilis often occurs in large aggregations at depths ranging from 50-1200

m, and has been characterized as the most abundant sea urchin off the California coast (Pearse, 1975),

I although borderlandThisompson from 150-480 et al. (19m. 87)Therepspecies orted that oftenthe dominates predominate(abundance depth rangeandwi/orthinbiomass) the southern trawl Ccatches aliforniaat

upper slope sites; densities have been recorded up to 6.2 individuals/mZ(Thompson et al., 1987). No correlation

i of to distribu very fine, tionsiltyand-clayabundance habitat, with but also grainoccurring size is evident, on hardwithsubstrate populations (Maluf,ranging 1988),primarily particularly frominfineecotone sand

regions (Lissner, pets. obs.). Their feeding type is characterized as predator-scavenger, consistent with the

i overlap of their highest abundances with areas of highest infaunal diversity. Allocentrotusfragilis also are common scavengers on food falls (e.g., fish and algae) observed within their depth range (Lissner, pers.

obs.).

I Distribution. The species is reported from Vancouver, Canada, to Isla Carmen, Baja California, ranging in depth from 50-1,200 m (Maluf, 1988).

i Order Spatangoida

Family Schizasteridae

i Brisaster laa'frons (A. Agassiz, 1898)

I Figures 8.6 and 8.7

Schizaster latifrons A. Agassiz, 1898.

I Sch Schizaster townsendi A. Agassiz, 1898. izaster (Brisaster) townsendi: Mortensen, 1907.

Brisaster latifrons: Clark, 1913, 1917, 1937.--Koehler, 1924.--Grant and Hertlein, 1938.--McCauley,

l 1967.

Brisaster to_sendi: Clark, 1917 (according to McCauley, 1967).

i Oppissaster latifrons: Lambert and Thiery, 1924.

Material Examined. California: Santa Maria Basin, Sta. 097 (USNM), 393 m (one specimen, 49

mm length, ratio of anterior/posterior petaloids is 26 mm/19 ram); Sta. 088 (SBMNH), 393 m (one specimen,

I mm length, anterior/posterior petaloids mm); Rep. 3, m (two specimens,

40 ratio of is 25 mm/15 Sta. R3 409

40 and 47 nun lengths, ratios of anterior/posterior petaloids are 27 mm/15mm and 27 mm/17 mm,

respectively).

i Description. Petaloids deeply sunken; anterior petaloids long, reaching almost to margins of test;

posterior petaloids shorter, approximately one-half length of anterior ones (1.0:0.5) (Word and Charwat,

i 1975; Figures 8.6 and 8.7); however, ratios measured in material examined approximately 1.0:0.6. Anterior ambulacram deeply notched. Peripetalous fasciole present (Figure 8.7), ranging even in preserved specimens

from dark brown (thereby contrasting sharply with the lighter brown test) to light brown (blending with the

test color).

I Biology. Brisaster latifrons is a very common heart urchin of the California continental shelf and

slope, occurring in "kilometer-scale herds" on the slope (Thompson, 1993), and sometimes representing the

t d1987). ominant Densities (abundonance and/or biomass) megabenthic invertebrate in these communities (Thompson et al.,

the upper slope can range up to 30 individuals/m 2.Maximum lengths from Thompson et

al. were 73 mm. Individuals burrow up to 15 cm below fine-grained sand or muddy substrates, feeding on

i organic material in the sediments (Nichols et al., 1989).

I 189

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I Distribution. This species is panamic, extending along the entire Pacific coast of North America

(Clark, 1914), with an overall range from the Bering Sea to the Galapagos Islands (Maluf, 1988). The

l reported depth range is from 9-2,817 m.

i Family Brissidae

Brissopsis pacifica A. Aggasiz, 1898

I Figures 8.8 and 8.9

Brissopsis pacifica A. Aggasiz, 1898.

I Toxobrissus pacificus: A. Agassiz, 1904.

Brissopsispacifica: Mortensen, 1907.--Clark, 1917, 1923, 1948.---Grant and Hertlein, 1938.-- Zeisenhenne,

1937.

i Kleinia pacifica: Lambert and Thiery, 1925.

,, Material Examined. California: Santa Maria Basin, Sta. 066 (USNM), 201 m (two specimens, 29

I and 49 mm lengths, ratios of anterior/posterior petaloids 10 ram/10 mm and 13mm/13 ram, respectively,

and one broken specimen, approximately 32 mm length, no ratio determination possible); Sta. 025 (SBMNH),

390 m (one specimen, 38 mm length, ratio of anterior/posterior petaloids 13 mm/10 ram); Sta. PJ-l Rep. 1,

m (one specimen, mm length, anterior/posterior petaloids mm).

145 47 ratio of 17 mm/17

Description. Body and spines light green to brownish in fresh specimens, fading to light green to

i light tan in preserved material O)urhametal., 1980; Lissner, pers. obs.); prominent brown to black peripetalous fasciole (Word and Charwat, 1975) and reniform subanal fasciole are present (Figure 8.7 and 8.9); petaloids

not deeply sunken, anterior and posterior petaloids approximately same length (see material examined

i ratios); anteri Biology. or amb Brissopsispacifica ulacram relativelyis one flushofwith three test echinoids edge. (along withAllocentrotusfragilis and Brisaster

latifrons) that dominate (abundance and/or biomass) their respective communities on the California

c1993). ontinental Highest slopeabundances (Thompson(e.g., et alapproximately ., 1987), sometim20 eindividuals s occurring/m ine, but "kilometer-scale ranging up toherds" 42 individuals/m (Thompson_),

occur at mid-slope depths from 300-600 m. Maximum test lengths from Thompson et al. were almost 60

I mm. The Distri speciesbutiisonchar . This acteristic speciesof t'me-grai ranges from ned substrates, the Channelingesting Islands organic and coast material of southern from theCalifornia sediments.to

Panama and the Galapagos Islands (Feder, 1980), and the western Indo-Pacific (Maluf 1988), at depths

I from 9-3,279 m.

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i Literature Cited

l Barnes, R. 1980. Invertebrate Zoology. 4th Edition, Philadelphia, PA., Saunders. 1089 pp.

Boolootian, R. and A. Moore. 1956. Hermaphroditism in echinoids. Biological Bulletin 111:328-335.

I Cameron, R. 1984. Two species ofLytechinus (Toxopneustidae: Echinoidea: Echinodermata) are completely cross-fertile. Bulletin of the Southern California Academy of Sciences 83:154-157.

i Clark, H. 1914. Hawaiian and other Pacific echini. Memoirs of the Museum of Comparative Zoology, Vol.

XLVI, No. 1, Harvard College, Cambridge, MA.

Clark, H. 1948. A report on the echini of the warmer eastern Pacific, based on the collections of the Velero

I 1II. Allan Hancock Pacific Expeditions 8:225-352.

Dean, T., S. Schroeter, andJ. Dixon. 1984. Effects of grazing by two species of sea urchins,Strongylocentrotus

i franc Macrocystis iscanus and Lytechinus anamesus on recruitment and survival of two species of kelp, pyrifera and Pterygophora caIifornica. Marine Biology 78:301-314.

Durham, J. and R. Melville. 1957. A classification of echinoids. Journal of Paleontology 31:242-272.

l Durham, J., H. Fell, A. Fischer, P. Kier, R. Melville, D. Pawson, and C. Wagner. 1966. Echinoids. Pp.

U211-640. In: R. Moore (ed.), Treatise on Invertebrate Paleontology, Echinoderms. Geological

l Durham,Society of America. New York, NY. J., C. Wagner, and D. Abbott. 1980. Echinoidea: The Sea Urchins. Chapter 11. In: R. Morris, D.

Abbott, and E. Haderlie (eds.): Intertidal Invertebrates of California. Stanford University Press.

I Stanford, CA. Pp. 160-176.

Gotshall, E. and L. Laurent. 1979. Pacific Coast Subtidal Marine Invertebrates. Sea Challengers. Los Osos,

CA.

I Grant. U.S., IV and L. Hertlein. 1938. The west American Cenozoic Echinoidea. University of California,

Los Angeles, Publications in Mathematics, Physics, and Science 2:1-225.

Hartman, O. 1955. Qualitative survey of the benthos of San Pedro Basin, Southern California. Allan Hancock

Pacific Expeditions 19:409-411.

I Hartman,Hancock O. and J.L. PacificBarnard. Expeditions 1958. The 22:56-58. benthic fauna of the deep basins off southern California. Allan

Ill Hyman, L. 1955. The Invertebrates: Echinodermata. McGraw-Hill Book Company. New York, NY. 763 pp.

I_' Jensen, M. 1974. The Strongylocentrotidae (F__chinoidea),a morphologic and systematic study. Sarsia 57:113-

48.

I Lissner, A. 1980. Echinoidea collected during the BLM survey of the Southern California Bight (1975-

1978). Taxonomic Proceedings of the Southern California Coastal Water Research Project, May

1980. 5pp.

I Lissner, A. and J. Dorsey. 1986. Deep-water biological assemblages of a hard-bottom bank-ridge complex

of the southern California continental borderland. Bulletin of the Southern California of Sciences

i 85:87-101.

Luke, S. 1982. Catalog of the benthic invertebrate collections: Echinodermata. Scripps Institution of

i Oceanography Reference Series 82-5:2-24, 44-54.

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Maluf, L. 1988. Composition and Distribution of Central Eastern Pacific Echinoderms. Technical Report •

Number 2, Natural History Museum of Los Angeles County, Los Angeles, CA. 242 pp.

Mortensen, T. 1928-1951. Monograph of the Echinoidea. 5 Vols., 17Parts. Reitzel, Copenhagen, Denmark. l

Nichols, F., Cacchione, D., Drake, D., and J. Thompson. 1989. Emergence of burrowing urchins from

California continental shelf sediments--A response to alongshore current reversals? Bulletin of the In

Southern California Academy of Sciences 86(3): 113-25. a

Pearse, J. 1975. Class Echinoidea. Pp. 621-622. In: R.I. Smith and J.T. Carlton (eds.), Light's Manual: roll

Intertidal Invertebrates of the Central California Coast. University of California Press, Berkeley, 1

CA. II

Pequegnat, W. 1964. The epifauna of a California siltstone reef. Ecology 45:272-283. i

m Pierce, S. and T. Maugel. 1987. Illustrated Invertebrate Anatomy. Oxford University Press. New York, NY.

307 pp.

n

Schroeter, S., J. Dixon, and J. Kastendiek. 1983. Effects of the starfish Patiria miniata on the distribution of i

the sea urchin Lytechinus anamesus in a southern California USA kelp forest. Oecologia 56:141-

147. I

Swan, E. 1952. The Strongylocentrotidae (Echinoidea) of the northeast Pacific. Evolution 7:269-273

Thompson, B. 1993. Megabenthic assemblages of coastal shelves, slopes, and basins off southern California. •

Bulletin of the Southern California Academy of Sciences 92:25-42. II

Thompson, B., G. Jones, J. Laughlin, and D. Tsukada. 1987. Distribution, abundance, and size composition I

of echinoids from basin slopes off southern California. Bulletin of the Southern California Academy i

of Sciences 86:113-125.

i

Thompson, B., D. Tsudada, J. Laughlin, and T. Moylen. 1987a. Biology of the white urchin, Lytechinus ll

pictus, off southern California. Abstract, Western Society of Naturalists, 68th Annual Meeting, II

December 1987.

Turner, C., E. Ebert, and R. Given. 1968. The marine environment offshore from Point Loma, San Diego B

County. California Department of Fish and Game, Fish Bulletin 140:1-85. RI

Tyler, A. 1944. Sexual dimorphism in sea urchins. Anatomical Record 89:573. •

Word, J. and D. Charwat. 1975. Invertebrates of Southern California Waters. I. Select groups of annelids,

arthropods, echinoderms, and molluscs. Southern California Coastal Water Research Project. E1

Segundo, CA. i

Ziesenhenne, F. 1941. Some notes on the distribution records of little-known Southern Californian

echinoderms. Bulletin of the Southern California Academy of Sciences 40:117-120. II

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194 I
i 9. CLASSHOLOTHUROIDEADE BLAINVILLE, 1834

I Including Keys and Descriptions to all Continental Sheff Species from California

by

i Mary Bergen 1

Introduction

I Holothuroids are common members of benthic communities in California. They are found in both

hard- and soft-bottom habitats, from the intertidal zone to abyssal depths.

i with Holothuroids are most often soft-bodied and cylindrical, with tentacles at one end. Their kinship the echinoids and asteroids is not immediately apparent because bilateral symmetry has been

superimposed over pentamerous symmetry and the plates of the exoskeleton have been reduced to microscopic

i part number icles ofin tenta the body cles wall. is a multiple One external of five. clueOn, howe dissection, ver, thatthe a worm-like presence creature of the water is a hvas olothuroid cular system is thatand the

muscle bands in multiples of five clearly show that holothuroids are, indeed, echinoderms.

I Morphology

I Most holothuroids have cylindrical bodies, with the tentacles and mouth at one end and the anus at

the other. Some holothuroids are somewhat U-shaped; in extreme cases, the body wall becomes foreshortened

i so that the mouth and anus are adjacent. Some holothuroids are flattened dorsoventrally, with a distinct soft sole sometimes developed.

The number and shape of the tentacles is an important taxonomic character. Most California

I holothuroids be a simple digitate, have 10 opirn15nate, tentacles; shaped hlike owevaer,shield the aorpodprofusely ous holothbranched urians have(Figure 12 tentacles. 9.1). InTedendro ntacleschirote may

holothurians, the tentacles can be retracted into the body; the area between the tentacles and the body is

I called theTheintrobodyvert.wall is divided by five radii that extend from the mouth to the anus. In each radius there

is a longitudinal muscle, a nerve strand and a water-vascular canal. The water-vascular canal is a vessel

l which canals carries are absent. fluidIntomost the tube cases,feetthere (= podia are three ). In the ventral apodradii ous holothurians that contact the bothsubstrate tube feetandandtwowater- dorsal vascuradii. lar

The portion of the body wall with the longitudinal muscle and tube feet is called the ambulacrum or

i radius. The spaces between ambulacra are called the interambulacra or interradii.

Tube feet may be restricted to the ambulacra or scattered over the entire body wall. In most cases,

interambulacral feet are not present in small holothuroids; rather, they first appear when the animal is about

one centimeter long. The podia are modified into papillae in some aspidochirote holothurians.

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I _SouthernCaliforniaCoastalWaterResearchProject,7t71FenwickLane,Westminster,CA92683.

i I 195

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Figure 9.1. Types of tentacles:A, peltate;B, digitate;C,tentaclewith 2pairsof digits andno terminaldigit, m

I

Internally, the gut is divided into an esophagus, intestine and, sometimes, a cloaca (Figure 9.2). The

esophagus is surrounded by a calcareous ring that provides support and is a point of attachment for the i

radial muscles. In dendrochirote holothurians, retractor muscles, which pull the tentacles into the body, also i

attach to the calcareous ring. w

Immediately posterior to the calcareous ring is the water-vascular ring. Connecting to the watervascular ring is one or more polian vesicles which act as reservoirs. In some groups, e.g., the Molpadiidae II

and the Stichopodidae, reservoirs for the tentacles, called tentacle ampullae, are also present. There is also

one or more stone canals terminating in a madreporite which allow exchange of fluid between the water- il

vascular system and the coelom. •

The gonad is a tuft of tubules which, when ripe, can occupy much of the coelol_ The gonoduct runs

anteriorly along the middorsal interradius to an external opening near the ring of tentacles. Except in etasipods am

and apodous holothurians, the coelom also contains respiratory trees which arise from the cloaca. The II

respiratory trees assist in gaseous exchange.

Some holothuroids, such asthe Psolidae, have a test of large overlapping plates. In others, the plates I

are small, but so closely packed as to form a hard surface. In most cases, with the exception of the Psolidae, II

the plates are embedded in the body wall, and are microscopic in size. These microscopic ossicles (sometimes

also called spicules or calcareous deposits) are a primary character used in identification.

The ossicles assume a wide variety of shapes, including anchors, tables and plates (Figure 9.3). I!

Plates are more or less fiat while tables have a spire attached to a flat disk. The key to species has illustrations

of the different ossicle types. The ossicles in the body wall, tube feet and the tentacles are generally different, i

The tube feet have suppo__ingtables and, in some cases, an end plate. The disk of the supporting table is

curved. The end plate is a sieve-like plate found in the end of the tube foot. The ossicles in the tentacles are

usually rods. They are so variable in size and shape that they are rarely used in identification. I

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Figure 9.3. Types of ossicles: A, anchor; B, anchor plate; C, wheel; D, miliary body or granule; E, cup; F, table; I

G, supporting table; H, plates; I, buttons.

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I Glossary

I AmbulThe acrum (Plural: Ambulacra). The radius. Ossicles. Microscopic calcareous particles, also portion of the body wall that includes known as spicules, embedded in the body

the longitudinal muscle andtube feet. wall which act as a skeletal structure.

I Anchor. An ossicle shaped like an anchor. Phosphatic body. Round to oval granules of iron

phosphate formed from ossicles in

Apodous. Lacking tube feet. Molpadiida.

I Button. A roundthick ossicle. Pinnate. Shaped like a feather.

Calcareous ring. A ring of calcareous plates Papillae. Tube feet lacking a suction cup at the

i surrounding the pharynx; the ring supports

end.

the water vascular ring, and provides

attachment for the longitudinal muscles. Peltate. Shaped like a shield.

! Caudal appendage. A tail-like extension of the Plate. A flat ossicle usually with perforations.

posterior body wall found in molpadiid Podia. See Tube feet. holothurians. it

I Ciliated funnels. Small funnel-shaped structures Polian vesicle. A sac-like protrusion from the

attached to the mesenteries and/or the body water vascular ring that acts as a reservoir

i wall in apodous holothurians, Radiu for the water vascular system. s (plural: radii). Ambulacrum. A

Cup. An ossicle with raised sides shaped like a

longitudinal division of the body wall

i Cuvie cup. having a muscle, a nerve strand and, except rian organs. Sticky tubules that can be in apodous holothurians, a water-vascular

ejected from the anus as a defense against canal. I

I predators, trees, tubules from Respiratory Branching arising

! Digitate. Shaped like a finger, the cloaca that assist gaseous exchange.

I End plate. A sieve-like plate found at the end of a Rod. A cylindrical ossicle. tube foot. Rosette. A small ossicle with short rounded lateral

Interambulacrum. Interradius. The portion of the branches.

t body wall between the ambulacra. Sigmoid body. An ossicle in the form of curved

Introvert. In dendrochirote holothurians, the part rod with both ends recurved. Found in some

I of mainthebody bodywhich wall bet allows ween inver the tensiontacoflesthand e apodous holothurians.

tentacles. Spicules. See ossicles.

I Madreporite. A calcareous structure at the Stone canal. A tube arising from the dorsal side of terminus of the stone canal which allows for the water vascular ring which terminates in

passage of fluid between the water vascular madreporite.

l system and the coelom. Supporting table. A table found in the tube feet;

Miliary granules. Small, compact ossicles found supporting tables usually have a curved disk.

inholothuroids. the longitudinal muscles of apodous Table. An ossicle having a spire attached to a

disk.

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Tentacles. Feeding structures, variously shaped, Tube feet. Cylindrical projections from the body •

surrounding the mouth, wall with suction cups at the end, which aid

Tentacle ampullae. Sac-like protrusions from the in locomotion. Also known as podia. I

water vascular system that act as reservoirs Water vascular ring. A part of the water vascular

for fluid supporting the tentacles, system that encircles the esophagus. 1

Test. A framework of overlapping plates forming Water vascular system. A system of vessels I

an exterior skeleton, filled with fluid that acts as a hydrolic

Tower. A hollow cylindrical ossicle with straight system, providing skeletal support and the I

sides, means to move the tentacles and tube feet.

Wheel. A circular ossicle with spokes.

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Taxonomic History i

The California holothuroids have a long taxonomic history. The literature is scattered, andto date II

there has been no comprehensive study of temperate eastern Pacific holothurians. Between 1829 and 1886, II

Eschscholtz (1829), Brandt (1835), Ludwig (1881) and Stimpson (1857, 1864) described some 16 species

from the Pacific coast, mostly from Alaska and PugetSound. Between 1900 and 1925, Hubert Lyman Clark i

described approximately seven species from California. Between 1935 and 1965, Elizabeth Deichmann l_

1

worked extensively on eastern Pacific holothurians, publishing descriptions of numerous species, including

six species from California. Several authors, including Cowles (1907), Pawson (1967), and Yingst (1972) lit,

have described additional species, n

The taxonomic classification used herein is that proposed by Pawson and Fell (1965). This

classification is based on the shape of the tentacles, the form of the ossicles, and the shape of the calcareous I

ring. In earlier classifications for dendrochirote holothurians, the number of tentacles was given taxonomic

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precedence over the shape of the calcareous ring. Based on examination of recent and fossil species, Pawson

and Fell concluded that the shape of the calcareous ring was more important than the number of tentacles. /I

InPawson and Fetl's classification, the dendrochirote holothurians that do not have a test are divided I

into three families: (1) the Cucumariidae, which have a simple calcareous ring, (2) the Phyllophoridae,

which have a ring with long projections divided into many small pieces, and (3) the Sclerodactylidae, which III

have undivided processes (Figure 9.4). Unfortunately, California species do not exactly fit into this scheme U

because in species with a calcareous ring with long projections, the projections are divided into two or three

big pieces. In this chapter, I have changed the keys and descriptions to reflect this difference, assuming that I

the California species belong in the Phyllophoridae. Further study is needed to determine if the California I

species should be put into a different family.

The generic classification used for dendrochirote holothurians in this chapter is that proposed by [11

Panning (1949) as modified in Panning (1964, 1966, 1971). In a postscript to his 1949 paper, Panning noted II

that Havelockia has processes divided into several large pieces, and moved this genus into the Sclerodactylidae.

Given that all the California species in this group, not just Havelockia, have this characteristic and that the lilt

species clearly do not belong in this family, this change in the classification has not been followed. II

In his papers, Panning assigns California species to genera. However, because he based his

assignments on published papers, some species were misplaced. In this paper, I have reassigned the species t

as best I can. However, the relationship of California species to the generic classification and the genetic II

classification itself are in need of additional study.

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I Figure 9.4. Types of calcareousrings: A, simple ring;B, ring with short processes; C, ring with longprocesses.

I The classification of apodous holothurians used herein is that proposed by H.L. Clark (1907). In

addition, the genus Rynkatorpa, proposed by Rowe and Pawson (1967) is included.

I Heding (1928) published a paper on the Synaptidae (apodous holothurians with ossicles as anchors

or wheels) including descriptions of several new species from California. He modified the classification of

the Synaptidae by elevating the importance of two characters: (1) the number and structure of ciliated

i funnels attached to the inside of the body wall or the mesenteries and (2) the of the

shape miliary granules,

small ossicles found in the body wall and longitudinal muscles. He also differed with H.L. Clark (1907) on

synonyms for several species. Since it has not been possible to sufficiently study Heding's characters and

I the identity of the species is unclear, I have not identified the Synaptidae in this chapter to species. More information about nominal species and their descriptions is given in the generic description.

I Dissection and Slide Preparation

I The calcareous and other internal be the lateral wall

ring organs can exposed by cutting along body

from the tentacles to the posterior end. By making the cut along the side of the specimen, dorsal mesenteries

and other important organs are left intact.

i In dendrochirotids when the tentacles are not retracted, it may be necessary to cut open the introvert

to expose the calcareous ring. When the tentacles are retracted, the introvert forms a tube between the top of

the body wall and the calcareous ring. The tentacles are inside the calcareous ring. To see the tentacles, the

ring must be cut and laid open.

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To make a temporary slide of the ossicles in the body wall, introvert or tube feet, dissolve a piece of B

the tissue in household bleach. For specimens with thin tissue and relatively few ossicles, a small piece of

tissue can be put directly on the slide and a drop or two of bleach added. When the tissue is dissolved, and I.

no more bubbles are being formed, a cover slip can be added and the slide examined on the microscope. For !1

specimens with thick tissue or abundant ossicles, it is easier to dissolve the tissue in a spot plate or watch

glass. Then a sample of the ossicles can be transferred to a slide with a pipette. El

To make a permanent slide, the bleach must be washed from the ossicles. When there are relatively II

few ossicles, it is best to allow a drop of bleach with ossicles to dry in the center of the slide. Then a drop of

distilled water is added to each side of the drop of bleach so that the distilled water barely merges with the

bleach. This procedure dilutes the bleach without removing the ossicles from the center of the slide. The E

slide is allowed to dry, preferably on a drying plate. A squirt bottle with distilled water is then used to wash

the edges of the slide, removing bleach but not the ossicles. This procedure is repeated until the bleach is

sufficiently diluted that mounting medium and a cover slip can be added. II

When there are sufficient ossicles, it is easier to wash them in a small vial. Distilled water is added

and then decanted with a pipette, with care being taken to keep the ossicles at the bottom of the vial. When

sufficiently clean, the ossicles are transferred to a slide in a drop of water. When the slide is dry, mounting II

medium and a cover slip added.

h

Ossicles in some species degenerate as the animal grows. Adults of some species, such as Caudina E

arenicola, have no ossicles at all. In molpadids, the ossicles become converted into phosphatic bodies, 411

round to oval granules of iron phosphate that are red in color. However, ossicles can usually be found in the

tip of the tail. In chiridotids, ossicles are often scattered in bundles on the surface of the body wall; the •

bundles, if present, may be found by scanning the body wall with a dissecting microscope.

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Biology I

are largely benthic, although some deep-sea species are known to make swimming I

Holothurians

excursions. They occur in both hard- and soft-bottom habitats. Some are members of the epifauna, others

nestle in cracks or under rocks and others burrow or form tubes in soft substrates. tim

Many species of holothurians are filter feeders. These species extend tentacles covered with mucus I

into the water. Then, from time to time, they insert a tentacle into the esophagus, removing the mucus with

its meal of planktonic organisms and detritus. Other species, such as Parastichopus, sweep the tentacles m

across the substrate, picking up detritus. These species also use the esophagus to remove food from the |

tentacles. Some species, such asLeptosynapta, burrow through mud and sand, engulfing the substrate whole.

Other species, such as Molpadia, feed head down in a burrow. These feed on mud at the bottom of the lib

burrow, passing the mud through the intenstine to be deposited in a mound on the surface. I

Most holothurians are dioecious broadcast spawners. Many of these species raise the anterior portion

of the body away from the substrate while spawning; spawning may be accompanied by swaying or tentacle m

waving (McEuen, 1988). Some species, such as Pseudocnus lubricus and Cucumaria pseudocurata, brood I

eggs under the ventral surface of the body. Other species such as Pachythyone retain the eggs, giving birth

to the young through special pores or by rupture of the body wall. 'm

As a defense against predators, some species of aspidochirotid holothurians have sticky tubules, the II

Cuvierian organs, that are ejected through the anus when the animal is disturbed. Other species of holothurians

will eject the whole viscera through the mouth or anus when disturbed. A new viscera is regenerated in a

few weeks. II

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I Key to Families, Genera and Species from Shallow Water

i and Continental Shelf Depths of California

I IA. Tube feet and papillae absent .......................................................................................................... 2 lB. Tube feet and/or papillae present .................................................................................................... 8

I 2A. Body cylindrical; ossicles as anchors and plates or wheels or sigmoid bodies, but not as tables or

phosphatic bodies ....................................................................................... Order Apodida .......... 3

I 2B. Body perforated moreplates, or lessanchors taperingsometimes posteriorlypresent, to a caudal wheelsappendage; and sigmoidossicles bodiesasabsent, tables,phosphatic fusiform rods bodiesor

often present ............................................................................................... Order Molpadiida ...... 6

i 3A. Ossicles as anchors and anchor plates ........................................................ Family Synaptidae ..... 4

3B. Ossicles as wheels ........................................................................................................................... 5

4A. Anchor plates elongate, irregular in outline ........................................................ Genus Rynkatorpa

i 4B. Anchor plates oval, regular in outline Genus Leptosynapta

I 5A. 5B. Wheels Wheels with with 86 or more spokes .................................. Family Myriotrochidae: Genus Myriotrochus spokes ........................................................... Family Chiridotidae: Genus Chiridota

I claw-shaped; phosphatic present; appendage as with round

6A. Tentacles bodies ossicles in caudal tables

to fusiform disk or as anchors and racquet-shaped plates .................................................................

i 6B. Tentacles ..................................................................... with 2 pairs of digits and no terminal Familydigit;Molpadiidae: phosphatic bodies Genus absent Molpadia............................. : M. intermedia

.................................................................................................................... Family Caudinidae ..... 7

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7A. Pseudomesenteries absent; ossicles as large spiny plates with large perforations ............................

I ......................................................................................................... Genus Itedingia: H. californica 7B. Pseudomesenteries present; ossicles as perforated plates or rods or, injuveniles, small tables; ossicles

often reduced or entirely lacking ........................................................ Genus Cau_dina:C. arenicola

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8A. Tentacles peltate, ventrally placed; retractor muscles absent; tube feet restricted to ventral surface;

i dorsal surface with papillae ...............................................................................................................

................................ Order Aspidochirotida: Family Stichopodidae: Genus Parastichopus ......... 9

8B. Tentacles digitate; retractors present ............................................. Order Dendrochirotida .......... 11

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9A. Dorsal papillae mostly large and conical; mean number of spines on spire of tables from dorsal li

body wall greater than 12 .............................................................................................................. 10

9B. Dorsal papillae mostly small, black tipped in live specimens; mean number of spines on spires of

tables less than 12; diameter of table disks less than 75 _rn .................. Parastichopusparvimensis i

10A. Mean diameter of table disks greater than 100 _rn; yellowish white when alive ............................. I

..................................................................................................................... Parastichopus johnsoni

10B. Mean diameter of table disks less than 100 lain;mottled brown to reddish brown when alive ........ i

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................................................................................................................ Parastichopus californicus

11A. Body flattened, bilaterally symmetrical; distinct sole present ................... Family Psolidae ....... 12 I

llB. Body cylindrical; distinct sole absent ........................................................................................... 15

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12A. Dorsal scales present, but covered by a layer of deposits including hour-glass shaped ossicles; tube

feet present on the dorsal surface ............................................................... Genus Lissothuria .... 13

12B. Dorsal scales naked; tube feet absent from the dorsal surface ................... Genus PsoIus ........... 14 II

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13A. Dorsal scale covering incomplete; dorsal scales overlain with towers, and hourglass-shaped ossicles; •

ossicles of sole perforated plates with raised marginal projections and cups ...Lissothuria nutriens !1

13B. Dorsal scale coveting complete; dorsal scales overlain with hourglass-shaped ossictes; towers absent; I

ossicles of sole large heavily knobbed plates, which tend to become 2-layered, smaller plates with II

knobbed margin and hourglass-shaped ossicles ........................................ Lissothuria deichmannae

14A. Approximately 12 scales between oral and anal scales; scales granulated; ossicles in sole plates I

with 2-4 holes and marginal projections, often reduced with age ......................... Psolus squamatus

14B. Approximately 4 scales between oral and anal scales; scales smooth; ossicles in sole oval buttons i

with multiple perforations, often forming a reticulated mass .............................. Psolus chitinoides IP

15A. Calcareous ring simple, lacking posterior processes ............................ Family Cucumariidae ..... 16 I

15B. Calcareous ring complex, with paired or unpaired posterior processes ........................................ 21

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16A. Ossicles thin, flat, sometimes with spines .................................................. Genus Cucu_aria .... 17

16B. Ossicles three-dimensional, knobbed, sometimes with a dentate handle .. Pseudocnus: P. lubricus I

17A. Ossicles with spines ...................................................................................................................... 18

17B. Ossicles without spines ................................................................................................................. 19

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I 18A. Ossicles mostly rounded, with large holes and spiny margins; single stone canal ...........................

........................................................................................................................... Cucumaria piperata

i 18B. Ossicles mostly oblong; perforations small, margin of ossicle smooth; multiple stone canals ........

............................................................................................................................ Cucumaria miniata

I 19A. Tube feet not restricted to the ambulacra; maximum length 2.5 cm; dorsal side black, ventral side

yellowish; ossicles mostly small rounded buttons with 4-8 holes ....................... Cucumaria curata

i 19B. Tube feet msWicted to the ambulacra ............................................................................................ 20

i 20A. B. Ossictes Ossicles mostly round small buttons plates with with154-8orholes; more maximum perforations;sizemaximum 10-15 cmsize............. 2-3 cmCucumaria ........................ salma

................................................................................................................... Cucumaria pseudocurata

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21A. Processes short, entire ............................................................ Family Sclerodactylidae ............... 22

I 2lB. Processes long, in pieces ........................................................ Family Phyllophoridae ................. 23

22A. Tube feet restricted to ambulacra; ossicles knobbed plates and baskets or cups ..............................

I ................................................................................................ Genus Eupentacta: E. quinquesemita

22B. Tube feet not restricted to the ambulacra; ossicles reticulated plates and regularly-knobbed 4-holed

I buttons; no baskets or cups present ...................................................... Genus Pachythone: P. rubra

23A. Ossicles in body wall predominantly plates .................................................................................. 24

I 23B. Ossicles in body wall mostly tables or a mixture of tables and plates .......................................... 28

I 24A. Lozenge-shaped plates, 0.05-0.15 mm long, common; supporting tables in tube feet with well- developed spire ............................................................................................... Pentamera lissoplaca

24B. Lozenge-shaped plates rare or absent ............................................................................................ 25

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25A. Spire on supporting tables in tube feet well developed ................................................................. 26

I 25B. Spire on supporting tables in tube feet reduced or lacking ........................................................... 27

26A. Spire on supporting tables with 4 pillars ending in a spiny mass; body wall ossicles plates, usually

I oblong, often covered with spines and knobs ..................................................... Stolus trachyplaca

26B. Spire on supporting tables with 2 pillars, sometimes elongate and flattened; body wall ossicles

i large, irregularly-shaped plates, 0.2-0.4 mm in diameter; star-shaped tables occasionally present. .............................................................................................................. Pentamera pseudocalcigera

I 27A. Body wall ossicles oval buttons with 2 cenwal and up to 8 marginal knobs ........ Thyone montereyensis

27B. Body wall ossicles flat, irregularly shaped plates and smaller, more delicate rectangular plates,

often with two enlarged central holes; buttons sometimes present ........................ HaveIockia benti !

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28A. Lozenge-shaped plates common .................................................................... Pentamera lissoplaca I_

28B. Lozenge-shaped plates absent ....................................................................................................... 29

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29A. Discs on tables oblong, regular in outline; spire long with 3-6 crossbars ..... Havelockia charlottae

29B. Discs irregular in outline; spire short with 1-2 crossbars .............................................................. 30 l

30A. Teeth on spire of supporting tables bifurcate ...................................... Pentamera pse_dopopulifera

30B. Teeth on spire of supporting tables not bifurcate .......................................................................... 31 !

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31A. Four-holed tables common; star-shaped tables absent ........................................... Havelockia benti _l

3lB. Four-holed tables not common; star-shaped tables present ........................... Pentamera populifera U

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Descriptions of Species I

As noted earlier, there has been no comprehensive review of the systematics of northeastern Pacific

holothurians. Information about holothuroids is scattered in a variety of publications, many of which are II

difficult to obtain. For this reason, I decided to expand the scope of this chapter to include all species in II

California, either reported in the literature or personally identified, that are found from the intertidal zone to

adepth of 200 meters. In addition, some species not reported from this area are included. Ihave not included

groups such as the Elasipodida or the Synallactidae that are found only in deep-water habitats. It

The following descriptions are based on an extensive review of the literature, including a manuscript

on the eastern Pacific holothurians that was being written by Elizabeth Deichmann at the time of her death. 1

This manuscript was kindly given to me by the American Museum of Natural History. U

The descriptions are also based on examination of the large collections of specimens in the U.S. l

National Museum of Natural History (USNM) and the Allan Hancock Foundation (AHF) collection which i[

is now housed in the Natural History Museum of Los Angeles County. I also examined material from the

American Museum of Natural History(AMNH), Museum of Comparitive Zoology, Havard (MCZ), the m

Santa Barbara Museum of Natural History (SBNMH), the British Columbia Provincial Museum (BCPM)

and Scripps Institute of Oceanography (SIO). In addition to the material collected in Bureau of Land

m

Management (BLM) and Minerals Management Service (MMS) surveys, I have included material collected

by individuals and groups, including Marine Ecological Consultants (MEC), the Southern California Coastal mE

Water Research Project (SCCWRP), Pacific Biomarine (PBM), and myself (MB).

Order Apodida Brandt, 1835 I

Diagnosis. Body cylindrical, worm-like. Body wall smooth, rough or with warty surface. Tube feet I

absent. Anal papillae, tentacle ampullae, and respiratory trees absent. Tentacles 10-25, pinnate or digitate in

form. Ossicles include wheels and anchors, may be reduced with age. II

Remarks. Two families of Apodida, the Synaptidae and Chiridotidae, are common in shallow II

water soft-bottom habitats in California. I have included a third family, the Myriotrochidae, in the key. This

group primarily occurs in deep water; however, occasional specimens axe taken in shallower water. I

206 I
I

I Family Synaptidae Burmeister, 1837

Diagnosis. Tentacles cylindrical or terete, not widened distally, either pinnate or digitate in form.

I Ossicles as anchors and anchor plates.

Remarks. Only one genus (Leptosynapta) of Synaptidae is known to occur in California. However,

• I have included the genus Rynkatorpa Rowe and Pawson, 1967 in the key because there is a possibility that

I Rynkatorpa duodactyla (H.L. Clark, 1907) may occur species distinguished by

in California. This is the

irregular shape of the anchor plates and the presence of two digits on the tentacles. It is known to occur from

the Aleutian Islands to Grays Harbor, Washington. !

Genus Leptosynapta Verrill, 1867

I Type Species. Holothuria inhaerens O.F. MiJller, 1776, designated by Verrill (1867).

i Leptosynapta spp. Figure 9.5

I Mugu Lagoon, (1), (BLM sandy beach, 1977,

Material Examined. California: intertidal Winter

Sta 4A); Corona del Mar, intertidal (1), (BLM sandy beach, Winter 1977, Sta 13c); Pt Dume, 60 m, (3),

(SCCWRP R15-60); Pt Arguello, (1:0.3 cm) (MMS CAMP 1-1, PJ7); Pt Arguello, (1:0.5 era), (MMS

I CAMP 1-2, R-I); Pt Arguello, (13), (MMS CAMP 1-2, R-8); Pt Arguello, (2), (MMS CAMP 2-3, R-8); BLM 23201; BLM 23206; BLM 23149; USNM BSS-63; SBMNH BSR-79; (MMS Camp 3-4, R1); (MMS

CAMP 3-1, R-8).

I Diagnosis. Tentacles pinnate, 10-13 in number. Sense organs on tentacles as small cups, pigment

spots absent. Stocks of anchors with f'me teeth, not branched. Anchor arms usually with small teeth, vertex

of anchor smooth. Anchor plates oval or somewhat elongated, usually with 6-7 large dentate holes; several

I smaller smooth present at narrow of plate.

holes end

Biology. Species of Leptosynapta live in sandy or muddy habitats. They are thought to burrow

i beneath the surface, engulfing sediment as they move. However, during the spawning season in Puget Sound (November-December), McEuen (1988) found Leptosynapta in vertical burrows in aggregations of

2 to 4 individuals. Fertilization and early development occurs within the ovary in Leptosynapta.

I H.L. ClarkRema (I 907) rks. recognized As noted inonlytheoneIntrodu species ction, fromthetheidenwest tiycoast of speci of North es in America: this genusLepistosynapta uncertain ainhaerens t present.

He considered Synapta albicans Selenka, 1867, a synonym of inhaerens, a circumpolar species occurring in

I Europe, Heding the western Atlantic and eastern Pacific. (1928) considered the European, western Atlantic and Pacific forms of inhaerens to be

distinct species. However, he did not believe that the Pacific forms could be referred to albicans; rather he

I described them as a new species: L. clarki. Heding recognized five species of Leptosynapta from the eastern Pacific: albicans, clarki, lens,

roxtona, and transgressor. He considered lens and albicans to be closely related, possibly the same species,

I being differentiated primarily by the presence (albicans) or absence (lens) of rods in the tentacles.

The two provisional species designations (Sp. A and Sp. B) given for specimens of Leptosynapta

are based on the size of the anchors: Species A has relatively large anchors compared to Species B. However,

i anchor size be variable; sometimes the anchors in the anterior and of the

Can quite posterior portions body

are distinctly different. A thorough study of many specimens from a variety of locations is needed to establish

the range of variability in species characteristics and determine the identity of the species of Leptosynapta

I occurring in California.

I 207
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I

i

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I

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0 ,i

Figure 9.5. Leptosynapta spp.: A, anchors; B, anchor plates; C, miliary granules. .t

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208 I
I

I Family Chiridotidae Ostergren, 1898

I Ossicles Diagnosis. as 6-spokedTentacle wheels;stalks sigmoidshort,or widened C-shapeddistally rods may , withor3-1may0 digits not beonpresent; each sideossicles of the sometimes distal end.

lacking.

I Genus Chiridota Eschscholtz, 1829

I Type Species. Chiridota discolor Eschscholtz, 1829, by original designation.

Chiridota spp.

I Figure 9.6

Material Examined. California: BLM 24242; Santa Catalina Basin, 1300 m (NOFAC Sta. 1).

U Diagnosis. Twelve tentacles present, rarely 13-14. Polian vesicles numerous. Ossicles as 6-spoked

wheels collected into papillae; sigmoid ossicles absent; small curved rods with enlarged ends may be present;

I tiny oval particles may be present in the longitudinal muscles.

Remarks, As noted in the Introduction, the identity of species in this genus is uncertain at present.

In his discussion of the genus Chiridota, Clark (1907) notes that the characteristics used to differentiate the

I species (the number and arrangement of wheel papillae, the number and length of digits on the tentacles, the

presence or absence of miliary granules) are highly variable. He provisionally identified two species as

ranging from Alaska to Baja California: C. discolor Eschscholtz, 1829 and C. laevis (Fabricius, 1780).

I However, he noted that the identifications were tentative and that additional species may be present.

I

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I 100pm

I Figure 9.6. Chiridota spp.: Wheels from body wall.

I 209
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Heding (1928) also recognized the variability of many characteristics used to identify species in the

genus Chiridota. However, he believed that the size, arrangement and shape of the ciliated funnels, the

shape and arrangement of miliary granules in the body wall and the shape of the calcareous ring could be I

used to differentiate species. II

Heding (1928) described two new species from the north eastern Pacific: C. pacifica, from La Jolla,

California, and C. nanaimoensis, from Nanoose Bay, British Columbia. He notes that (7.pacifica is closely II

related to C. albatrossi Edwards, 1907 and C.laevis, both of which occur in the northern Pacific. He indicates n

that the specimens identified by Clark (t907) represent three species: C. albatrossi, C. pacifica, and C.

laevis. Since there is no discussion of C. discolor, the status of this species is unclear. B

II Heding (1928) differentiates C. pacifica from C. albatrossi and C. laevis by the shape and size of

the ciliated funnels, the shape of the madreporite, and the presence or absence of rods in the anterior portion

of the body wall. A study to determine the range of variability in these characters is needed in order to I

determine the status of these species. U

Order Molpadida Haeckel, 1896 i

Diagnosis. Body stout, tapering to a more or less distinct tail Fifteen tentacles present. Respiratory g

trees and tentacle ampullae present. Ossicles may include tables, anchors, fusiform rods, or perforated

plates. Red phosphatic ossicles often present. I

Family Molpadiidae Miiller, 1850 il

n

Diagnosis. Tentacles claw-shaped or with lateral digits. Tentacle ampullae long. Ossicles in tail

tables or fusiform rods. Phosphatic bodies often present. 1

II

Genus Molpadia Risso, 1826

Type Species. Molpadia musculus Risso, 1826, by original designation. I

Diagnosis. Tentacles with 1, sometimes 2, pairs of digits and a terminal digit. Ossicles as tables,

plates or fusiform rods. In young individuals, racquet-shaped plates and anchors sometimes present, n

Phosphatic bodies usually present. U

MoO_zdia intermedia (Ludwig, 1894) I

Figure 9.7

i Trochostoma intermedium Ludwig, 1894:161, pl. 16, figs. 7-21.

Molpadia intermedia H.L. Clark, 1907:33, pl. 12,figs. 5-15.--Heding, 1931:279.--Deichmann, 1937:174. II

Material Examined. West coast tropical America: (syntype, USNM 18202).---Califonia: USNM

Albatross 2937, 33°04'30"N, 117°42'00"W, 464 fms, mud (1:6.5 cm); USNMAlbatross 4475, 9.7 mi S15°W

Pt. Pifios lighthouse, 58-142 fms, mud (3:7.3-7.8 cm); USNM Albatross 4509, 8.6 mi S13°W Pt. Pifios N

lighthouse, 152-286 fms, mud (1:10 cm); MEC, Pt. Arguello (1); MBC, N side of Coronado Canyon (1); U

MBC, offPt. Loma, 187 m (1:3.5 cm); Chambers, Pt. Arguello (1); SCCWRP, Pt. Dume (1); CAMP 1, RI

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I 1 (2:0.5, 2 cm); PJ-1 (1:0.5 cm); CAMP 1-2, R-1 (1:2 cm); R-4 (1); R-8; CAMP 1-3, R-5 (1); CAMP 2-3,

R-1 (2); R-9 (1); CAMP 2-4, R-2 (1:5.5 cm); R-1 (1:2.4 cm); R-5 (1:1.8 cm); CAMP 2-5, R-I; R-8 (1);

CAMP 3-1, R-1 (1); CAMP 3-4, R-1 (2); USNM BSS-058; SBNMH BSS-102. Ii Description. Body stout, globose, tapering to distinct caudal tail, brown in smaller specimens,

mottled magenta in larger specimens. Tentacles with unpaired terminal digit. In small individuals ossicles

I iwall n bodconve y wallrtedtabtles o pho andsphatic racquet-sha bodiepes.dOplat ssicles es anindcaudal anchors,appenda sometimgeestabaslesrosewitthtes.roun Witdh toage, fusoifssicles orm diinsk.body

Biology. Molpadia intermedia is one of the most common holothurians in muddy bottoms in

I California from 60 to at least 900 m. The species lives head-down, engulfing mud at the bottom of the burrow. The mud is passed through the gut and deposited in a mound on the surface of the substrate. In

Puget Sound, M. intermedia spawns during the winter (McEuen, 1988). To spawn, the adults emerge from

I the watersubstra in onet,short raisinburst. g the anterior end of the body 4-8 cm above the substrate. Eggs are expelled into the

Type Locality and Type Specimens. West coast of tropical America, syntype (USNM 18202).

i Distribution. From Alaska to South America, 60-900 m.

i Family Caudinidae Heding, 1931

i Diagnosis. Tentacles with 1-2 digits, no terminal digit. Skin may show coloration, but does not

contain phosphatic bodies. Ossicles tables, plates, small crossed cups or irregularly-shaped bodies.

i Genus Caudina Stimpson, 1853

Type Species. Caudina arenata Gould designated by Stimpson (1853).

I Diagnosis. Body stout, tapering toward the posterior end. Fifteen tentacles with 2 pairs of digits, no terminal digit. Pseudomesenteries present. Ossicles astables, or plates; ossicles sometimes lacking. Phosphatic

bodies absent.

I

Caudina arenicola (Stirapson, 1857)

I

Lioso_a arenicola Stimpson, 1857:525.

i Molpadia Trochostomaarenicola arenicolaH.L.The_l, Clark,1886. 1907:33, 171, pl. 7, figs. 1-2.--Ricketts and Calvin, 1968:292, fig. 219.

Caudina arenicola Deichmann, 1938:112, text-fig.

i Material Examined. Off Pt. Loma, California, 195 ft, (1) (City of San Diego, Sta. B-3).

Description. Large form, with stout body tapering slightly inposterior end. Skin smooth, slippery,

I with yellowish-brown mottling. Ossicles in posterior end small perforated plates, with 2-4 holes, and branched rods. Ossicles in juveniles small tables. Ossicles reduced with age, often absent.

Remarks. Because of its shape and color, this species is commonly known as the "sweet potato".

i Surprisingly, little is known about its distribution or habits.

very

Type Locality and Type Specimens. San Pedro, California, types missing.

I Distribution. In sandy areas from San Pedro south to northern Baja California.

I 211
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II

Figure 9.7. Molpadia intermedia: A, tables from caudal appendage; B, tables from body wall of small individual, ill

table at fight forming phosphatic bodies; C, rosette of raquet-shaped plates with anchor (not to scale,

copied from H.L. Clark, 1907); D. racquet-shaped plate. I

212 I
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I Genus Hedingia Deichmann, 1938

Type Species. Trochostoma albicans Thb.el, 1886, designated by Deichmann (1938b).

I Diagnosis. Fifteen tentacles, with 2 pairs of digits, without terminal digit. Pseudomesenteries absent.

Ossicles huge tables or ptates with large perforations and smooth or spinous surface.

I Hedingia cah'fornica (Ludwig, 1894)

i Figure 9.8

Caudina californica Ludwig, 1894.--H.L. Clark, 1907:37; 1935:279.

i Hedingia californica Deichmann, 1938b:112.

Material Examined. Gulf of California, syntype (USNM 18164).

I Diagn .osis. Ossicles large spiny plates with many perforations; spines sometimes mounded.

Remarks. Very little is known about this species. It was described from deep water in the Gulf of

California. Clark (1907) ascribes two other specimens to this species: one from 85 m off Baja California and

I the other from "Southern California probably". Deichmann (1938) considers all members of the genus

Hedingia to be deep-water forms. It may be that the shallow-water specimens identified by Clark (1907)

were young Caudina arenicola.

I Type Locality and Type Specimens. Gulf of California, syntype, Albatross Station 3434 (USNM

18164)

I Distribution. Uncertain, see Remarks.

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I Figure 9.8. Hedingia californica: ossictesfrom caudal appendage of syntype.

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Order Aspidochirotida Grube, 1840 J

Diagnosis. Tentacles peltate, 15 to 30 in number. Respiratory trees present. Mesentery of posterior I

loop of intestine attached to right ventral interradius. Gonad in 1 or 2 tufts. Ossicles usually include tables.

Family Stichopodidae Haeckel, 1896 I

Diagnosis. Tentacle ampullae present. Gonads in 2 tufts, one on either side of dorsal mesentery. I

Dorsal side of body usually with papillae or warts. Ossicles as tables and sometimes C-shaped rods and

buttons. I

Genus Parastichopus H.L. Clark, 1922

Type Species. Stichopus tremulus Gunnerus, 1770, designated by Deichmann (1937). I

Diagnosis. Body cylindrical, mouth ventrally placed, anus terminal. Dorsal side with large conical

papillae and smaller feet; ventral side with cylindrical tube feet more or less arranged in bands. Ossicles _

tables and buttons with several holes along sides, no terminal hole. iN

Remarks. This genus was proposed but not described by Clark (1922). Deichmann (1937) defined

the genus. I

Parastichopus cah'fornicus (Stimpson, 1857) I

Holothuria californicus Stimpson, 1857:524. •

I

Stichopus californicus H.L. Clark, 1901b:164, 1922:70, pl. 1,figs. 8-12.

Stichopus fuscus Therl, 1886:5. I

Parastichopus californicus Deichmann, 1937:163.--Lambert, 1986:2267, fig. 3B. U

Material Examined. Califonia: USNM Albatross 4534, 4 mi S80°E Pt. Pifios Lighthouse, 76-86 lm

fms, hard grey sand (1:15 cm); USNMAlbatross 2906, 34°23'30"N, 129°19'30 '', 96 fms (1:13 cm); USNM I

2964, 34°22'45 '', 119°40'00"W, 21.5 fms, sand, stones (1); Bodega Bay, intertidal (2:11.6, 16.4 cm) (AHF

U

295.36); USNM BRC-001; SBMNH BRA-1.

Description. Body cylindrical, with numerous large dorsal papillae. In life, mottled brown to reddish I

brown, rarely white; papillae flesh colored. Ossicles tables, 0.06-0.12 mm in diameter, and oblong plates,

0.13-0.24 mm long. am

Biology. Parastichopus californicus is a common member of the epifauna in both hard- and soft- II

bottom habitats. It feeds by sweeping its tentacles across the substmte, picking up surface detritus. In Puget

Sound, P. californicus spawns in the spring (McEuen, 1988). Oogenesis in this species has been described •

by Smiley (1988). II

Type Locality and Type Specimens. California, Tomales Bay, types lost.

Distribution. Alaska to Cedros Island, Baja California, intertidal to 216 m. •

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I Parastichopusjohnsoni (The_l, 1886)

Figure 9.9

i Stichopus johnsoni The61, 1886:4.--H.L. Clark, 1922:69, pl. 1, fig s. 15,16.

i Parastichopus johnsoni Lambert, 1986:2271.

Material Examined. California: 30 mi S of SantaBarbara, 22 fms (1) (holotype, MCZ 767); USNM

BRA-17.

i Description. Body cylindrical, form and color uncertain, possibly yellowish in life. Ossicles tables

0.10-0.17 mm in diameter, oblong plates, 0.14-0.22 mm long.

I califomicus Re.marHowever, ks. Deichmaim after measuring (1937) cthe onsidered ossiclesParastichopus in a number ofjohnson specimens, i to beLambert an immature (1986) determined form of P.

that tables in P.johnsoni were significantly larger than the tables in P. californicus. He considered these two

I species tType o be distinct. Locality and Type Specimens. California, 30 mi S of Santa Barbara, holotype (MCZ 767).

Distribution. California, Point Conception south to Baja Califomia, 40-140 m. I

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I Figure 9.9. Parastichopusjohnsoni: ossiclesfrom the body wall of the holotype.

I

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Parastichopus parvimensis (H.L. Clark, 1913) I

Stichopusparvimensis H.L. Clark, 1913:234, 1922:70, pl 1, fig. 13-14. I

Parastichopus parviraensis Deichmann, 1937:163.--Lambert, 1986:2267, fig. 3C.

Matedai Examined. Ca_fonfia: Four& of July Cove, Cata_na I, 1-2 firm (1:11 cm) (AHF 339.29); I

Santa Momca Bay, 60 m (1) (SCCWRP SMB 6.4 No. 1).lMe_co: Ce&os I, Baja Ca_fon_a _ototype,

USNM Ace. No. 55337). Ii

Description. Body cylindrical with numerous small dorsal papillae and 3-6 large papillae. In life, II

body light brown, small papillae with black tips. Ossicles tables, 0.04-0.06 mm in diameter, and oblong

plates, 0.07-0.14 mm long. •

Biology. The habits and habitat of Parastichopus parvimensis and P. californicus are similar;

however, P. parvimensis has a more southerly distribution.

Type Locality and Type Specimens. Mexico, Baja California, Cedros I, 1 m, holotype (USNM i

Ace. No. 55337). W

Distribution. Central California to Baja California, intertidal to 64 m. I

Order Dendrochirotida (Grube, 1840) I

(Restricted Pawson and Fell, 1965)

m

Diagnosis. Tentacles highly branched, 10-30 in number. Calcareous ring simple or complex; retractor I

muscles present.

I

Family Psolidae Perrier, 1902

I

Diagnosis. Dorsal body wall more or less completely covered by scales. Ventral body wall flat, not

covered by scales, functioning as a "sole". Mouth and anus dorsal, usually protected by valves.

!

Genus Psolus Oken, 1815

Type Species. Holothuria phantapus Strussenfeldt, 1765, designated by Oken (1815). I

Diagnosis. Body oval with distinct sole, with marginal tube feet; tube feet sometimes present in the

midventral ambulacrum. Dorsal side covered with scales; oral and anal apertures with or without operculum III

of triangular scales. Dorsal side without appendages except for anal papillae and, in some cases, a few tube |

feet on introvert. Ten tentacles present. Ossicles in sole plates or buttons, smooth or knobbed; ossicles

sometimes develop reticulations or may be reduced with age. I

I

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216 I
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I Psolus chianoides H.L. Clark, 1901

Figure 9.10

!

PsoIus chitinoides H.L. Clark, 1901a:335, pl. 3, figs. 4-5, pl. 4, figs. 6-10; 1924:57.

l Material Examined. Washington: 74 m, rocky (1:4.9 cm) (USNM E1992).--California: Pt Arguello, 1 (Nekton).

Description. Medium-sized form, up to 7.0 centimeters long. Oral and anal opening surrounded by

i triangular plates. Dorsal plates smooth, without granulations. Approximately 4 rows of plates between oral

and anal scales. Ossicles in sole knobbed buttons, approximately oval in shape, with many knobs and

perforations; reticulated mass often present.

I Biology. Psolus chitinoides is found on top of rocks and in cobble fields. They are generally sessile,

but may move at times. In Puget Sound, P. chitinoides spawns in March and May (McEuen, 1988; McEuen

and Chia, 1991). The females extrude ropes of brick red eggs, which float to the surface. After 10 to 15

I minutes, the gelatinous material binding the eggs dissolves, releasing the eggs.

Type Locality and Type Specimens. Type locality unknown, holotype (MCZ 309)

I Distribution. British Columbia to Pt Arguello, California; intertidal to at least 74 m, rocky habitats.

I

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I

Figure 9.10. Psolus chitinoides: buttons from sole.

I

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Psolus squamatus (Koren, 1844) I

Figure 9.I 1

m

Cuvieria squamata Korea, 1844:211, pl. 2-3. i_

Psolus squamatus H.L. Clark, 1901b:165, 1901c: 491.--Mitsukuri, 1912:225, pl. 7, figs. 61-62, text-fig. I

42.--Oshima, 1915:280. |

Psolus squamatus var. segregatus Pettier, 1905:59.--Ekman, 1925:136, fig. 33.--Deichmarm, 1941:147,

pl. 30, fig. 7. I!

Psolus pauper Ludwig, 1894:139. l

Material Examined. Califonia: Gull Islet, S coast of Santa Cruz I, 97-281 fms, sand, rock, pebbles

(3:6.1-7.6 cm) (USNMAlbatross 4430); 7.6 mi S of East Pt, San Nicolas I, 216-339 fms, sand, pebbles, rock II

(1:9.2 cm) (USNMAlbatross 4423); 6.9 mi N8°E SW rock, Santa Barbara I, 238-310 fms, mud, sand, rock

(2:6.0-6.8 cm) (USNMAlbatross 4418); off southern California, 33°49'45"N, 119°24'30"W, 603 fms, green •

mud (1) (USNM Albatross 2980); 276 fms (1:5.3 cm) (USNM E2000); northern California, 239 fms (2: |

3.6, 4.6 cm) (USNM E2001); USNM BRA-02; SBNMH BRA-04.

Description. Large form, up to 13 cm long. Oral and anal opening surrounded by triangular scales. 1

Dorsal scales granulated; approximately 12 scales between oral and anal plates. Ossicles of sole plates with

2-4 holes and marginal projections; ossicles often reduced with age.

I

Type Locality and Type Specimens. Patagonia, types missing.

Distribution. Southern Bering Sea to Cape Horn; in northern hemisphere, 194 to 1206 m.

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Figure 9.11. Psolus squamatus: platesfrom sole. I

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I Genus Lissothuria Verrill, 1867

Type Species. Lissothuria ornata Verrill, 1867, by original designation.

i Diagnosis. Small to medium-sized form with distinct sole. Dorsal scales present, covered by a layer

of ossicles, including hour-glass shaped bodies or towers or both. Ossicles in sole as knobbed or smooth

I plates; cups sometimes present.

Lissothuria nutriens (H.L. Clark, 1901)

I Figure 9.12

I Thyonespolu pl. 26, s nufigs. triens1-5.H.L. Clark, 1901a:169, 1901b:491.--Deichmann, 1930:193, 1937:174, 1941:138,

Psolidium nutriens Ludwig, 1904:689.

I Lissothuria nutriens 1967:6.

Pawson,

I Material Examined. California: Talcott Shoals, Santa Rosa I (2) (MB); SE of Carrington Pt, Santa

Rosa I (1) (MB); E of pier, Johnson's Lee, Santa Rosa I (1:1.2 cm) (MB); Pacific Grove (1: 1.8 cm) (USNM

l

22458).

Description. Small, up to 2 cm long. Dorsal scale coveting incomplete, with scales absent from

I towers hourglass-shaped deposits; cups absent. Ossicles sole perforated

dorsal midline. Dorsal ossicles and in

plates with raised marginal projections and cups. Ossicles in tentacles perforated plates or buttons and rods;

rosettes lacking. !

I Figure 9.12. Lissothuria nut-dens: ossicles from sole,A, plates; B, cups.

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Biology. Lissothuria nutriens is red in life with a pinkish sole. The female produces a small number •

of large eggs that are brooded in pits on the dorsal surface. When ready to leave the mother, the young have

ten tentacles and six tube feet. By the time the animal has grown to 2 mm in length, the sole is fully •

developed. l

Type Locality and Type Specimens. California, Monterey Bay, Pacific Grove, intertidal (MCZ 244).

Distribution. In rocky habitats, Monterey Bay south to Catalina Island, intertidal to 20 m. I

Lissothuria deichmannae Pawson, 1967 1

Lissothuria deichmannae Pawson, 1967:14, fig. 5. U

Description. Dorsal scale covering complete. Dorsal ossicles hourglass-shaped deposits and fiat I

plates; towers absent. Ossicles in sole large plates with conspicuous knobs which become multilayered with

growth. Small hourglass-shaped plates also present.

Remarks. Pawson (1967) described Lissothuria deichmannae from one specimen collected in the n

intertidal zone on Santa Cruz Island. It is distinguished from L. nutriens by the large plates in the sole that H

tend to become multilayered. At present, little is known about its distribution or habits.

Type Locality and Type Specimens. California, southwest shore of Smugglers Cove, Santa Cruz i

Island, intertidal, holotype (AHF 325.1).

Distribution. At present known only from the type locality, nn

Family Cucumariidae Ludwig, 1894 Hen

emend. Pawson and Fell, 1965 •

U

Diagnosis. Body wall not enclosed by test of imbricating plates. Calcareous ring simple, lacking •

posterior prolongations.

ll

Subfamily Cucumariinae Panning, 1949

Diagnosis. Ten tentacles. Ossicles in body wall as knobbed or smooth plates; no cups or plates with i

spil'es. 1

Remarks. In Panning' s (1949) classification, the genera of Cucumariinae are differentiated by the

form of the plates in the body wall. Accordingly, species in the genus Cucumaria have thin, spiny plates, i

Species inPseudocnus have thick knobby plates. These plates are either round or are oblong with a tendency

to develop a dentate handle. I have attempted to allocate California species to the appropriate genus; however,

in some cases, the allocation is uncertain. In particular, C. piperata is problematic. While most ossicles in i

this species are flat and spiny, some are knobby with a dentate handle. U

Genus Cucumaria Blainville, 1834 1

emend. Panning, 1949

Type Species. Holothuriafrondosa (Gunner, 1767) designated by Panning (1949). •

I

Diagnosis. Ossicles thin plates, smooth or with spines.

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I Cucumaria curata Cowles, 1907

Figure 9.13

i Cucumaria curata Cowles, 1907:2.--Panning, 1949:416.--Rutherford, 1977: 165, figs. 3-5.--Lambert,

1984:437, fig. 1.

I Material Examined. California: Carmel Bay, intertidal (1: 2.4 cm) (AHF Acc. 1234); San Clemente,

intertidal (multiple) (BLM 2A28376).

I Description. Small form, up to 2.5 cm long. Ten tentacles, approximately equal in size. Dorsal

body wall and tentacles purplish brown to black, with tube feet scattered in interambulacra. Ventral body

wall yellowish. Single stone canal. Ossicles of the body wall as small buttons with 4-6 holes and larger

I plates with 20 or more holes.

Remarks. Rutherford (1977) considered Cucumaria pseudocurata to be a synonym of C. curata.

However, Lambert (1985) concluded that both C. curata and C.pseudocurata were valid species. He believed

I that C. curata was very limited in distribution, only occurring at Pacific Grove in Monterey Bay. He,

therefore, believed that Rutherford had only collected and studied C. pseudocurata.

I

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I Figure 9.13. Cucumariacurata: ossicles from body wall.

I 221
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The specimens listed under Material examined above agree well with Cowles' and Lambert's •

description of C. curata. Since they seem distinctly different, I have herein treated C. curata and C.

pseudocurata as separate species. Additional studies are needed to determine the status of these two species. •

Cowles named this species "curata" for its habit of brooding its eggs under the ventral surface of II

the body wall.

Type Locality and Type Specimens. Cypress Point, Monterey Bay, intertidal, types missing. I

m

Distribution. Uncertain, presumably near Pacific Grove, Monterey Bay, intertidal in rocky habitats

(see Remarks). I

Cucumaria miniata (Brandt, 1835)

Figure 9.14 I

Cladodactyla (Polyclados) miniam Brandt, 1835:44. I

Cucumaria miniata Selenka, 1867:350.--H.L. Clark, 1924, p. 56.--Edwards, 1910:604, pl. 19, figs. 17-21. II

Cucumaria albida Selenka, 1867:350, pl. 20, fig. 109.

i

Cucumariajaponica Lampert, 1885:143.--H.L. Clark, 1902:562-564.--Edwards, 1907:61-62. I

Material Examined. California: Midway between Twin and Platts Harbor, Santa Cruz I, 10-50 ft.,

rock (1:7.3 cm); Santa Barbara I, 321 fms (1:3.9 cm) (VeIero 1856-49); off UCSB marine lab, Santa II

Barbara, 30 ft (1:5 cm) (SBNMH 49001); 1.5 mi E of Cardwell Pt, San Miguel I, 20-21 fms (3:4.0-10.0 cm)

(Velero 1415-41); Anacapa Passage, 33°59'00"N, 119°32'08"W, 15-50 fms (2: 7.4, 11.1 cm) (Velero 1190-

40); 1 mi S of East Pt, Santa Rosa I, 15-16 fins (2: 7.0, 9.4 cm) (Velero 1284-41b); 5 mi 152° from San Pedro II

breakwater, 18-19 fins (1) (Velero 1232-41b); S side of North jetty, Bodega Bay, intertidal (8:1.0-9.0 cm) E

(AI-IF342.15); 1.5 mi E of Cardwell Pt, San Miguel I, 20-21 fms (3:7.4-12.0 cm) (Velero 1415-41); E of

pier, Johnson's Lee, Santa Rosa I, (1:6.7 cm) (MB); Campbell Cove, Bodega Harbor, (2:7.3, 8.8 cm) (AHF I

342.6); Cambria, San Luis Obispo Co, (1:2.3 cm) (Velero 1581-47); North jetty, Bodega Bay, intertidal

(4:6.9-11 cm) (AHF 342.1); Point Arena, Mendocino Co (1) (Velero E.B. 41-49); 1.5 mi East of Cardwell

Pt, San Miguel I, 20-21 fms (2:3.4, 8.5 cm) (Velero 1415-41).--Mexico: 5.5 mi S of San Benito I, 66-81 fms •

(1: 1.9 cm) (AHF 403.9).----Oregon: North Bay, Cape Arago State Park, intertidal (1:12.5 cm) (Velero 1489- E

42).

Description. Large form, upto 20 cm long. Body cylindrical, with 10 equal tentacles. Color in life II

red, brownish in preserved specimens. Tube feet restricted to ambulacra. Twenty or more stone canals II

present. Ossicles in body wall small oblong plates, with 3-7 holes and spines scattered on the surface. Plates

often narrowed at one end. I

Biology. Cucumaria miniata is found under rocks and in crevices in hard-bottom habitats. In Puget

Sound, it spawns in spring (McEuen, 1988).

Remarks. This species is differentiated by it large size, red color, lack of interambulacral feet, II

multiple stone canals and small fiat spiny ossicles.

Type Locality and Type Specimens. Sitka, Alaska, types missing. •

Distribution. Alaska south to San Benito Island, Baja California, intertidal to 35 m, rocky habitats. II

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Figure 9.14. C_:_na miniata: ossicles from body wall.

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Cucum pipe_m (Stimpson, ]864)

I Figure 9.15

Pentacta piperata Stimpson, 1864:161 .--H.L. Clark, 1901b: 171.

I Cucumaria piperata H.L. Clark, 1924:56.--Deichmann, 1937:169.--Panning, 1962:77, fig. 20.-Lambert,

1990:913, fig. 1,4.

i Cucumariafisheri Wells, 1924:114, text-fig. 1, pl. 3, fig. I-2.

Material Examined. Washington. Friday Harbor, Puget Sound, (1) (USNM Acc. 102730).--

California: 18.5 mi S of San Nicolas I, 57 fms (1:4.3 cm) (AHF 403.13); I mi N of west end of Anacapa I,

I 45-46 fms (3:0.9-1.7 cm) (AHF 403.15); 4.25 mi NE of Sandy Pt, Santa Rosa I, 35-36 fms (1:3.2 cm) (AHF

403.12); 4 mi SE of east end of Santa Catalina I, 104-135 fms (2:3.6, 4.6 cm) (AHF 403.16); 0.5 mi S of

west end of Anacapa I, 25-26 fms (4:0.8-1.5) (AHF 403.7); 0.35 mi SW of Ship Rock, Santa Catalina I, 34-

I 43 fms (1:1.8 cm) (AHF 515.1); off Del Monte, Monterey Bay, 5 fms (2:1.5, 2.3 cm) (USNM Acc. 152709); 3.25 mi NE of San Pedro Pt, Santa Cruz I, 46-47 fms (I :1.7 cm) (Velero 1418-41); 10.75 W ofPt Dume, 47-

48 fm (1:3.6 cm) (Velero 1276-41a); 1.5 mi NW of Cavern Pt, Santa Cruz I, 54-56 fins (1:2.5 cm) (Velero

I 1300-41b); E of pier, Johnson's Lee, Santa Rosa I, (1:2.4 cm) (MB); SE of Carrington Pt, Santa Rosa I, (2:2.8, 4.4 cm) (MB); East Pt, Santa Rosa I, (1:2.0 cm) (MB); Tajiguas, Santa Barbara Co, 20 ft (3) (SBMNH

49004m); W of Dutch Harbor, San Nicolas I, 8-9 fms (1:1.7 cm) (VeIero 1204-40); Pt Dume, 150 m (1:1.8

I

I 223
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cm) (SCCWRP R15-150).---Mexico: Ranger Bank, off Cedros I, Baja California, 81-83 fms (4:0.5-1.5 cm) I

(AHF 403.2); 5 mi S of San Benito I, Baja California, 87-95 fms (50:1.5-4.7 cm) (Velero 1119-40); off San

Benito66_81 fmsI' Baja(l:2.3Calif(AHF92-95403.9);fms ornia,cm) (5s: 0.9-7.1 cm) (Velero 1010-39); 5.5 mi S of San Benito I, Mexico, I

Description. Medium-sized form, up to 7 cm long. Body cylindrical with 10 equal tentacles. Color

inOssicles life yellowish with brown spots; tentacles brown. Tube feet restricted to ambulacra. Single stone canal. I

round to oblong plates with a spiny margin and spiny or slightly knobbed surface. Some plates with

dentate handle.

Biology. Cucumaria piperata is found in both hard- and soft-bottom habitats, sometimes in great I

numbers. McEuen (1988) reports seeing C. piperata in "vast beds on soft bottoms" in Puget Sound. On

Santa Rosa Island C. piperata was found clinging to coralline algae in a mixed aggregation with Pseudocnus

lubricus and Pachythone rubra (M. Bergen, unpublished). I

Remarks. Lambert (1990) synonymized Cucumaria fisheri fished Wells, 1924 with Cucumaria

piperata. A discussion of the status of Cucumaria fisheri astigmata Wells, 1924 is included under the

remarks for Pseudocnus lubricus. I

Type Locality and Type Specimens. Cucumariapiperata: Washington, Puget Sound, type missing;

C.fisherifisheri Distribu: tionCalifornia, . Washington, MtereyPugetBay,Sound 24 m,south holotypeo San(USNM Benito E1198). Island, Baja California, 6-250 m. I

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Figure 9.15. Cucumariapiperata: ossicles from body wall.

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I Cucumaria pseudocurata I)eiehmann, 1938

Figure 9.16

I Cucumariapseudocurata Deichmarm, 1938:104, fig. 1.--Rutherford, 1977: 165, figs. 3-5.--Lambert, 1985:

437, fig. 2.

I Cucumaria curata: Smith, 1962:233-246. Not Cowles, 1907.

Material Examined. California: San Mateo, (1) (cotype, MCZ 356). Velero EB23, Zone 3,

I Mendocino Co, Cormorant Cove, intertidal; Velero M1-48, Dillon Beach, intertidal. Oregon: Velero 1492- 42, Cape Arago State Park, intertidal (multiple); Agate Beach, intertidal (multiple); Agate Beach, under

Yaquina Head Light, intertidal (21:0.7-1.8 cm) (Velero 1477-42).

I Description. Small form, 2-3 cm long. Ten tentacles, approximately equal in size. Dorsal body wall

purplish brown to black; ventral body wall yellowish. Tube feet restricted to ambulacra. Ossicles in body

wall oblong to round plates, often with 20 or more holes, and rods.

I Biology. Cucumaria pseudocurata is found in dense aggregations in the lower intertidal zone in

rocky intertidal habitats. Spawning occurs in December and January (McEuen, 1988; Rutherford, 1973;

Smith, 1962). When spawning, the females deposit the eggs under the ventral surface of the body wall,

I where they are brooded. After about a month the eggs hatch and the young move away from the female.

Remarks. According to Lambert (1985), the small black holothuroid commonly found in rocky

I interti Grove,daMonterey l areas in nBoarthern y (SeeCali Remarks fornia under is C. pseudocurata C. curata). ;Lambert C. cura(I985) ta is restalsorictedsuggests to a smal thatl are C. pseudocurata a near Pacific

and C. vegae The61, 1886, a species described from the Aleutian Islands, may be synonyms.

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I Figure 9.16. Cucumariapseudocurata: ossicles from body wall.

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Type Locality and Type Specimen. Southern California, San Mateo, cotype (MCZ 2356). (Note: I

The type locality for this species is uncertain as San Mateo is in northern, not southern California).

Distribution. Queen Charlotte Islands, British Columbia south to Monterey Bay, California, l

intertidal, rocky habitats. II

Cueumaria salma Yingst, 1972 I

Figure 9.17

Cucumaria salma Yingst, 1972:142, figs. 1-3. I

Material Examined. California: 1 mi S of East Pt, Santa Rosa I, 15 fms, loose rock, sand (1:4.5 WI

cm) (Velero 1284-41); San Miguel Passage, 37-39 fms, shale, gray sand (1:0.9 cm) (Velero 990-39); 0.5 mi l[

E San Pedro Pt, Santa Cruz I, 26-40 fms, rock (1:1.9 cm) (Velero 1297-41); 1 mi N of San Pedro Pt, Santa

1

Cruz I, sand, broken shell (1:1.5 cm) (Velero 1298-41); off Anacapa I, 2-20 ft(1:2.5 cm) (SBMNH 49049m); 1

Becher's Bay, Santa Rosa I, (1:2.2 cm) (AHF Hubbs 47-86a); 0.5 mi N of Platt Pt, Santa Cruz I, 36-47 I

(2:0.9, 1.1 cm) (Velero 1303-41a); 0.5 mi N of Platt Pt, Santa Rosa I, 36-47 fms (3:0.5-1.1 cm) (Velero I1

1303-41b); Smugglers Cove, Santa Cruz I, 40 ft (1:4.1 cm) (SBMNH 49010); 5 mi 152° from San Pedro =.

breakwater, 18-19 fins (6:2.5-5.0 cm) (Velero 1232-41 a); Becher_ s Bay, Santa Rosa I, 13-16 fms (8) (Velero 1

1004-39b); 4.25 mi NE of Sandy Pt, Santa Rosa I, 46-47 fms (1:1.5 cm) (Velero 1417-4Ib); E of pier,

Johnson's Lee, Santa Rosa I (1:3.9 cm) (MB); off Pt La Jolla, 55 m (1:6.1 cm) (Scripps E2659); N face of

Bird Rock, Catalina I, 40 ft (1) (holotype, AHF 61). 1

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Figure 9.17. Cucumaria safma: ossicles frombodywall. I

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I Description. Medium-sized, up to 10 cm long. Body cylindrical with 10 equal tentacles. Color in

life salmon; anterior end black; tentacles and buccal membrane black mottled with white. Tube feet restricted

to ambulacra. Three to 4 stone canals present. Ossicles in body wall smooth plates with undulating margin

I and 2-20 or more perforations.

Remarks. Cucumaria salma lives in crevices in rock reefs. Black dendrochirote tentacles protruding

I frCucumaria om a rockcrax wallDeichmann are often ,the1941, onlya evidence species described that thisfromspeciesBajaisCalifornia, present. Theareossicles similar. oItf isCpossible . saIma and that

further study will show these two species to be synonyms.

Type Locality and Type Specimens. California, Catalina I, Bird Rock, 13 m (holotype and paratypes,

I AHF 61).

Distribution. California, Santa Rosa Island south to Point Loma, San Diego County.

!

Genus Pseudocnus Panning, 1949

I Type Species. Cucumaria dubiosa Semper, 1868, designated by Panning (1949).

Diagnosis. Ossicles in body wall thick, knobby, either round or oval with one end tending to form

I a dentate handle.

Pseudocnus lubricus (H.L. Clark, 1901)

I Figure 9.18

I Cucumaria lubrica H.L. Clark, 1901a:334, 1924:55.

Cucumariafisheri astigmata Wells, 1924:114, text-fig. 1, pl. 3, fig. 1-2.

i Pseudocnus astigmatus Lambert, 1990:913, fig. 2,3. Material Examined. Califonia: Paradise Cove, (1) (USNM Acc. No. 261891); 3.5 mi S of Fraser

Pt, Santa Cruz I, 25-28 fms, sand, broken shell (1:0.8 cm) (Velero 1287-41); 0.5 mi S of Gull I, Catalina I,

I 34-41 fms, sand, shell (7) (Velero 1294-41); 2.6 mi E of East Point, Santa Rosa I, 47-49 fms, mud

green

(2:0.9, 1.7 cm) (Velero 1289-41); 3 mi NW Anacapa I light, 47 fms, gray, green sand (multiple) (Velero

1267-41a); W of Dutch Harbor, San Nicolas I, 8 fms, rocky (7:1.4-2.4 cm) (Velero 1204-40); S. side of

I Santa Cruz I, 38 fms (12:1.2 - 3.7 cm) (Velero 1191-40); off Del Monte, Monterey Bay, 5 fms (10:0.4-3.5 cm) (USNM Acc. 152709); Monterey Bay, 16 fms (2:1.2, 2.3 cm) (USNM Acc. 152709); Chinese Harbor,

Santa Cruz I, 25 ft (1:2.0 cm) (SBMNH 49001m); Becher's Bay, Santa Rosa I, 13-16 fms (17) (Velero

I 1004-39a); 2.5 mi E of South Pt, Santa Rosa I, 17-18 fms (11) (Velero 1282-41b); 2.25 mi E of South Pt, Santa Rosa I, 23-28 fms (7:0.8-2.5 cm) (Velero 1283-41b); Monterey Bay, 8 fms (2:0.9, 2.1 cm) (Velero

1456-42); San Luis Obispo Bay, (1:1.8 cm) (Velero 885-38a); S of San Miguel I, 5-15 fms (2:0.9, 1.9 cm)

i (ofVelero buoy, 894-38); Cortez Bank, 1 mi50S offmsEa(sVelero t Pt, Santa 1342-4Rosa1); Nifty I, 15-16Rock, fmsSan(3:0Miguel .8-2.3 I,cm)(1:2.2 (Velero cm) (M1284-4 B); SE1b);of9.5Carrington mi NW

Pt, Santa Rosa I, (7:0.8-2.2 cm) (MB); East Pt, Santa Rosa I, (12:0.5-1.5 cm) (MB); E of pier, Johnson's

I Lee, S o Santa Rosa I, Ca (6:1.0-3.8 cm) (MB); W of pier, Johnson's Lee, Santa Rosa I, (1:2.3 cm) (MB); 4 mi f Gaviota, (1:2.0 cm) (SBMNH 48998); Pelican Cove, Santa Cruz I, 20-60 fi (2) (SBNMH); reef midway

between Twin and Platts Harbors, Santa Cruz I, 10-50 ft (1) (SBNMH 48845m); Tajiguas, Santa Barbara

I CBarbara o, 20 ftCo(3), (1) (SB(SBNMH NMH 48849); 49000);Chinese 1 mi NHarbor, of W endSanta of Anacapa Cruz I, (I,1)46(SBNMH fms (10:0.7-1.6 49004m); cm)Ho(Velero pe Ranch, 1419-41); Santa

4.25 mi N of Sandy Pt, Santa Rosa I, 35 fms (2:1.2, 1.7 cm) (Velero 1417-41); 2 mi W of Cardwell Pt, San

I Miguel (Hubbs I47-85); , 35 fms1(mi 1:3.1NWcmof)W(Velero end, Anacapa 1413-41);I, 413.6fins mi offS (1:1.7coast, cm) (VeIero near E end, 1937-50); Santa 3.5Rosmi a I,NE(2:0.5, of San0.9Pedro cm)

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PL Santa Cruz I, 46 fms (4:1.1-1.4 cm) (Velero 1418-41); W of Dutch Harbor, San Nicolas I, 8-9 fms I

(23:0.5-3.1 cm) (Velero 1204-40); 3 mi E of South Pt, Santa Rosa I, 18 fms (2:1.7, 2.9 cm) (Velero 1410-

41); 1.5 mi NW Cavern Pt, Santa Cruz I, 55 fms (42) (Velero 1300-41); 3.6 mi off East Pt, Santa Rosa I, 53 •

fms (2:2.0, 2.1 cm) (VeIero 1388-41); 2 mi S of Crook Pt, San Miguel I, 42 fms (3:1.3-1.9 cm) (Velero 1

1412-41); 9.5 mi SW of Tanner Bank, 72 fms (1:I.0 cm) (Velero 1346-41); 3 mi N of Arch Rock, Santa

Cruz I, 55-60 fins (3:0.3-0.7 cm) (Velero 1304-41); USNM BRC-002; SBMNH BRA-016.--Puget Sound, •

Washington (2:2.3, 3.2 cm) (syntypes, MCZ 202);--Oregon: Middle Bay, Cape Arago State Park, (2:2.3, 1

2.8 cm) (Velero 1487-42);

Description. Medium-sized, cylindrical, blunt on both ends, up to 4 cm long. Tube feet retractile, •

not restricted to ambulacra. Single stone canal and polian vessel present. Ossicles mostly round, mounded,

knobbed plates and buttons, smallest with 4 symmetrically placed holes. Some larger plates with one end

drawn out into handle. •

Biology. From the specimens examined, Pseudocnus lubricus apparently lives in both hard- and

soft-bottom habitats. In hard-bottom habitats, P. lubricus broods its eggs under the ventral surface of the

body. When spawning, the female folds the anterior end under the body to release the eggs. While the n

females were sedentary, the males were apparantly errant during the spawning season. In Puget Sound,

spawning occurred in winter (McEuen, 1988).

Remarks. After viewing the type specimens of Cucumaria fisheri Wells 1924, Lambert (1990) 1

recognized that the specimens described as C.fisherifisheri and C.fished astigmata were distinct species, II

belonging, respectively, to the genera Cucumaria and Pseudocnus. Cucumariafisherifisheri was synonymized Illl

with Cucumaria piperata and C.fisheri astigmata was referred to the genus Pseudonus. 1

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Figure 9.18. Pseudocnus lubricus: ossiclesfrom body wall. I
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I Lambert (1990) noted that Pseudocnus astigmatus was similar to Cucumaria curata and C. lubrica,

inasmuch as the shape of the ossicles was similar to some degree and also because these species brooded

i their eggs under the ventral surface of the body. Cucmaria curata and Pseudocnus astigmatus, however, are quite distinct in terms of size (3 versus 5 cm maximum length), color (black versus white or white with

brown spots) and habitat (intertidal versus subfidal). On the other hand, P. astigmatus and C. lubrica are

i quite similar. When I examined the type specimens of C. piperata astigmata and C. lubrica, I found them to be identical. For this reason, I consider P. lubricus a junior synonym of P. lubricus.

Type Locality and Type Specimens. Cucumaria lubrica: Washington, Puget Sound, syntypes

I (MCZ 202); DistribC.ufisheri astigmata: California, Monterey Bay, 12 fms, (USNM E1196). tion. British Columbia, Queen Charlotte Sound, Alert Bay south to Cortez Bank, southern

California, 1.5-72 fms.

I Family Sclerodactylidae Panning, 1949

i emend. Pawson and Fell, 1965

Diagnosis. Body wall not enclosed in a test of imbricating plates. Calcareous ring complex with

I single or paired processes; processes entire, not divided into pieces.

Genus Eupentacta Deichmann, 1938

I Type Species. Cucumaria quinquesemita Selenka, 1867, by original designation.

Diagnosis. Ten tentacles, 2 ventral tentacles smaller. Body wall cylindrical. Tube feet non-retractile,

I restricted to ambulacra. Ossictes knobbed buttons or plates and cups or baskets or reticulated bodies.

Eupentacta quinquesemita (Selenka, 1867)

I Figure 9.19

I Cucumaria Cucumaria quinquesemita Selenka, 1867:351, pI. 20, fig. 107a,b. chronhjelmi The61, 1886:105.--H.L. Clark, 1924:55.

Eupentacta quinquesemita Deichmann, 1938:109.

I Material Examined. California: off Ford Pt., Santa Rosa I, 92 ft (1) (USNM Acc. 205678); Talcott

Shoals, Santa Rosa I, (1:5.2 cm) (MB); SE of Carrington Pt, Santa Rosa I, (1:6.0 cm) (MB); W of pier,

I Johns Mendocino, on's Lee, Santa Rosa I, (1:2.8 cm) (MB); Puget Sound, Washington (1:4 cm) (USNM 33091); (1) (cotype, MCZ 217).

Description. Medium-sized, up to 10 cm long. Body wall cylindrical with broad interambulacra;

I numerous tube feet in ambulacra. Ossicles as knobbed plates and cups.

Biology. Eupentacta quinquesemita occurs on top of rocks and boulders in rocky habitats. In Puget

Sound E. quinquesemita spawns in spring (McEuen, 1988).

I Type Locality and Type Specimens. California, Mendocino, cotype (MCZ 217).

Distribution. Alaska south to Santa Rosa Island, California, intertidal to 30 m, rocky habitats.

I Genus Pachythyone Deichmann, 1941

I Type Species. Thyone rubra H.L. Clark, 1901, by orginal designation.

I 229
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Figure 9.19. Eupentacta quinquesemita: ossicles from body wall, A, plates; B, cups; C, supporting tables from I

tube feet.

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I Diagnosis. Small form, up to 3 cm long, with numerous interambulacral feet. Ossicles as oval

buttons or plates with external side reticulated and regularly-knobbed 4-holed buttons.

I Pachythyone rubm (H.L. Clark, 1901)

i Figure 9.20

Thyone rubra H.L. Clark, 1901a:494, 1901b:166, figs. 1-5. Deichmann, 1939:5, fig. 3.

I Pachythyone rubra Deichmann, 1941:115, pl. 23, figs. 10-12.

Material Examined. California: S 6°W 2.5 miles north of anchorage, East Point, San Nicolas I,

I I,291-298 mussel fms, bed No. gray5Bmu(1) d, rock (BLM); (23)Pelican (Albatross Cove, 442Santa 1); SaCruz nta Cruz I, 20-40 I, intertidal ft (1) (SB(23)(AHF MNH 48848m); 294.18);reef Sanmidway ta Rosa

between Twin and Platts Harbors, Santa Cruz I, (1) (SBMNH 48845m); Chinese Harbor, Santa Cruz I, 25 ft

I M(1)CZ(SB244). NMH 49004m); Los Angeles Harbor, (2:1.5, 1.9 cm) (USNM 261891); Monterey Bay, (1) (cotype,

Description. Small form, up to 2 cm in long. Dorsal body wall orange red; ventral body wall white.

I Tbuttons. ube feet scattered in interambulacra. Ossicles in body wall as oval reticulated plates and regular 4-holed

Biology. This species may be locally very abundant in rocky habitats. It broods it eggs internally,

I producing fully-formed juveniles.

Type Locality and Type Specimens. California, Monterey Bay, cotype (MCZ 244).

Distribution. California, Monterey Bay south to Los Angeles Harbor, intertidal to 300 fins (548

I m), rocky habitats.

I Family PhyUophoridae Ostergren, 1907

(emended Pawson and Fell, 1965)

I Subfamily Thyoninae Panning, 1949

I Small medium-sized forms, often Ten 2 ventral tentacles smaller. Diagnosis. to U-shaped. tentacles,

Tube feet non-retractile, restricted to the ambulacra. Calcareous ring with long processes divided into pieces.

i was Remarks. In Pannlng's classification, the presence or absence of tables and plates in the body wall

used to divide the genera of Thyoninae. Accordingly, the genus Stolus has only plates, Pentamera and

Thorsonia have tables and plates, andAllothyone, Havelockia, Thyone and Hemiathyone have only tables in

i the body wall. Thorsonia andAllothyone are distinguished by having 4-pillared tables while Pentamera and Havelockia have two-pillared tables. Thyone is distinguished by having rosettes and no tables in the introvert

while Havelockia has tables and no rosettes.

i

feet as aIngehis nericclassification, character. Ther Panni eforeng, (1s94pecie 9)sdidin not the use genustheThy presence one sensu or abPanning sence ofmainytemmbu or malyacral not have tube

interambulacral feet. In addition, many former members of the genus Thyone, e.g., Thyone benti, were

i moved into It isother not clear genera. at present that Panning's classification really works. Tables commonly degenerate

with growth in this group, making practical application of the classification difficult. It is often not clear if

I a ossicle is a plate or a degenerate table.

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Figure 9.20. Pachythyone ruhra: ossiclesfrom bodywall, A, reticulated plates;B, 4-ho]edbuttons; C, supporting

tables from tube feet. I

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I To some degree Panning's classification does seem to work for California species. Species in the

genus Pentamera generally have tables and plates in the body wall and only have plates in the introvert.

I Spe andcintrovert. ies in the However, genus Havelockia there is considerable seem to haveuncertainty either tablesabout or ossicles the placement derived offromsometables species. in theInbody addition, wall

the importance ofinterambulacral feet as a character is uncertain. Pawson and Miller (1981) use the presence

i of mayinterambulacral have feet in their definition of the genus Thyone. Species in Thyone sensu Pawson and Miller rosettes only, tables only, or rosettes and tables in the introvert.

It is clear that this group is in need of revision. For the purposes of this chapter, I have followed

I Pan difficult ning'sto(apply 1949) theclassificat genericiondefinitions and allocated to individual the species specimens, to the genera I havetonothe t included best of my the genera ability. inSince the key. it is

I Genus Havelockia Pearson, 1903

Type Species. Havelockia herdmani Pearson, 1903, by original designation.

I Diagnosis. Ossicles in body wall and introvert tables with 2 pillars. Tables may become reduced

with age.

I Haveloekia benti (Deiehmann, 1937)

i Figure 9.21

Thyone benti Deichmann, 1937:170, fig 2, 1-11.

I Havelockia benti Panning, 1949:466. Thyone benti var. zacae Deichmann, 1938a:376, fig. 10.

Havelockia benti var. zacae Panning, 1949:466.

I Material Examined. Havelockia benti: California: 1.8 mi 299° to South Pt, Santa Rosa I, 22 f-ms,

sand, shell,(2: 1.1, 4.2 cm) (BLM 23192); 1mi 329° toSouth Pt, Santa Rosa I, 30 fins, sand, shell (2) (BLM

I 23045); East 3.55 mi 359.5 ° to East Pt, Santa Rosa I, 36 fms, sand, gravel, shell (1) (BLM 23205); 4.8 mi 349° to Pt, Santa Rosa I, 48 fins, sand, rocks (1) (BLM 22952); 46.3 mi 71 ° to China Pt, San Clemente I, 46

fms, sand, gravel, shell (1:0.7 cm) (BLM 24382); 4.25 mi 313° to South Pt, Santa Rosa I, 44 fms, sand,

I grS avel, shell (2) (BLM 22976); 2.6 mi 360 ° to East Pt, Santa Rosa I, 28 fms, sand, shell (2) (BLM 23206); side of Santa Cruz t, 39-40 fins, sand (6:0.8-3.4 cm) (Velero 119t-40); off Redondo Beach, 11-20 fms,

gravel (1:1.0 cm (Velero 1139-40); 11.5 mi SE of Long Beach, 32-52 fins, mud, sand, shell (1:1.1 cm)

I (Frasier VeleroPt,1160-40); San Santa Cruz Channel, 32-34 fms, sand, shell (1:1.9 cm) (Velero 1005-39); 3.5 mi S of ta Cruz I, 25-28 fms, sand, shell (1:1.9 cm) (Velero 1287-41); off Gull I, S side of Santa Cruz

I, 39-43 fms, sand, mud (4:0.6-0.8 cm) (Velero 1194-40); offSan Nicolas I, 29-30 fms, sand, shell (1:I.5

I cm1413-41); ) (Velero0.51120-4 mi E of 0);San2 miPedro SW Pt, of Cardwell Santa CruzPt,I, S26-40 anta Mfms, iguelrockyI, 34-35 (1:2.3fms, cm)m(Velero ud (3:0.8-1.1 1297-41); cm)off(VelBluff ero

Cove, 50-75 fms (1:0.9 cm) (Burch 4085); 1 mi E of Empire Landing, Santa Catalina I, 22 fins, sand, shell

I (1:0.8 of Archcm)Rock, (Velero Santa137Cruz 4-41I,); 55-60 4.5 mifms, SE ofmud, Eastshell Pt, Santa (1:1.2Ros cm)a I,(Velero 52 fms,1304-41); mud (4) (6Velero mi E of13South 87-41)Pt, ; 3Santa mi N

Rosa I, 28-30 fms, rock (7:1.2-4.2 cm) (Velero 1292-41); 2.5 mi SE of Bennett Pt, San Mignel I, 57 fms

I (1of:0.7 San cPedro m) (Vele Pt,roSanta1396-41); Cruz I, 1045-46 .75 mi fms,Wsand, ofPt shell Dume,(4:0.8-1.0 47-48 fins, cm)ro(cVelero ky (1:21298-41); era) (VeIero 1.5mi 1276NW-41of); Cavern 1 mi N

Pt, Santa Cruz I, 55 fms, mud, sand, shell (6) (Velero 1300-41); 0.5 mi S of Gull I, Santa Cruz I, 34-41 fms,

i sand, shell (21:0.4-1.7 cm) (Velero 1294-41; 1 mi E of Smugglers Cove, Santa Cruz I, 15-21 fms, sand,

I 233
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pebbles (2:0.5, 1.4 cm) (Velero 1295-41); 1.5 mi SW of Gull I, Santa Cruz I, 48 fins, mud (1:1.4 cm) (Velero i

1435-41); 0.35 mi SW of Ship Rock, Santa Catalina I, 34-43 fms, sand, rock (1:1.3 cm) (Velero 2955-54);

1.5 rni SW of Judith Rock, San Miguel I, 45-48 fms, sand, shell (4) (Velero 1411-41); 3.5 mi NE of San •

Pedro PL Santa Cruz I, 46 fins, sand (1:1.5 cm) (Velero 1418-41); Monterey Bay, 25 fms (1) (Burch 3711); l

0.5 mi N of Platt Pt, Santa Cruz I, 36-47 fms, shell (6:0.3-2.2 cm) (Velero 1303-41); off Abalone PL Laguna

Beach, 54-57 fms, mud (9:0.9-4.2 cm) (Velero 1131-40); USNM BSS-058. •

Havelockia bent• var. zacae: California: 20.3 mi S of East Pt, Santa Rosa I, 71 fms, sand, gravel, l

rock (1) (BLM 23071); 7.2 mi SSW of East Pt, Santa Rosa I, 33 fms, sand, shell (1) (BLM 22960); 6.2 mi

SSW of East Pt, Santa Rosa I, 37 fms, sand, shell, rock (1) (BLM 22958); off Abalone Pt, Laguna Beach, •

54-57 fms, mud (1:4.4 cm) (VeIero 1131-40); S side of Santa Cruz I, 37-40 fms, sand (1:4.1 cm) (Velero

1191-40); off Pt Pifios, Monterey Bay, 36 fins, shell (1:3.7 cm) (Velero 889-38); W side of Santa Barbara I,

,100 fms, sand (2:2.4, 2.7 cm) (Velero 1176-40); 2.5 mi NW of Anacapa Light, 48-51 fins, sand (2:3.1, 4.0 •

cm) (Velero 1268-41); off Bowen Pt, Santa Cruz I, 58-90 fms, sand, shell (5:3.1-4.5 cm) (Velero 1192-40); l

4.5 mi NNW of buoy, Cortez Bank, 60-61 fms, sand, rock (1:6.2 cm) (Velero 1341-41); 3 mi NW of

Anacapa Light, 47-52 fms, sand (2:4.0, 5.2 cm) (VeIero 1267-41); 1.5 mi off Dutch Harbor, San Nicolas I, •

63-83 fms, sand, mud, shell (2:4.1, 5.6 cm) (Velero 1220-40); 0.5 mi N of Platt Pt, Santa Cruz I, 36-47 fms, i

shell (2:0.8, 0.9 cm) (Velero 1303-41); 3 mi SE of Ship Rock, Santa Catalina I, 52-65 fins, sand, shell

(4:1.4-2.9 cm) (Velero 2953-54); 4.5 rni SW of East Pt, Santa Rosa I, 43-45 fins, mud (2:2.5, 2.7 cm) •

(Velero 1390-41); 0.35 rni SW of Ship Rock, Santa Catalina I, 34-43 fms, sand, rock (1:1.1 cm) (Velero I

2955-54); 1.5 mi SW of Judith Rock, San Miguel I, 45-48 fms, sand, shell (1:2.9 cm) (Velero 1411-4I); 1.5

mi S of Crook Pt, San Miguel I, 41-43 fms, sand, shell (1:4.8 cm) (Velero 1412-41); 1.5 mi SW of Gull I, •

Santa Cruz I, 48 fms, mud (2:2.4, 2.8 cm) (Velero 1435-41); 6 mi E of South Pt, Santa Rosa I, 28-30 fms, l

rock (1:2.2 cm) (VeIero 1292-41); 2.6 mi E of East Pt, Santa Rosa I, 47-49 fms, mud (1:1.7 cm) (Velero

1289-41); 0.5 mi S of Gull I, Santa Cruz I, 34-41 fms, sand, shell (1:2.5 cm) (Velero 1294-41); off Abalone •

Pt, Lagtma Beach, 25-27 fins, mud (1:1.9 cm) (Velero 1130-40); Pt Arguello, 450 ft (1:3.1 cm) (MEC Sta |

B2); Pt Argue•l•, (1) (Chambers sta 5); Pt ArgueUo, (4) (Chambers sta 1);Pt Arguello, (1:6.3 cm) (Chambers

sta 4); 4.5 mi SE of East Pt, Santa Rosa I, 52 fins, sandy mud (2:0.7, 1.0 cm) (Velero 1387-41); 6.5 ESE of i

South Pt, Santa Rosa I, 57 fms (1:1.6 cm) (Velero 1392-41); 6.1 mi 344 ° to East Pt, Santa Rosa I, 56 fms, |

coarse sand, shell (10).--Mexico: 5 mi S of San Ben•to I, Baja California, 87-95 fins, sand (2:3.8, 3.9 cm)

(Velero 1119-40); off San Ben•to I, Baja California, 92-95 fms, f'me sand (8:1.5-5.2 cm) (Velero 1010-39).

Other Material. California: 0.5 mi S of Gull I, Santa Cruz I, 34-41 fms, sand, shell (4:1.9-2.5 cm) i

(Velero 1294-41); 0.5 mi N of Platt PL Santa Cruz I, 36-47 fins, shell (2:3.1, 3.3 cm) (VeIero 1303-41); E of

Bridge Rock, La Jolla, 85 ft, under rocks (1:8.1 cm) (SIO E2657). •

Description. Small to medium-sized, up to 8 cm in long, somewhat U-shaped, slightly fusiform, l

anterior end blunt, posterior end tapering to a tail. Body wall cream, tan, orange, rust brown, grey or black.

Tube feet orange, scattered in interambulacra.

In Havelockia bent•: body wall ossicles tables with round to irregularly-shaped disk, 0.07-0.13 mm

in diameter, with 4-18 holes and a short spire (0.02-0.06 mm) ending in a few teeth. Supporting tables with

2-pillared spire. Iintrovert with tables and delicate plates sometimes looking like rosettes. Tentacles delicate i

rods, plates and occasional tables.

In Havelockia bent• var. zacae: body wall ossicles flat irregularly-shaped plates, 0.01-0.27 mm in i

diameter, and smaller, more delicate, rectangular plates, 0.10-0.15 mm in length, often with 2 enlarged i

central holes. Supporting tables with low 2-pillared spire or reduced to simple curved rod. Introvert with i

delicate plates, sometimes resembling rosettes; tables present, but rare. Tentacles with delicate rods and

plates; tables rare. i

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234 i
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,

, #

II F --200._--

I Figure 9.21. Havelockia benti: A, tables from body wall; B, supporting tables from tube feet; C, tables from introvert. Havelockia benti var. zacae: D, tables from body wall; E, supporting tables from tube

feet. Havelockia variant: F, biscuit-shaped plates and bar-shaped deposit.

i

i

i 235
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il

In "other" (see Remarks): body wall ossicles small biscuit-shaped plates, 0.04-0.09 mm indiameter, •

with 2-5 central holes, and bar-shaped ossicles 0.07-0.12 mm long. Occasionally irregular and rectangular

plates as in Havelockia bent• vat. zacae are present. Supporting tables, when present, curved bars as in H. •

bent• var. zacae. Introvert with delicate plates that sometimes resemble rosettes. Tentacles with delicate i

rods, plates and tables.

Biology. This is one of the most common holothurians in soft-bottom habitats on the continental

shelf in California.

Remarks. Most specimens of this species fall into one of three distinct groups: Havelockia bent•, I

H. bent• var. zacae and those I have allocated to "other". In specimens listed as H. bent•, tables are abundant i

in the body wall. In 1-1.bent• var. zacae ossicles are relatively scarce and most of the ossicles are plates. In I

specimens listed as "other", biscuit-shaped plates are common in the body wall.

I

There seems to be an association between the size of the specimen and type of ossicles, with H.

bent• tending to be smaller than H. bent• var. zacae, and H. bent• var. zacae tending to be smaller than

III

specimens with biscuit-shaped plates. Deichmann (1937, 1938) believed that body wall tables in H. bent• !

degenerated into the typical plates of H. bent• var. zacae. While intermediates between the three forms are iS

rare, there is no reason at present to consider them as separate species. All share a common anatomy and

il

form, including orange pigmentation in the tube feet that persists after treatment with bleach. At the present i

time, I believe that all the specimens belong to one extremely, variable species. The three groups are

differentiated with the hope that someone will study the different forms in greater detail

i

Type Locality and Type Specimens. Havelockia bent•: Washington, Friday Harbor, Rocky Bay, il

70 m (Museum of Comparative Zoology, MCZ 1810). l

Havelockia bent• var. zacae: Baja California, East of Cedros Island, 80 m (MCZ 1954).

Distribution. Puget Sound to northern Baja California, 36-200 m. I

Havelocida charlottae (Deichmann, 1938)

Figure 9.22 i

Pentamera charlottae Deichmann, 1938:108. •

Havelockia charlottae Panning 1949:466. I

Material Examined. California: Velero 886-38, S off Pillar Pt, Half Moon Bay, 16 fins, gravel i

(1:2.1 cm); Velero 1004-39, Becher's Bay, 13-16 fms, rock (1:2.5 cm); Velero 1856-49, Santa Barbara II

Basin, 321 fms (1:1.6 cm); MEC 206, Rep A, 49 ft (1); Chambers Station 5, Rep 2 (1); Pacific Biomarine,

Long Beach Harbor Breakwater (2:4.9-6.2 cm). it

Description. Medium-sized, cylindrical, up to 6.5 cm long. Tube feet crowded in ambulacra. Color il

of preserved specimens white or light coffee. Body wall ossicles tables with an oblong disk, 0.09-0.11 mm

long, and a spire 0.08-0.12 mm long with 3-5 cross beams and a few teeth on the tip. Supporting tables also •

with a long spire. Tables in introvert round to oblong, with 2 enlarged central holes and 8-16 additional I

holes, a dentate margin, and low spire. Tentacles with rods, plates and tables.

Type Locality and Type Specimens. California, Pacific Grove, tidepools (USNM E2391 and E2392). H

Distribution. California,Monterey Bay to southern California,hard bottom habitats,intertidalto 642 m.

Genus Pentamera Ayres, 1852 I

Type Species. Pentamera pulcherrima, 1852, by original designation.

,11

Diagnosis. Ossicles in body wall 2-pillared tables and plates; ossicles in introvert plates, i

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I Figure9.22. Havelockiacl_rlottae: A, tab]esfrombodywalt;B, tablesfrom introvert.

I

Pentamera lissoplaca (H.L. Clark, 1924)

I Figure 9.23

Cucumaria lissoplaca H.L. Clark, 1924:55.--Deiehmann, 1937:169.

I Pentame ra lissoplaca Deichmann, 1938:106 (passim).--Panning, 1949:465.

i Material Examined. Specimens with small tables: California: 2.25 mi E of South Pt, Santa Rosa I,

• 23-28 fms, sand, gravel (1:2.1 cm) (Velero 1283-41); Point Loma, (1:3.4 cm) (City of San Diego, E5); off

Ford Pt, Santa Rosa I, 92 ft (1:1.2 cm) (USNM Acc. No. 255678); (1:2 cm) (USNM E2293); off San Onofre

generating plant (1:1.3 cm) (MB).---Specimens with large tables: California: 10 mi SE of Long Beach, 23-

I 28 fins, mud, sand (1:1.3 cm) (Velero 1159-40); White Cove, Santa Catalina I, 36-41 fins, mud, sand (1:0.9

cm) (Velero 998-39); 1 mi E of Empire Landing, Santa Catalina I, 22 fms, shell, sand (4:0.7-0.8 cm) (Velero

1374-41); off Abalone Pt, Laguna Beach, 54-57 fms, mud (1:2.4 cm) (Velero 1131-40); off Huntington

i Beach, (1) (OCSD S-l-00); 0.75 mi off Mugu Pier, 50-60 fi (1) (SBMNH 48847m); King Harbor, 20 ft (2:6.3, 6.7 cm) (MB); Santa Monica Bay (1:4.7 cm) (Hyperion, IMIV).--Mexico: off Thurloe Head, Baja

California, 30 fms, sand (1) (VeIero 284-34); 4.25 mi of Redondo Pt. light, Magdalena Bay, Mexico, 9 fins,

I sand (1:I.1 era) (Velero 1717-49); off Thurloe Head, Baja California, 10-20 fms (1:1.2 cm) (Velero 285- 34).

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_'_ _1 i

'

A

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!

° !

200 _m

i

1.0 mm i

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Figure 9.23. Pentamera lissoplaca: A, lozenge-shaped plates and small tables, B, large tables, C, suppo_aag

tables; D, large plate from specimen from San Clemente. I
I

i Description. Medium-sized, up to 7 cm long, U-shaped, fusiform, tapering to a long tail. Body wall

ossicles as oblong plates with 4-6 holes and delicate tables 0.03-0.06 mm in diameter; tables sometimes

i ablunt bsent;teeth. large,Well rounddeveloped plates occasionally end plate present. present. Supporting tables with a 2-pillared spire ending in a few

One group of specimens, listed under "large tables" above, with oblong plates and supporting tables

i aschapter, described thesebutspecimens with tablecsonsidered relativelyPlarge . lissoplaca (0.05-0.16 (See mmRemarks). in diameter) and stout. For the purposes of this

Remarks. The specimens listed under "small tables" conform well to Clark's description of

_l Pentamera Iissoplaca. The specimens listed under "large tables" agree well with the description except that

the tables axe larger. While these tables are similar to those of P. pseudopopulifera, the white color of the

body wall, the presence of oblong plates and the shape of the spire on the supporting tables make it clear that

I these of P. lissoplaca specimens inareallnotrespects P. pseudoexcept populifforerathe. Since size ofthethespecimens tables, I amwithconsidering large tablesthemconformhereinto theas adesvariant criptionof

P. lissoplaca. Further study is needed to determine if these specimens are P. lissoplaca or an undescribed

i species. Type Locality and Type Specimens. British Columbia, Queen Charlotte Sound, Alert Bay (Victoria

Memorial Museum, No. 583), (Museum of Comparative Zoology, MCZ 1288).

I Distribution. Southeastern coast of Alaska to southern Magdalena Bay, Baja California, 18-82 m.

I Pentamera popuh'fera (Stimpson, 1857)

Figure 9.24

i Pentacta populifera Stimpson, 1857:161.

Cucumariapopulifera Theel, 1886:103; H.L. Clark, 1924:56; 1901b:171; Bush, 1921:76, text-figs. 40-41;

I CucumariaDeichmann, 1937:170. tenuicoriata Wells, 1924:121, pl. 3, fig. 4, text-fig. 4.

Pentamera populifera Panning, 1949:465.

I Allothyone tenuicoriata Panning, 1949:466.

Material Examined. California: 3.1 mi 106°to Pt Duroc, 27 fms, clay (2) (BLM 23981); 1/2 mi

I SW of Ribbon Rock, Santa Catalina I, 51-56 fins (2:1.9, 2 cm) (Velero 51-56); 11 112mi SE of Long Beach, 32-52 fms (1:1.2 cm) (Velero 1160-40); 2.6 mi E of East Pt, Santa Rosa I, 47-49 fms (6) (Velero 1289-41);

off Santa Barbara, 29 fins (1:1.5 cm) (USNM 30638); (1:3.5 cm) (BLM 22966); NE of Anacapa I, 45 fms

i (4:1.5-2.7 cm) (Velero 876-38); 3 mi N of Arch Rock, Santa Cruz I, 55-60 fms, mud (2:3.5, 3.7 cm) (Velero 1304-4I); off Abalone Pt, Laguna Beach, 28-29 fms, mud (2:1.5, 1.6 cm) (Velero 1130-40); off Bluff Cove,

50 fms (3:2.2-3.9 cm) (Burch 40125); 3 mi ESE off Ship Rock, Santa Catalina, 52-65 fms (6:1.1-2.3 cm)

I (Velero 2953-54); 4.5 mi SW X 0.5 mi W of East Pt, Santa Rosa I, 43-45 fms (2:1.8, 4.0 cm) (Velero 1390- 41); 2 mi W of Church Rock, Santa Catalina I, 45-53 fins (1:1.8 cm) (Velero 1321-41); S of Pyramid Cove,

San Clemente I, 55-69 fins (1:2.9 cm) (Velero 1012-39); 2.5 mi SE of Bennett Pt, San Miguel I, 57 fms

• (3:1.3- East Pt,1.9Santa cm) R(Vele osa roI, 47- 139409-41); fms (31)mi(Vel offeroSeal1289-41 Beach,);11offfinsGull(2:3.1, Island3.3I,cm) 39-40(Velero fins (3:2. 1158-4 1-2.30);cm2.6) (mi VelEeroof

1194-40); off Abalone Pt, Laguna Beach, 54-57 fms (13:1.2-2.6 cm) (Velero 1131-40); S side of Santa Cruz

I I, 39-40 fms (10:2.3-5.4 cm) (Velero 1191-40); 4.5 mi SE of East Pt, Santa Rosa I, 52 fms (9:2.5-5.1 em) (Velero 1387-41); S of San Miguel I, 130-138 fins (2:3.1-3.9 cm) (VeIero 986-39a); off Redondo Beach, 75

fms (4) (Bureh 3856); off Redondo Beach, 96-120 fins (1:3.1 cm) (Velero 1137-40); N of west Anacapa I,

I 44-45 fms (2:2.7, 3.0 cm) (VeIero 1857-49); Catalina Harbor, Santa Catalina I, intertidal (1:2.4 cm) (Velero

i i 239

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200pm 100//rn i

Figure 9.24. Pentamerapopulifera: A, ossiclesfrom body wall; B, supporingtables. !

I

1368-41 ); Avalon Bay, Catalina I, 98-116 fms (1:0.7 cm) (VeIero 1149-40); 1.5 mi NW of Cavern Pt, Santa

Cruz I, 54-56 fms (16:0.9-3.2 cm) (Velero 1300-41 c); off W end of Santa Catalina I, 134-150 fins (1:4.6 cm) 1

(Velero 1201-40); 3 mi NW of Anacapa I light, 47 fms (6:0.8-3.5 cm) (Velero 1267-41); 4.2 mi SE of East i

Pt, Santa Rosa I, 46 fms (2:3.0, 4.5 cm) (Velero 1290-41); 4.5 mi W of Church Rock, Santa Catalina I, 51

fms (1:3.9 cm) (Velero 1318-41); off White Cove, Santa Catalina I, 105 fms (2:1.4, 3.4 cm) (Velero 1373- _:,

41a); 1.5 mi SW of Gull I, Santa Cruz I, 48 fms (1:4.8 cm) (Velero 1435-41); 6.5 mi ESE of South Pt, Santa _|

Rosa I, 57 fms (1:3.9 cm) (Velero 1392-41); SW of Castle Rock, San Clemente I, 40-46 fms (1:3.1 cm)

(Velero 1328-41); 10 mi SE of Long Beach, 22-25 fins (1) (VeIero 1159-40); off Bluff Cove, 50-75 fms (1)

(Burch 40118); 1.5 mi S of Cron Pt, San Miguel I, 41-43 fms (6:2.3-5.3 cm) (Velero 1412-41); off Bluff i

Cove, 35 fms (4:2.0-2.6 cm) (Burch 40129); 1.5 mi SE of west end, Santa Catalina I, 40-50 fms (1:2.3 cm) I

(Velero 1311-41); 2.5 mi NW of Anacapa I light, 48-51 fms (4:3.1-5.4 cm) (Velero 1268-41); 2.75 N of

West Pt, Santa Cruz I, 55 fins (1:4.0 cm) (Velero 1436-41); N of Anacapa I, 15 fms (2:2.3, 3.1 cm) (Velero i

878-38a); 2 mi SW of Cardwell Pt, San Miguel I, 34-35 fms (3:1.3-2.1 cm) (Velero 1413-41); 10 mi S of

San Pedro breakwater, 81-83 fms (1:2.0 cm) (Velero 1229-41 ); 3.5 mi S of Hueneme, 29-30 fms (1:2.4 cm)

(Velero 1274-41a); off Ten Mile Creek, Or, 20-40 fms (4) (Velero 1471-42); San Miguel Passage, 37-39 !

fms (10:1.5-6.0 cm) (Velero 990-39b); 0.5 mi N of Platt Pt, Santa Cruz I, 36-47 fms (1) (Velero 1303-41a); i

1.5 mi SE of Point Mugu, 26-30 fms (8) (Velero 1275-41); (2) (BCPM FRB 63-204); sandy bay behind

Rugged Pt, Kyuquot Channel, 5 fms (1) (BCPM 980-344-1); Haul 2, San Pedro, Ca (1:1.5 cm) (CAS Station I 

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I XIV); off San Onofre, 91 m (1:1.6 cm) (SlO E2679); off Morro Bay, 99 m (1) (SIO E 2686); Pt Sal, 132 m

(2) (SIO E 2687); MMS CAMP 3-4, R-2.---Mexico: off San Benito I, Baja California, 92-95 fms (2:2.4, 5.7

i cm) (Velero Desc1010-39); 8.5 mi S of San Benito I, Baja California, 71-72 fms (1:4.6 cm) (Velero 1252-41). ription. Medium-sized, up to 6 cm long, body shape variable, often fusiform, tapering to

pointed ends; body wall thin, so packed with ossicles as to be rough to the touch. Podia cylindrical. Ossicles

I innumerous body wall teethmostl onytop;tables occasional with round platestopresent. star-shapedSupporting disk, 0.10-0.42 tables withmmlowin diameter, 2-pillared with spire,steither out spire bluntwithor

pointed, never flattened as in P. pseudocaIcigera. Well-developed end plate present. Introvert with oblong

i to roundBperforated plates. Tentacles with plates and rods, sometimes absent. iology. This species is common in soft-bottom habitats, intertidally to 150 fms. In Puget Sound

Pentamerapopulifera spawns in spring (McEuen, 1988).

I' Remarks. Small specimens (< 1.5 cm) ofPentamerapopulifera andP. pseudocalcigera are difficult

to differentiate. However the spires on the supporting tables are usually much longer and flattened in P.

pseudocalcigera than in P. populifera.

t Type Locality and Type Specimens. Washington, Puget Sound, holotype, lost; C. tenuicoriata:

California, Monterey Bay (USNM E1201).

l Distribution. West coast of North America: Puget Sound to San Benito Island, Baja California.

Pentamera pseudocalcigera Deichmann, 1938

I. Figure 9.25

i Pentamera pseudocalcigera Deichmann, I938:106.--Panning, 1949:465. Cucumaria calcigera H.L. Clark, 1901b: 165, 1901c:492.--Edwards, 1907:54, text-fig. 5-11. NotPentamera

calcigera (Stimpson).

I Material Examined. Oregon: USNM Acc. No. 108-05, 7 frns (2:4.1, 5.0 cm); USNM Oregon coll,

AD27 (1:5.0 cm); USNM Oregon coil, AD70, 200 m (1:7.0 cm); Velero I471-42, off Ten Mile Creek, 20-

40Bay,fins, 252sanfmsd (2:5. (1:3.01,5.5cm);cmUSNM ).--Alaska: E2294 Albat(1:2.0 ross cm); 4240,Alba(l)tross .--Calif4457, ornia:6 miUSNM 21°W 30581, Pt Pifiosofflighthouse, San Luis Obispo 40-46

fms (2:2.0, 5.0 cm); Albatross 4558, 2 mi 79° W Pt Pifios lighthouse, 28-40 fms (1); Albatross 4464,

I Montere 124°44%yV, ,21-3 200 1fmfms (1:5.(51);cmUS); NMUSNMBCFBCF , 46°08'N, , 46°02.4'N, 124°13'W, 124°3750.8'Wfins , 10(1:4.2 0 fins c(1m); :4.2UScm) NM; USBCF, NM 45Ac°c.60'N, No.

152709, Monterey Bay, 50-60 fms (1:6.4 cm); Velero 992-39, off Santa Rosa I, 130-230 fins, mud, grovel

t (3: 403.3-4.5 cm); MCZ 2080, 34°09'30"N, 120°40'00"W, 150 fins (1); BCPM 167-1, 8°41q_1123°29.1W¢, 38- ft (1); CS DOC Deep, G-16 (1); OCSD B41 (1); OCSD B27 (1); ACE dumpsite, BD2-4IA, 307 m; MEC

123a, 526 ft (1); MBC 081-BSS-01-TX, 34°21.263'N, 120°28.827%V, 980 ft (1); CAS, off Trinidad Head,

l 65 fins (1:3.0 cm); BLM 24018CH (2:2.2, 5.0 cm); BLM 23283 (1:4.2 cm); USNM BSS-038; SBNMH BSS-081.--Mexico: SIO E2586, between San Benitos and Cedros I, Baja California, 170-180 fins (4:1.5-

4.2 cm);

I, B Description. Medium-sized, globose, U-shaped form, tapering to blunt ends, up to 10 cm long. ase of tube feet conical. Body wall rigid, packed with ossicles. Ossicles in body wall of small specimens

mostly round to star-shaped tables, 0.15-0.48 mm in diameter, similar to those of Pentamera populifera.

i Wpredominate. ith growth, Spire numberof of tables decreases and large, irregularly shaped plates, 0.13-0.72 mm in diameter,

supporting tables sometimes elongate, flattened. Introvert with open, spiny plates,

0.20-0.40 in diameter. Tentacles with spiny rods.

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i 241
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Figure 9.25. Pentamera psuedocalcigera: A, plates from body wall; B, tables from body wall; C, supporting •

tables; D, plate from introvert. il

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I Remarks. The ossicles in small Pentamera pseudocalcigera and P. popuIifera are similar, but P.

pseudocalcigera can usually be differentiated by the globose body form, conical tube feet, and the flattened

i inspireP. onpseudocalcigera the supporting than tables.in The P. populifera plates in.theInintrovert two lotsarefromgenerally Baja California larger (0.20-0.40 in the Scripps vs. 0.08-0.29 Instituteram)of

i Oceanography, tables predominated in large as well as small specimens. In a few large specimens, tables

1_ predominated in the dorsal body wall but plates predominated in the ventral body wall. In two lots from

It Southern California, the dorsal and ventral body were fused so that the anus and mouth were contiguous.

These specimens were typical of P. pseudocalcigera in every other respect and are considered a variant of

I P. pseudocalcigera Type Loca.lity and Type Specimens. off California, syntypes (USNM 164I 1, 30568, 30569, 30572,

30573, 30578, 30580, 30581, 30585, 30669, 30681, E2294, E2386, E2387).

i Distribution. Southeast Alaska to San Benitos Island, Baja Califomia, from 7 to 250 fins (12 to 450 m).

i Pentamera pseudopopu Figure h'fera9.26Deiehmann, 1938

i Pentamera pseudopopulifera Deichmann, 1938:107.--Panning, 1949:465.

Material Examined. California: Velero 1131-40, off Abalone Pt, Laguna Beach, 54-57 fins, mud

i (10:2.8-5.2 897-38, off Point cm); Vele SantaroBarbara, 1130-40,33offfms, Abalone mud (3:4.5-6.3 PL Lagunacm);Beach, Velero25-2 1292-41, 7 fins, 6mud mi E(8of:0.8-3 East.9Pt,cm); SantaVel Rosa ero

I, 28-30 fms, rock (3:1.8-3.1 cm); Velero 1142-40, off Pt Vicente lighthouse, 17-41 fins, mud, sand (1:4.5

I, cm); SouthVelero Pt, Santa1191-40, Rosa S side of Santa Cruz I, 39-40 fms, sand (2:4.7, 5.5 cm); VeIero 1281-41, 3 mi E of I, 23-26 fms, sand (1:1.0 cm); Velero 1160-40, 11.5 mi SE of Long Beach, 32-52 fms,

mud, sand shell (3:1.1-2.5 cm); Velero I205-40, S side of San Nicolas I, 20-23 fms, sand (1:4.3 cm); Velero

l fms 1143-(1:4.1 40, offcm); Portuguese Velero 1275-41, Pt, 16-201.5fins, misand SE ofPt (1:2.0Mugu, cm); 26-30 Velerofms, 1619-48, mud (4:3.2-3.8 White Covcm); e, Santa MECCatalin 313A,a I,7226ft

(1:4.7 cm); MCZ 473, Corona del Mar, 77 fms (1:2.5 cm).

I Description. Medium-sized, up to 6.5 cm long. Body shape variable, often fusiform, curved, sometimes U-shaped. Preserved specimens often purplish brown. Body wall ossicles mostly small roundish

tables, 0.04-0.17 mm in diameter, with stout spire tipped with bifurcate teeth; plates rare.Supporting tables

i with approaching 2 pillars, also with bifurcate teeth. Well-developed end plate present. Introvert with delicate plates, rosettes. Tentacles with oblong plates and rods.

Biology. This species is common in soft-bottom habitats in the Southern California Bight, in shallow

i water to 114 m.

Remarks. Pentamera pseudopopulifera differs from P. populifera in that the disk of the tables is

smaller, more regularly shaped and never star-shaped. The bifurcate teeth on the body wall tables and the

i supporting tables are also diagnostic.

Type Locality and Type Specimens. California, Point Dume, holotype (USNM 39159)

l Distribution. Southern California, shallow water to 57 fins.

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Figure 9.26. Pentamera pseudopopulifera: A, ossicles from body wail; B, supporting tables.

!

Genus Stolus Selenka, 1867 II

II

Type Species. Thyone buccalis Stimpson, 1856, by subsequent designation in Panning (1949).

Diagnosis. Ossicles in body wall plates, tables absent. I

Stolus trachyplaca (H.L. Clark, 1924)

Figure 9.27 I

Cucumaria trachyplaca H.L. Clark, 1924:56. ,i

II

Cucumaria cosmotyrsitus Wells, 1924:119, text fig. 3, pl. 3, fig. 3.

Pseudothyone trachyplaca Panning, 1949:456. i

g Material Examined. California: SBMNH, Pelican Cove, Santa Cruz I, 20-60 ft (1); SBMNH 488,

45m, reef midway between Twin and Platts Harbor, Santa Cruz I, 10-50 ft; USNM E2276, Monterey Bay I

(1:2 cm); USNM Acc. 205678, off Ford Pt, Santa Rosa I, 92 ft (1:2.1 cm); USNM BRA-002.----Canada: i

BCPM FRB63-204; BCPM 978-199-4, Possession Pt, near Sooke, 48020.4 ' N, 42.8 ° W, < 30 ft (I). el

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I Description. Small, cylindrical form, up to 3 cm long, with stout tube feet. Body wall ossicles

plates, usually oblong, sometimes irregularly shaped, 0.1-0.2 mm long; undeveloped plates with 2 enlarged

1 central knobs, holes sometimes and 3-5built smaller into holes a massin resembling either end. More a spire. developed Supporting platestables with varying with 4 pillars numbersending of spines in spiny and

mass. Well-developed end plate present. Inlzovert with oblong reticulated plates. Tentacles with various

i sized plaRemarks tes often. with The2tyenlarged pe specimens centralofholes. Cucumaria trachyplaca and C. cosmotyrsitus are synonyms.

Clark's (1924) description antedates Well's (1924) description by several months. Since the ossicles in the

i body wall are derived from plates, I have placed this species in the genus Stolus. Type Locality and Type Specimens. Cucumaria trachyplaca: British Columbia, Queen Charlotte

Sound, Alert Bay, 20 m (National Museum of Canada, Acc. 85-4, cat. No. 272); C. cosmotyrsitus: California,

I off Monterey, shale, 24 m (National Museum of Natural History, USNM E 1200). Distribution. North Pacific Ocean, at least from British Columbia to Santa Cruz Island, California,

intertidal to 100 ft (30 m), rocky habitats.

1

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Figure 9.27. Stolus trachyplaca: A, plates from body wall; B, supporting table; 12,platesfrom introvert. !

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Genus Thyone Jaegger, 1833 i

Type Species. Thyonefusus (O.F. Mtiller, 1766), by subsequent designation. I

Diagnosis. Ossicles in body wall tables, sometimes reduced. Ossicles in introvert rosettes, tables B

absent.

m

Thyone montereyensis (Deichrnann, 1938) a

Figure 9.28 i

Pentamera montereyensis Deichmann, 1938a, p. 108, text-fig.

Thyone montereyensis Panning, 1949, p. 467. i

Material Examined. California: Long Beach breakwater, Ca (2:4.3, 4.8 cm) (PBM); E of pier, ./

Johnson's Lee, Santa Rosa I, 20-40 ft (1:2.2 cm) (MB). i

Description. Medium-sized cylindrical form, up to 5 cm long; interambulacral feet absent. Ossicles i

small oval buttons, 0.025-0.055 mm long, with 2 central and up to 8 marginal knobs. Supporting tables with

marginal knobs, spire reduced to knobs. Well-developed end plates and perforated rods present. Introvert i

with round to oblong perforated plates often with 2 enlarged central holes and 6-20 additional holes. Two

central knobs often present, additional knobs sometimes present. Tentacles with plates and rods.

I

I

o 0

A ooC ,|

i

0 0 '

Figure 9.28. Thyone montereyensis: A, buttons from body wall; B, supportingtables; C, irregular plate from i

specimen from Long Beach; D, plates from introvert. |

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I Remarks. The generic placement of this species is uncertain. Since interambulacral feet are lacking,

this species does not belong in the genus Thyone sensu Pawson and Miller (1981). The presence of two

I central genus Havelockia knobs on thewithplates the inspirethe onbodthey wall tablesandreduced. the introvert However, suggest because that thisit isspnot eciesclearis athat member this species of the

belongs in the genus Havelockia, I am retaining Panning's (1949) designation. Irregularly shaped plates,

I 0.06-0.15Tymmpe Linocdiameter, ality and were Type present Spec in a large specimen taken from Long Beach breakwater. imens. California, Pacific Grove, intertidal (Museum of Comparative

Zoology, MCZ 1840), ossicles dissolved.

l. to at leastDistribution. 14 rm California, from Pacific Grove to southern California, hard-bottom habitats, intertidal

I Literature Cited

i Brandt, J.F. 1835. Echinodermata ordo Holothurina. In: Podromus description_is animalium ab H. Mertensis in orbis terrarum circumnavigatione observatorum. 1:42-62.

l Bush, M. 1921. Revised key to the echinoderms of Friday Harbor. Publication of the Puget Sound Biological

Station 3:65-77.

Clark, H.L. 1901a. Echinoderms from Puget Sound: observations made on the echinoderms collected by the

I parties from Columbia University in Puget Sound in 1896 & 1897. Proceedings of the Boston

Society of Natural History 29:323-337.

i Clark, H.L. 190lb. The holothurians of the Pacific coast of North America. Zoologischer Anzeiger 24:162- 171.

Clark, H.L. 1901c. Synopses of North American invertebrates. 15. The Holothuroidea. American Naturalist

i 35:479-496.

Clark, H.L. 1902. Notes on some North Pacific holothurians. Zoologischer Anzeiger 25:562-564.

i Clark, H.L. 1907. The apodus holothurians. A monograph of the Synaptidae and Molpadidae. Smithsonian Contributions to Knowledge 35:1-231.

I Clark, H.L. of 1913. Echinoderms from Lower California with descriptions of new species. Scientific Results the Expedition to the Gulf of California in charge of Dr. C.H. Townsend by the U.S. Fisheries

Steamship Albatross 1911. Bulletin of the American Museum of Natural History 32:185-236.

i Clark, H.L. 1922. The holothurians of the genusStichopus. Bulletin of the Museum of Comparative Zoology 65(3):39-74.

i Clark, H.L. 1924. Some holothurians from British Columbia The Canadian Field-Naturalist 38:54-57. Clark, H.L. 1935. The Holothurian genus Caudina. Annals and Magazine of Natural History, series 10,

15:267-284.

l Cowles, R.P. 1907. Cucumaria curata sp. nov. Johns Hopkins University Circular Series 3, No. 195: 1-2.

Deichmann, E. 1930. The holothurians of the western part of the Atlantic Ocean. Bulletin of the Museum of

I Comparative Zoology 71(3):43-226.

Deichmann, E. 1937. The Templeton Crocker Expedition. IX. Holothurians from the Gulf of California, the

i west coast of Lower California and Clarion Island. Zoologica, New York 22:161-176.

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Deichmann, E. 1938a. Eastern Pacific Expedition of the New York Zoological Society. XVI. Holothurians g

from the western coasts of lower California and Central America and from the Galapagos Islands.

Zoologica, New York 23:361-387. •

Deichmann, E. 1938b. New holothurians from the western coast of North America and some remarks on the U

genus Caudina. Proceedings of the New England Zoological Club 16:103-115. Im

Deichmann, E. 1939. A new holothurian of the genus Thyone collected on the Presidential Cruise of 1938. I

Smithsonian Miscellaneous Collections 98(12):1-7.

Deichmann, E. 1941. The Holothuroidea collected by the Velero 11Iduring the years 1932-1938. Part I. i

Dendrochirota. Alan Hancock Pacific Expeditions 8(3):61-153.

Edwards, C.L. 1907. The holothurians of the north Pacific coast of North America collected by theAlbatross II

in 1903. Proceedings of the United States National Museum 33(1558):49-68. II

Edwards, C.L. 1910. Four species of Pacific Ocean Holothurians allied to Cucumariafrondosa (Gunnerus). m

Zoolologisches Jahrbuch, Abteilung Systematik 29:597-612. I

Eschscholtz, F. 1829. Zoologischer Atlas: enthaltend Abbildungen und Beschreibungen neuer Thierarten

w_ihrend des Flottencapit_s von Kotzebue zweiter Reise um die Welt. II. Heft: 12-13. _

It Heding, S.G. 1928. Synaptidae. Videnskabelige Meddeleser fra Dansk Naturhistorisk Forening, K_benhavn

85:105-323.

m

Heding, S.G. 1931. On the classification of the Molpadids. Videnskabelige Meddeleser fra Dansk I

Naturhistorisk Forening, Kcbenhavn 92:18-284.

Koren, J. 1844-1845. Beskrivelser over Thyone fusus og Cuviera squamata. Nyt Magazin for I

Naturvidenskaberne (Christiana) 4:203-225. il

Lambert, P. 1985. Geographic variation of calcareous ossicles and the identification of three species of sea Ii

cucumbers (Echinodermata: Hotothuroidea) from the eastern Pacific.In: Echinodermata. Proceedings il

of the Fifth International Echinoderm Conference, Galway, 24-29 September 1984: 437-443.

Lambert, P. 1986. Northeast Pacific holothurians of the genus Parastichopus with a description of a new i

species, Parastichopus leukothele (Echinodermata). Canadian Journal of Zoology 64:2266-2272. 21

Lambert, P. 1990. A new combination and synonomy for two subspecies of Cucumaria fisheri Wells n

(Echidodermata: Holothuroidea). Proceedings of the Biological Society of Washington 103(4):913- |

921.

Lampert, K. 1885. Die Seewalzen, Holothuroidea, eine systematische Monographie. Wiesbaden. 310 pp. N

u

Ludwig, H. 1881. Uber eine lebendiggeb_ende Synaptidae und zwei andere neue Holothurienarten der

Brasilianischen Kiiste. Archiv fiir Biologie 2:41-58.

Ludwig, H. tl 1894. Reports on an exploration off the west coasts of Mexico, Central and South America and

off the Galapagos Islands, in charge of Alexander Agassiz by the U.S. Fish Commission Steamer

Albatross. Memoirs of the Museum of Comparative Zoology 17(3):1-183. n

m,

Ludwig, H. 1904. Brutpflege bei Echinodermen. Zoologisches Jahrbuch, Supplement 7:683-699.

McEuen, F.S. 1988. Spawning behaviors of northeast Pacific sea cucumbers (Holothuroidea: Echinodermata). ii

Marine Biology 98:565-585.

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I McEuen, F.S. and F.-S. Chia. Development and metamorphosis of two psolid sea cucumbers, Psolus

chitinoides and Psolidium bullatum, with a review of reproductive patterns in the family Psolidae

I (Echinodermata: Holothuroidea). Marine Biology 109:267-279. Mitsukuri, K. 1912. Studies on actinopodus Holothuroidea. Journal College of Sciences, Tokyo 29(2): 1-

234.

I Oken, L. 1815. Lehrbuch der Naturgeschichte. Zoologie 3:351-352.

Oshima, H. 1915. Report on the holothurians collected by the U.S. Fisheries Steamer Albatross in the

i northeastern 48:213-291. Pacific during the summer of 1906. Proceedings of the United States National Museum

i Panning, A. 1949. Versuch einer Neuordnung der Familie Cucumariidae. Zoologisches Jahrbuch, Systematik 78:404-470.

Panning, A. 1964. Bemerkungen iiber die Holothurien -Familie Cucumariidae (Ordnung Dendrochirota) 4.

I Teil. Die Gattungen Steroderma, Staurothyone und Trachythyone. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 61:159-174.

i Panning, A. 1966. Bemerkungen fiber die Holothurien -Familie Cucumariidae (Ordnung Dendrochirota) 5. • Tell. Die Gattungen Heterothyone Panning 1949 und Leptopentacta H.L. Clark 1938. Mitteilungen

aus dem Hamburgischen Zoologischen Museum und Institut 63:51-69.

I Panning,Mitteilungen A. 1971. Bemerkungen iiber die Holothurien - Familie Cucumariidae (Ordnung Dendrochirota). aus dem Hamburgischen Zoologischen Museum und Institut 67:29-51.

i Pawson, D.L. 1967. The psolid genus Lissothuria. Proceedings of the United States National Museum 122:1-17.

Pawson, D.L. 1970. The marine fauna of New Zealand: Sea Cucumbers (Echinodermata: Holothuroidea).

i Bulletin of the New Zealand Department of Scientific and Industrial Research 201:1-69.

Pawson, D.L. and H.B. Fell. 1965. A revised classification of the dendrochirote holothurians. Breviora

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of two new species (F_,chinodermata: Holothuroidea). Proceedings of the Biological Society of

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species from Tasmania. The Papers and Proceedings of the Royal Society of Tasmania 101:31-35.

I Rutherford, J.C. 1973. Reproduction, growth and mortality of the holothurian Cucumaria pseudocurata.

Marine Biology 22:167-176.

I Rutherford, holothurian J.C. 1977. Cucumaria Geographical curata. variation Marine Biology in morphological 43:163-174. and electrophoretic characters in the

Selenka, E. 1867. Beitriige zur Anatomie und Systematik der Holothurien. Zeitschrift fiir Wissenschaftliche

I Zoologie 17:291-372.

Smith, E.H. 1962. Studies of Cucumaria curata Cowles, 1907. Pacific Naturalist 3(5):233-246.

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Stimpson, W. 1857. The crustacea and echinodermata of the Pacific shores of North America. Journal of the I

Boston Society of Natural History 6:524-526.

Stimpson, W. 1864. Descriptions of new species of marine invertebrata from Puget Sound collected by the I

naturalists of the Northwest Boundary Commission. Proceedings of the Academy of Sciences,

Philadelphia 16:153-161.

Therl, H. 1886. Report on the Holothuroidea. Report on the Scientific Results of the H.M.S. Challenger I

during the years 1873-1878. Part 39, Zoology 14:1-290.

Wells, H.G. 1924. New species of Cucumaria from Monterey Bay. Annals and Magazine of Natural History, I

series 9, 14:113-21.

Yingst, J. 1972. A new species of rock dwelling dendrochirote holothurian from Catalina Island. Bulletin of I

the Southern California Academy of Sciences 71(3): 145- I50.

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I 10. HEMICHORDATA: ENTEROPNEUSTA

1 by

Keith H. Woodwick _

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i Introduction

Enteropneusts (acorn worms) are soft bodied marine worms of the Phylum Hemichordata. Most

I species have been described as burrowers in sand and mud of the intertidal and subtidal. They average 10 to 50 cm in length and are about the diameter of a pencil. The first species, Ptychoderaflava, was described by

Eschscholtz (1825) and there are now more than 70 known species in 13 genera and 4 families.

!

Morphology

I Enteropneusts are fragile burrowing animals often collected as incomplete specimens. Complete

specimens have three main divisions of the body, an anterior muscular proboscis, a central cuff-like collar,

I and a posterior elongate trunk including branchial, esophageal, hepatic, and intestinal regions. Collections usually contain specimens with only proboscis, collar, and the branchial region of the trunk. External features

used in placing specimens in families and genera include the shape and size of the proboscis, shape and size

i of the collar and its ratio to that of the proboscis, presence or absence of genital wings in the anterior trunk

• and hepatic caeca in the mid-trunk area. Presence of branchial slits externally versus small branchial pores

as well as gonad positioning are other useful characteristics. Histological features and internal structures,

i their presence or absence, and variations, are important in placing specimens in family, genus, and species. Proboscis muscle pattern, glomerntus, proboscis skeleton and crura, buccal diverticulum, neurocord and

epidermal zones of the collar, shape and size of gill bars and presence or absence of synapticules as well as

i epithelial variations throughout the gut of the trunk, all provide information and suites of characteristics important to classification.

I Taxonomic History

I Ptychodera flava, the first known enteropneust, was described by Eschscholtz (1825) as a new species of sea cucumber from the Marshall Islands. In the next 30 years only two more species were described,

Balanoglossus clavigerus Delle Chiaje (1829) from Naples and Stimpsonia aurantiacus by Girard (1853)

i from After the Kowalevsky Carolina coast. The latter was described as a nemertean and later transferred to Balanoglossus.

(1866) described the gill slits in B. clavigerus, Gegenbaur (1870) used this characteristic

in establishing Enteropneusti (Enteropneusta). Bateston (1885) proposed using Hemichordata for

i Enteropneusti phylum (Hyman, and1959). placed it in Chordata. Most present day workers consider Hemichordata to be a separate

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I 1Department of Biology, California State University, Fresno, California 93740-0073.

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A period of intensive systematic investigation of entempneusts occurred between 1891 and 1901; i[_

eight new genera and 36 new species were described. They comprise about 75% of the presently known

genera and 50% of the known species. Spengel (1893, 1901) was the dominant worker of that period. Since _,

1901 systematic research has been meager and sporadic. Classification now includes 73 species in 13 genera l

and the following families: Ptychoderidae, Spengeliidae, Hardmaniidae, and Saxipendiidae (Hyman, 1959;

Woodwick and Sensenbaugh, 1985). In the last 30 years there have been a number of special subject studies, lira

for example, histology and cytology (Norrevang, 1965; Welsch and Storch,1970; Saita et al., 1978), |

development (Hadfleld, 1975), larval dispersal (Scheltema, 1987), ecology (Thistle, 1980), and presence of

organohalogens (Higa et al., 1980; King, 1986; Corgiat et al., 1993). 1

Biology I

Nearly all known species of enteropneusts are found in the intertidal zone or in shallow waters.

Generally they inhabit burrows in sand and/or mud, although a few species occur under rocks or are associated J

with marine plants. They feed upon the sediment using cilia and mucus to move materials into and through II

the gut as they extract items of food. Undigested materials are deposited at the surface as piles of castings

and in quiet waters are useful indicators of entempneust populations. Sexes are separate and gametes are

released into the environment; development may be indirect with tornaria larvae or direct, as it is in all l

members of Harrimaniidae.

Only four species have been reported from depths greater than those of the immediate subtidal. I

They are Spengelia sibogae (275 m: Spengel, 1907), Glandiceps talaboti Marion (30-350 m), Saxipendium U

coronatum (2478 m: Woodwick and Sensenbaugh, 1985), and Glandiceps abyssicola (4500 m: Spengel,

1893). I

Methods n

Many genera can be identified using external characteristics but identification of other genera and

all species is dependent on individual and suites of characteristics of the internal anatomy. The internal

anatomy is revealed through study of serial cross and sagittal sections. For this reason, straight, clean, well- U

preserved complete specimens are desirable. Hyman (1959) summarized the usual situation in enteropneusts

wherein most species are known only by the original description which in addition was usually based on one

or a few and often imperfect specimens. Imperfection results from a number of enteropneust features. They I

are fragile and likely to be within the sediment so that whole specimens are rarely taken; it is also likely that

even if many parts of specimens are collected they will not include all trunk regions. Enteropneusts are

ordinarily covered with mucus and sediment which remain attached to their bodies on fixation. Sediment U

moving through the gut is also retained on fixation. These features limit the taking of clean complete specimens

when collections are made individually in the field and they may be magnified during benthic surveys. [_

When II possible, live specimens should be sorted from the sediment. Sometimes breaking the sediment

into large pieces will reveal the burrow and the organism and further careful breaking will free the specimen.

Live specimens should then be placed in large containers of clean sea water to permit evacuation of the gut. n

Several changes of the sea water may be needed and surface sediment may be released with their movement It

or assisted utilizing a camel's hair brush. Clean specimens should then be fixed by holding them at the

posterior end with a pair of forceps and dipping them in a tall jar of fixative (e.g., Bouin's fluid). Gravity jlJ

will assist in straightening the specimen as fixative is added to the posterior end and allowed to flow down II

into the jar. When fixation is complete release the specimen into the jar and store on its side for several days.

Transfer specimen to 50% alcohol, and then wash several times in 70% alcohol (not water) to clear the I

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I picric acid (Galigher and Kozloff, 1971). Specimens can then proceed through an alcohol, toluene, paraffm

series for embedding. Prepared paraffin blocks are then serially sectioned at 10-15 pm and slide sections

I s

sertainiaeldcrwith oss and Harris' sagitHemat tal secoxylin tions. and MosEosin t publiY. sheThe d deanascrtoiptions my is rhav econs e included tructed frlabe om llmicrosc ed drawing opic sstudy of critica of thel

diagnostic features and areas.

i Systematic Account

i Specimens from the benthic survey of the Santa Mafia Basin and the Western Santa Barbara Channel

include members of four genera and three families. It is likely that they represent four new species. Genera

I represen Glossobalanus ted include in Ptychoderidae. Saccoglossus Woodwick and Stereobalanus (1955 ) found in Harrimaniidae, representativesSchizocardium of the same four in Spengeliidae, genera in samples and

of the benthos of San Pedro Basin.

I A total of 54 identifiable specimens were taken at 8 of the 28 MMS Phase 1I soft-substrate stations. (A unit which included proboscis, collar, and some part of the trunk was considered a specimen. Parts of the

trunk, collar, or proboscis were considered fragments.) Stereobalanus sp. was the dominant form. It was

I taken R-7. Balanoglossus at Stations R- 1,R-4, R-7, and R-8. Seventeen of the 28 specimens came from a depth of 565 m at Sta. sp. was represented by 19 specimens and 17 of these were from 90 m at Sta. R-8.

Schizocardium sp. was the largest form but was represented by only a total of 4 specimens at 3 stations (RI were 8, R-l,collected and R-4atat390, different 91, andstations 92 m, respectively). (R-I, PJ-1, andSaccoglossus PJ-11 at 91,145, sp. wasandthe136smam, llesrespectively). t form. The 3There specimens were

3 unidentifiable specimens and many fragments.

I morphology. For purposes They willof survey work, specimens of the four genera can be sorted on the basis of external either have or lack genital wings or lappets, tong or short probosces, and large or

small eggs (KEY I). Because the muscular proboscis is the most easily recognized and regularly collected

I part of an enteropneust, its muscle pattern was chosen as the most diagnostic internal characteristic manifested in serial cross sections (KEY ID. In addition to the two keys, a list is provided which includes many positive

family characteristics, as well as, characteristics specific for each genus. The selected family characteristics

I are useful in further separating the four genera of this study and will be helpful in identification of specimens taken in other surveys or present in existing collections.

I Family Ptychoderidae Spengel, 1893

'l Diagnosis. Proboscis short,conical; longitudinal muscle,radial pattern of many thin bands. Collar

,It as long as broad. Trunk with pronounced external regionation; short curved gill bars; synapticules;

parabranchial ridges;ventaal digestive area in pharynx; wide long genital wings; plate-shaped hepatic caeca;

ciliated intestinal grooves (ridges). Development indirect, with tomaria larvae.

Genus Balanoglossus Delle Chiaje, 1829

I Diagnosis. Cauliflower organ lacking; lateral septa extend only half way into genital wings; small

external branchial pores; gonads dorsal and lateral.

I Representatives Remarks.of There the genus are nohavedescribed been found species in theof Balanoglossus intertidal at several from localities the west coast and atofabout North50mAmerica. in the

San Pedro Basin. Comparison of the Santa Maria form with the 22 described species of the genus is in

I progress.

I 253
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Ii

Family Spengeliidae Willey, 1899 I!

Diagnosis. Proboscis short, ovate; longitudinal muscle, homogeneously arranged; circular muscle,

thick layer; vermiform process; chondroid tissue well-developed. Collar shorter then broad. Trunk with U

circular muscle, internal and well developed. Development indirect, with tornaria larvae.

II

Genus Schizocardium Spengel, 1893 I

Diagnosis. Vermiform process long; gill bars long; hypobranchial area narrow; gonads lateral only; an

esophageal pores present, single and paired; hepatic caeca present and finger-shaped. II

Remarks. There are two species ofSchizocardium, S. brasiliense, and S. peruvianum, described by

Spengel in 1893. The species from Peru is represented by a single incomplete specimen. The Santa Maria n

form fits the characteristics of the genus but preliminary observations suggest that it has a suite of il

characteristics different from the described species. In comparison to S. brasiliense its proboscis is more

elongate in shape (less pyramidal), circular muscle layer of the proboscis is thinner, vermiform process I

begins in the posterior one-third of the proboscis, auricles are longer than the vermiform process, and the I

proboscis skeleton has a high sharp keel. In comparison toS. peruvianum its proboscis is longer, the vermiform

process begins in the posterior one-third of the proboscis, and the auricles are longer than the vermiform II

process. Presence of the Santa Maria Basin form at a depth of 91m and in the northeast Pacific are additional 11

important factors.

Im

Family Harfimaniidae Spengel, 1901 I

Diagnosis. Proboscis lacks cauliflower organ and vermiform processes. Collar lacks neurocord i

nerve roots and peripharyngeal cavities. Trunk lacks genital wings, hepatic caeca, parabranchial ridges, and

,qp

synapticules. Eggs large (150-400 m); development, direct.

II

Genus Saccoglossus Schimkewitsch, 1892

Diagnosis. Proboscis long, cylindroid; longitudinal muscle of proboscis in concentric rings; collar i _

broad as long; gonads simple. U

Remarks. One species of Saccoglossus, S. pusillus, has been described from intertidal areas in i

California. Specimens of the genus were reported from 40-50 m in the San Pedro Basin. The Santa Maria !

form is very small. The proboscis, collar andbranchio-genital region of an incomplete specimen measured

1.05 cm; total length is estimated to be 2.0-3.0 cm. The number of concentric rings (3) of longitudinal

muscle in the proboscis is smaller than that of most species and most similar to S. otagoensis (3-4) and S. i

cambrensis (4-6). The Santa Maria form is different in being taken at a depth of 91 m and in having small

ova (160 0m). The most similar species is S. horsti with ova measuring 230 × 70 pare.Other species have

ova of 250, 310, 375, and 400 lam. Additional morphological comparisons are underway, i

Genus StereobaIanus Spengel 1901 i

Diagnosis. Proboscis short, conical, and tinged anteriorly; longitudinal muscles, radial pattern n

(heavier bands); collar shorter than broad; elongate dorso-ventral gill slits open directly to the exterior; two

pairs of genital lappets; gonads lateral and dorsal, i

Remarks. Stereobalanus was established in 1901 by Spengel for a species he described as n

Balanoglossus canadensis in 1893.The specimens from Sta. R-7 in the Santa Mafia Basin fit the characteristics

for Stereobalanus listed above; however, they do not fully match the geographic distribution, habitat, and n

morphological features of S. canadensis of this monotypic genus, i

254 . I
!

i Characteristics found in preliminary studies which suggest that the Santa Maria Basin form is a new

species include the following: thick layer of circular muscle in the proboscis, heavy central area of radial

i pattern of longitudinal muscle, glomerulus poorly developed, appearing as a series of isolated islands surrounding the buccal diverticulum, well-developed chondroid tissue sheath encases the buccal diverticulum

and the sinus organ in the peduncle, longer peduncle, skeletal crura develop in the peduncle, conrtective

tissue and muscle well-developed in the collar coelom, ova about 500 pm but nucleus and nucleolus larger at 200 and 100 gin, respectively, nucIeolus lunate. In addition, the Santa Mafia form was found in the

Pacific, at 565 m, and in a very fine silty sediment with high clay density.

l Keys to Enteropneusts

I I. Key Based on External Characteristics

Figure 10.1

I 1A. Genital wings or enlarged genital lappets absent ............................................................................ 3

l lB. Genital wings or enlarged genital lappets present ........................................................................... 2

2A. Lateral genital wings (A) in the branchial and esophageal region ............................. BaIanoglossus

I 2B. Four enlarged genital lappets (B), not present in esophageal region; lappets present below and above branchial openings ........................................................................................... Stereobalanus

I 3A. Elongate cylindroid proboscis (C); small with large eggs (150 _-n) visible externally .... Saccoglossus

3B. Proboscis not elongate, generally ovate (D); large with small eggs (<100 _rn); proboscis 3 x length,

I 5 x width of Saccoglossus ......................................................................................... Schizocardium

I II. Key Based on Internal Characteristics Figure 10.2

I 1A. Longitudinal muscles of proboscis arranged in definite patterns; thin layer of external circular

muscle .............................................................................................................................................. 2

t lB. Longitu externaldicircular nal musmuscle cle of proboscis not in patterns but is homogeneously arranged; thick layer of (E) ....................................................................................... Schizocardium

/I

I! 2A. Muscle arranged inradial pattern .................................................................................................... 3 m

2B. Muscle not arranged in radial pattern; concentric rings common (F) .......................... Saccoglossus

I 3A. Many (>100) thin radial bands closely packed and concentrated peripherally; large central proboscis

coelom (G) .................................................................................................................. BaIanoglossus

I 3B. Less than 50 bands, narrow peripherally with much open space between bands, thicker and more concentrated near the center where the proboscis coelom is small (H) ..................... StereobaIanus

I

i 255
Figure

10.1. Plate showing anterior ends of four different genera of enteropneusts. Letters explained inkey.1

!
I

i

I

i

Ii E F

Schizocardium Saccoglossus

!

i

Stereobalanus Balanoglossus

!

Figure 10,2. Plate showing cross sections of the proboscis in four different genera of enteropneusts. Letters

I explained in Key.

I

i

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i

I

I 257
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Literature Cited i

Bateson, W. 1885. The later stages in the development of Balanoglossus kowalevskii with suggestion as to n

the affinities of the Enteropneusta. Quarterly Journal of Microscopical Science 25 (Supplement):81- il

122.

Corgiat, J.M., F.C. Dobbs, M.W. Burger, and P.J. Scheuer. 1993. Organohalogen constituents of the acorn $

worm Ptychodera bahamensis. Comparative Biochemistry and Physiology 106B:83-86.

DeUe Chiaje, S. 1829. Memorie sulla storia e notomia degli Animali senza. Vertebre del Regno di Napoli. i

Volume 4:117-120, 141,154. 4;

Eschscholtz, F. 1825. Bericht Ciberdie Zoologische Ausbeute der Reise von Kronstadt bis St. Peter und iiil

Paul. Oken's Isis for 1825, Heft 6:734-747. |

Galigher, A.E. and E.N. Kozloff. 1971. Essentials of Practical Microtechnique. Second Edition. Philadelphia.

Lea and Febiger. pp. i-viii + 1-531. n

U

Gegenbaur, C. 1870. Grundzuge der Vergleichenden Anatomie. Second Edition, pp. 158.

Girard, C. 1853. New nemerteans from the coast of the Carolinas. Proceedings of the Academy of Natural i

Sciences, Philadelphia, 6:365-367. II

Hadfield, M.G. 1975. Hemichordata. In: Giese, A.C. and J.S. Pearse (eds.), Reproduction of Marine i

Invertebrates. Entoprocts and Lesser Coelomates. Vol. 2. Academic Press, New York. pp. 185-240. |

Higa, T., T. Fujiyama, and P.J. Scheuer. 1980. Halogenated phenol and indole constituents of acorn worms.

Comparative Biochemistry and Physiology 65B:525-530. i

Nil

Hyman, L.H. 1959. The Invertebrates: Smaller Coetomate Groups, Vol. 5. McGraw-Hill, New York. Pp.

72-207. II

King, G.M. t986. Inhibition of microbial activity in marine sediments by a bromophenol from a hemichordate. I

Nature 323:257-259.

Kowalevsky, A. 1866. Anatomic des Balanoglossus Delle Chiaje. Memoirs Academie Imperial Science St. I

Petersbourg, Section 7, 10:7-25.

Norrevang, A. 1965 Fine structure of nervous layer, basement membrane, and muscles of the proboscis in

Harrimania kupfferi 0Enteropneusta). Videnskabelige Meddelelser fra den naturhistoriske Forening, U

Copenhagen 128:325-337.

Saita, A., L Castellani, and S. Tripepi. 1978. The integument of Glossobalanus minutus Kowalevsky i

(Enteropneusta Ptycoderidae (sic)). Ultrastructural Analysis, Monitore Zoologica Italien (new series)

12:155-t79.

II

Scheltema, R.S. 1987. Adaptations of Ptychodera flava and Ptychodera bahamensis t

(Enteropneusta:Ptychoderidae) for dispersal and colonization. Biological Oceanography 5:85-97.

Spengel, J.W. 1893. Die Enteropneusten des Golfes Neapel. Fauna und Flora des Golfes Neapel Monograph 1

U

18:1-757.

Spengel, J.W. 1901. Die Benennung der Enteropneusten-Gattungen. Zoologica Jahrbuchen Abteilung System l

15:209-218. 11

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I Spengel, J.W. 1907. Studien tiber die Enteropneusten der Siboga Expedition. Siboga Expedition Monograph

26:1-126.

' I Thistle, D. 1980. The response of a harpacticoid copepod community to a small-scale natural disturbance. Journal of Marine Research 38:381-395.

I Welsch, U. and V. Storch. 1970. The true structure of the stomochord of the enteropneusts Harrimania kupfferi and Ptychodera tiara. Zeitschrift ftir Zellforschungen Mikroskopic Anatomie 107:234-

239.

I Woodwick, K.H. 1955. A list of Enteropneusta identified from samples of the benthos of San Pedro Basin, California. In O. Hare'nan, Quantitative survey of the benthos ofSan Pedro Basin, Southern California.

Allan Hancock Pacific Expeditions 19:166-167.

i Woodwick, K.H. and T. Sensenbangh. 1985.Saxipendium coronatum, new genus, new species (Hemichordata:

Enteropneusta): The unusual spaghetti worms of the Gahipagos Rift hydrothermal vents. Proceedings

!' i of the Biological Society of Washington 98:351-365.

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II

II 11. PHYLUMCHORDATA"

I SUBPHYLUM UROCHORDATA, CLASS ASCIDIACEA

by

I Gretchen Lambert 1

I Introduction

l affmitiesThe are ascidians, expressed or primarily tunicates, in thearelaarval groustage. p of exclusi The early vely development marine invertebr of theateeggchoshares rdatesmany whosesimilarities chordate

with vertebrate early development. The larva is a tiny (< 1mm long) short-lived, non-feeding tadpole (Fig.

i 11.1A) with a notochord, dorsal tubular nerve cord, and the non-functioning rudiments of pharyngeal gill

slits that become greatly proliferated in the adult branchial sac that is used for filter feeding. The notochord

is lost when the swimming tadpole settles and undergoes metamorphosis into the sessile saclike adult form;

i all that remains of the nerve cord in the adult is a dorsal ganglion. Another chordate feature is the endostyle, considered to be an evolutionary forerunner of the thyroid gland, which produces a thyroxin-like hormone.

Detailed overviews of the ascidians are well presented by Van Name (1945), Abbott (1975, 1980), and

i Monniot et al. (1991).

i Morphology and General Biology

Adult ascidians are sessile f'dterfeeders (Figs. 11.1B, 11.2). Each animal has two siphons, incurrent

I and oral excurrent, for the passage of water through the pharynx or branchial sac. At the base of the incurrent siphon are a number of oral tentacles that sort the incoming food particles and may reject large ones.

The oral siphon opens into the branchial sac which is pierced by a number of rows of ciliated gill slits. A

i mucous sheet is formed by the ventral endostyle during feeding and spreads across the inner surface of the • branchial sac, coveting the gill slits and catching filtered particles down to the size of bacteria. When the

mucous sheet reaches the dorsal side of the sac, a longitudinal series of curved dorsal languets form the

mucus into a string which moves by ciliary action into the esophagus and stomach. Fecal pellets accumulate in the rectum and are expelled periodically through the excurrent siphon.

The ascidian body is covered by a mantle or body wall which contains longitudinal and circular

I mcellulose-like uscle bands.material; The mantle ascidian is loos sely are often attached referred to a totouasghtunicates outer covbecause ering, cofalletheir d thetunic. tunic, composed of a

All ascidians are hermaphroditic, possessing both ovaries and testes. Although a few species routinely

I self fertilization -fertilize may their beowninternal eggs, mwithost the species larvaedobeing not. The incubated eggs anund tisperm l theyarereachusually the swimming spawned inttadpole o the sea, stageor

(Fig. 11.1A). A few species breed all year, but most have a definite breeding season, usually in the summer.

I Asc several idianscentimeters may be either in length, solitarythough or coloni the smallest al. Solitary, species or siaremple, interstitial forms reproduce and just asexu fewammlly; intheylength. may Some reach

l

. t Department of Biological Science, California State University, Fullerton, California 92634-9480.

i 261
!

solitary forms are stalked, although the majority of solitary species attach directly to the substrate by their i

posterior end or left side. Colonial forms are composed of many small zooids a few mm in length embedded

in a common tunic which in some species, may spread out in a sheet to cover considerable space. Each I

colony arises from the settlement of a single tadpole which then buds asexually to form a clone of zooids. I

There are two types of colonial species: (1) social ascidians in which the zooids are attached to one another

only by their base, with the rest of the body independent and covered by its own tunic (Fig. 11.6), and (2) m

compound ascidians in which the zooids are completely contained within the common tunic with only one U

or both siphonal openings visible on the tunic surface.

Ascidians can be found in almost every marine habitat: interstitially in mud and sand, attached to

rocks on exposed coasts, epibiotic on molluscs or other ascidians, and as common fouling organisms on II

floats, pilings and boat hulls. They occur from the intertidal to abyssal depths, and in the wannest and

coldest waters of the planet. In the tropics some compound ascidians contain symbiotic algae, primarily the _i

unicellular Prochloron (Lewin and Chang 1989). A newly discovered family of abyssal ascidians do not I

filter-feed; their pharynx is modified to catch large prey such as crustaceans. Another recently discovered

group of benthic deep-sea tunicates (not covered in this chapter) is so different from the Ascidiacea that they i

have been placed in their own class, the Sorberacea (Monniot et al. 1975). Undoubtedly there are still many U

undescribed species yet to be collected, especially from deeper waters; this Atlas contains three new ascidian

species, described separately (Lambert, 1993).

Commensals and parasites are common in ascidians. The clam Mytilimeria is never found living II

freely but only embedded in the tunic of certain compound or solitary ascidians. Certain species of burrowing

amphipods are sheltered in the tunic. Free living amphipods are found in the branchial sac of most solitary

species, apparently feeding on the nutritiously rich mucous sheet. Ascidians are frequently parasitized by II

specially adapted copepods called ascidicoles (Ooishi, 1991). Although predation on ascidians does occur,

it is not common because the ascidian tunic appears to be a deterrent. The tunic is often highly acidic, and in III

some species contains large numbers of calcareous spicules (Lambert, 1979). A number of solitary forms II

also have spicules in their body tissues (Lambert, 1992). Nevertheless, flatworms, fish, various echinoderms,

and molluscs have been observed feeding on both solitary and colonial forms.

The I! ascidians collected as part of this study come mostly from collections made along hard bottom

transects that were surveyed with a submersible. Specimens were taken as part of these surveys and were

collected with the aid of manipulator arms on the submersible. The coordinates of each transect are provided am

in the Appendix (Table A.3) at the end of this volume. A list of ascidian species found at each station is II

provided in Table 11.1.

!

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I

I

I

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262 i
I

I

Figure 11.1. A, ascidian tadpole larva greatly enlarged (from Koz|off, 1990); B, a typical compound ascidian

zooid with postabdomen, x16. (from Abbott, 1975). !

I 263
!

!

y

_Y.'i/'] ,oog,..se_.ol ,

__! transversevessels

/

tentacles -- oral siphon (cut open) I

pe_iphoryngeol _ _

groove dorsol tubercle,

beoring operture I

endostyle---___//[_ _°f neurol gland

large folds , . . . . . . " " dorsal _1 in ' ..' .. , • .-' " ' ' lamina

phorynQeol ,, • _ : . :. !..; _ with I!

wall ; i " _. • : _ _ Iong_ets

, .. •, .-.- . • _ ., esophogus I

B

se m olrioE siphon _/ \

,riQht _.t-common ,octspermotic/_ t_- '\\ \ \ _/ _ ---- rectum ao.s |

rightgo_ad / __ '._'_1 le_tgonod I

t ,

v __ ,ton. II

esophogus / • stOmoch

!

Figure 11.2. Pyura haustor, an example of a typical solitary ascidian (approximately life size) showing major

body parts. A, method of dissection; B, animal spread open to show inner surfaces of oral siphon m

and branchial sac (pharynx); C, same view as in B but with branchial sac removed, oral tentacles not II

shown; D, a small part of branchial sac, enlarged. (from Abbott, 1975).

I
!

I Taxonomy

I Aplousobranchia The class Ascidiacea is divided into three orders based on the structure of the branchial sac. The axe all compound species with small zooids possessing a simple branchial sac with no

transverse and longitudinal internal blood vessels. The Phlebobranchia includes both social and solitary

I fbesides orms; thecontaining pharynx cvessels, ontains isinternal greatly vessels. expandedIn theintoStollongitudinal idobranchia folds. the inne Identifi r surfcations ace of the are branchial based on sacthe,

presence or absence, and number, of branchial folds, and number of longitudinal vessels, as well as the

l relevant structure differences of the stomachare, listed numberin and the arrangement Key to Species. of theFigs. ovaries l l.IBandandtestes, 11.2 andshowa hothest ofgeneral other details. anatomy Theof

compound and solitary ascidians.

I Table 11.1. Assemblages of Ascidian Species on Hard Substrate Stations in the Santa Maria Basin and

Western Santa Barbara Channel. Listed by Hardbottom Transect Number (See Appendix

i A.3).

i Transect No. Species assemblages

1 Chelyosoma columbianum, Styela milleri, Didemnum carnutentum var. lacteolum, Styela sp.

I

2 Boltenia polyplacoderma, Chetyosoma cotumbianum

I 4 Chelyosoma columbianum, Halocynthia igaboja, Corella sp., Pyura haustor

I 6 Aplidium var. lacteolum, californStyela icum,

tesseris Pyura ,

spHalocynthia ., Chelyosomaigaboja, columbianum Trididemnum , Styela opacu gibbsimi, Didemnum carnulentum

l Aplidium sp., Halocynthia igaboja, sp., Pyura haustor, polyplacoderma, Pyura sp.,

13 Ascidia Boltenia

Chetyosoma columbianum, Styela milleri, Eudistoma carolinense (?)

I 14 Bolteniapolyplacoderma, Pyura haustor, CheIyosoma columbianum, StyeIa milleri, MotguIa regufaris

i 16 Aplidium polyplacoderma, spauldingPyura i, Chelyosoraa haustor columbianum, Ascidia sp., Halocynthia igaboja,Boltenia

I 20 Chelyosoma columbianum, Pyura haustor, Halocynthia igaboja, Styela milleri

21 Chelyosoma columbianum, Halocynthia igaboja, Distaplia occidentalis, Styela wsseris, Eudistoma

I carolinense (?)

i 25 Chelyosoma columbianum, Styela milleri, Corella sp.

27 Aplidium californicum, Halocynthia igaboja, Chelyosoma columbianum, Pyura haustor, Euherdmania

I claviformis

i 265
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Methods of Preservation and Dissection i

Ascidians should be relaxed and narcotized before fixation to best observe the anatomical details i

It necessary for taxonomic identification. However, this is often a problem because these organisms are

extremely difficult to relax. A common method involves leaving the animals in a bowl of ambient temperature

sea water until they are fully expanded. Then a few drops of a concentrated solution of menthol in 95 % n

ethanol is added and gently stirred in; a solution of 7% magnesium chloride in diluted sea water can also be g

gradually added to the bowl. Alternatively, the chemical MS-222 will quickly anesthetize most compound

species and solitary phlebobranchs when a few milligrams are sprinkled on the surface of the dish in which i

the ascidians have already relaxed; it does not seem to work well on stolidobranchs. U

When the animals do not contract their siphons in response to prodding (after 4-12 hours or more),

they are then transferred to a solution of 10%buffered sea water formalin. One animal should be transferred i

f'lrst; if it contracts in the fixative, then add about two drops of 10% formalin to the bowl of sea water and i

wait until the animals are dead before transferring them to the fixative. They can be stored in the formalin,

or if they have calcareous spicules, they should be transferred after a few hours to permanent storage in 70% I

ethanol. I

In order to observe the zooids in a compound species, thin slices of the colony can be made with a m

blade, or if the tunic is not too tough, it can be torn with forceps to release the zooids. Certain details •

razor

of the branchial sac may require cutting this tiny structure open with small iridectomy scissors or a scalpel, w 

Some zooids are opaque and must be cleared in glycerin for several hours before observation. Sometimes m

staining the zooids in a very dilute solution of methylene blue or toluidine blue will enhance the outlines of i

difficult-to-see structures such as the branchial stigmata. w

Solitary ascidians are opened by cutting posteriorly from the oral siphon along the ventral side m

through the tunic, body wall and branchial sac to the posterior end of the animal (Fig. 11.2). The animal is i

then laid out in a dish of water with its two halves still connected dorsally.

Key to Genera and Species of Ascidians n

This key includes those species collected to date from the Santa MariaBasin, and is not meant to be I

used as a general key for ascidians. For more comprehensive keys to American Pacific coast species the

reader is directed to Van Name (1945), Abbott (1975), or Lambert et al. (1987). I

m

IA. Solitary; not attached to other individuals by a common tunic or by stolons ................................ 8 h

lB. Social (zooids connected at their base by stolons or a sheet of tunic, their bodies separate and !

contained within their own tunic) or compound (zooids contained within a common tunic) ........ 2

2A. Social, zooids attached by stolons at their base; body very slender, about 2 mm wide and 20-35 mm I

in length .................................................................................................... Euherdmania claviformis

2B. Compound, zooids embedded in a common tunic; zooids never more than 10 mm in length ....... 3 I

3A. Tunic with stellate calcareous spicules ...................................................... Didemnidae ............... 4 _L

3B. Tunic without calcareous spicules ................................................................................................... 5 II

266 |
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I 4A. Zooids with 3 rows of branchial stigmata ...................................................... Trididemnum opacum 4B. Zooids with 4 rows of branchial stigmata ........................... Didemnum carnuIentum var. lacteolum

I 5A. Zooids with 3 rows of stigmata ................................................................ Eudistoma carolinense(?)

5B. Zooids with more than 3 rows of stigmata ...................................................................................... 6

i

6A. Zooids with 4 rows of stigmata ...................................................................... Distaplia occidentalis

6B. Zooids with 8 or more rows of stigmata ........................................................................................ 7

7A. Zooids with 8-12 rows of stigmata; stomach with 12-23 longitudinal folds .......Aplidium californicum

I 7B. Zooids with 18 rows of stigmata; stomach with 3 or 4 longitudinal folds .............. Aplidium spauldingi

I 8A. Branchial sac without longitudinal folds ......................................................................................... 9

8B. Branchial sac with 4 or more longitudinal folds on each side ...................................................... 11

I 9A. Branchial stigmata straight ............................................................................................... Ascidia sp.

9B. Branchial stigmata coiled ............................................................................................................. 10

I

10A. Oral and atrial apertures surrounded by horny disk composed of variable number of plates arranged

i 10B. inTunic definite relatively patternsmooth, ................................................................................... without plates ........................................................................... Chelyosoma columbianum CorelIa sp.

I 11A. Branchial sac with 4 longitudinal folds on each side; oral tentacles not branched

.................................................................................................................... Styelidae .................. 16

I 11B. Branchial sac with more than 4 longitudinal folds per side; oral tentacles branched .................. 12

12A. Stigmata straight, both oral and atrial apertures 4-lobed; without renal sac (="kidney") ................

I .................................................................................................................... Pyuridae ................... 14

12B. Stigmata coiled, oral aperture 6-lobed, atrial aperture 4-lobed, with renal sac (="kidney") on right

I side of body below gonad ........................................................................... Molgulidae ............... 13

13A. Ovaries very long, sinuous tubes; animal with only short rootlet hairs, attached primarily by posterior

I region of tunic ...................................................................................................... Molgula regularis

13B. Ovaries compact, oval; tunic covered with short hairs ptus a very long branched rootlike process

that projects from posterior end; siphons widely separated, the atrial anteriormost and the branchial

I about halfway back Molgula napiformis

I 14A. Stigmata tunic ratherwiththinlongbut diameter composedtransverse of flat irregular to bodypolygo axis,neach al plates; crossedanimal by narrow flattenedlongitudinal anteroposteriorly vessel;

................................................................................................................... Boltenia polyplacoderma

I 14B. Stigmata with long axis arranged anteroposteriorly in branchial sac 15

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15A. Tunic thick, ridged, without obvious spines; siphons tong, reddish; gonads one on each side ........ ._

.................................................................................................................................... Pyura haustor

15B. Tunic densely covered with long fleshy thorn-like spines, branched at their tips and with recurved I

spinelets along the sides; spines may obscure short siphons; several gonads on each side ..............

I

.......................................................................................................................... Halocynthia igaboja i

1

16A. One ovary on each side of animal, attached to body wall ........................................... Styela miUeri

I6B. Two ovaries on each side of animal, attached to body wall .......................................................... 17 I

17A. Tunic irregularly thickened into flattened polygonal tessellations with groove separating each one; I

5-6 longitudinal vessels on each branchial fold ........................................................ Styela tesseris I

17B. Tunic with numerous longitudinal or oblique wrinkles, may also be warty or bumpy but not tessellated;

5-15 longitudinal vessels on each branchial fold ......................................................... Styela gibbsii I

Additional species not in the key, but which are known from the area are listed below:

Family Polycitoridae I

Cystodytes lobatus (Ritter, 1900); 79-203 m

Family Ascidiidae I

Ascidia clementea Ritter 1907; 1196-2012 m Ill

Benthascidia michaelseni Ritter 1907; 3990 m I

Ciona mollis Ritter 1907; 2012 m

Family Pyuridae I

Bathyperafeminalba Young and Vazquez, I995; 60-100 m off Santa Catalina Island I

Bathypera ovoida (Ritter, 1907); 1829 m !

Boltenia villosa (Stimpson, 1864); 38-88 m /

CuIeolus pyramidalis Ritter, 1907; 4131 m I

1

Pyura millari Rodrigues, 1966; 91 m

Pyura mirabilis (von Drasche, 1884); 60 m I

With two exceptions, allthe above species were collected by Ritter in 1904 off the coast of California I

and described in his 1907 paper. See Young and Vazquez (1995) concerning Bathyperafeminalba. Pyura 1

milIari (?) was collected by Dr. Richard Emlet and myself by dredging at 91 m off Santa Catalina Island, II

April 1992. The species was abundant and was provisionally identified by Dr. Claude Monniot, Museum

Nationale d'Histoire Naturelle, Pads. This species has previously been recorded only from the Atlantic, !

near Sao Paulo, Brazil, its type locality, at 140 m. I

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I Description of Species

I Order Aplousobranchia

Family Didemnidae

I Didemnum carnulentum var. lacteolum Ritter and Forsyth, 1917

i Figure 11.3 A-B

Didemnum carnulentum Ritter and Forsyth, 1917:470, pl. 39, fig. 11, pl. 44, figs. 57-59.--Johnson and

i DidemnumSnook, carnulentum 1927:596,var. fig.lacteoIum 696.--Abbott, Riter 1975:649.-- and Forsyth,Abbott 1917:471, and Newberry, pl. 40, fig. I980:187, 23, pl. 44, fig. fig. 12.11. 60.--Van

Name, 1945:94-95.

I Material Examined. California, off Point Arguello, Sta. BRA-6, 61 m, sample 2 (3 pieces,

SBMNH).--Westem Santa Barbara Channel, Stat BRA-1, sample 4, 69-73.5 m (1 colony piece, USNM).

I pure whiteDesdue cription. to numero FromusRitter calcareoandusFospicules rsyth (1in917). test,C0.015 olony-0.065 encrusting, mm in1diameter mm or less(staintedthiincorrec ckness; tlcoy loasr

0.15-0.065 mm). Spicules in both upper and lower strata of colony. Bladder cells not conspicuous but

I presen mm long, t around with close 4 rows ly packed of stigmata. zooids,Zooids which similar extend intotoDboth . carnulentum upper and lbut owersmaller, strata. Zooids with atrial onlyopening about 1

proportionately much larger, extending over half length of branchial sac and sometimes even larger than

i dep sac.icCoil ted byof vas Ritterdeferens and Fowith rsythseven (1917). turns. TestisRipealmost ova gigantic, sphericallarger whenproportionately fully ripe and half thanasinlarge D. carnulentum as branchial.

Breeding season in June.

I species. RInemar the USNM ks. Thiscolony, varietyonediffers zooidbyhadenough a testischaracters cleft intofrom2 halves D. carnulentu and the vasm deferens that it mawith y be 9-10 a sepcoils, arate

while others had fewer turns and an undivided testis. In the SBMNH colonies all the zooids had a testis with

I 2thatlarge"neither lobes isanthed thenumber vas defere of parts ns with into 7which coils. theVanteName stis is cleft (1945,always p. 80 constant under thewithin genusthedescriptio same species." n) states

Distribution. Southern California, on under sides of rocks and in kelp holdfasts, from the zero tide

I level to a few meters; Santa Barbara Channel, 69-73.5 m; Santa Maria Basin, 61 m.

i TrididemnumFigure opacum 11.3 C-D(Ritter, 1907)

I Didemnum opacum Ritter, 1907:42, pl. 3, figs. 40, 41.

Trididemnum opacum Van Name, 1945:106-107, text fig. 49 (synonymy).--Abbott, 1975:649.--Abbott

and Newberry, 1980:I87-188, fig. 12.12.--Lambert et al., 1987:473.

I Material Examined. California, off Point Arguello, Sta. BRA-6, sample 2, 61 m (several pieces of

probable single colony, USNM).

I Description. From Van Name Colonies reach 15 and

(1945). encrusting; may cm or more across

10-15 mm thick, opaque pale flesh color or white, with the stellate calcium carbonate spicules showing

through as dense white specks. Test tough and firm. Spicules abundant throughout the test but especially in

I superficial region, 0.03-0.035 mm in diameter, with spines than half spicule diameter, their points

greater

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Figure 11.3. Didemnum carnulentum var. lacteolum. A, zooid, x50 (a.o.: atrial opening; m.b.: muscle band; o.:

ovary;s.:stomach;t.:testis; v.d.: vas deferens); B, spicule,x650 (from Ritter andForsyth, 1917). CD, Trididemnum opacum. C, zooid, X30; D, spicules, x900. (from Van Name, 1945). •

|

long but often truncated at tips. Zooids 2 mm or less in length in contracted specimens. Both branchial and m

atrial tubes well developed, atrial far back on thorax, slightly funnel-shaped and commonly directed obliquely I

backward. Three rows of stigmata. Stomach large and globular. Testis round and undivided; vas deferens 1

with about 6 coils on surface but may be fewer in young zooids. I

Remarks. The colony in this collection has zooids that are brownish, visible through the tunic

which is not as opaque as Van Name (1945) indicates, but he does state that the zooids become a dull

brownish or grayish white after preservation. The vas deferens has eight coils. The atrial siphon is not quite

as far back as stated by Ritter (1907), but it is tubular and flared like a funnel as stated by Van Name. I

Distribution. Common in central California intertidally on overhanging rocks, and dredged offSan I

Nicolas Island, California, from 61 m. Also recorded from southern California; Santa Maria Basin, 61 m.

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I Family Polycitoridae

I Distaplia oecidentalis Bancroft, 1899

Figure 11.4 A

I Distaplia occidentalis Bancroft, 1899:59-112, pls. 1-6.--Ritter and Forsyth, 1917:464-467, pl. 45, figs. 64,

65.--Van Name, 1945:149-150 (synonymy).--Abbott, 1975:650.--Abbott and Newberry,

1980:194-195, fig. 12.21.--Lambert et al., 1987:473.

Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA-21, sample 2, 87

m (1 small colony, USNM).

I Description. From Van Name (1945). Colony form variable; may be fiat and encmsting or, more

usually, pedunculate or mushroom shaped with heads up to 1 cm or more across. Several systems in head,

zooids closely arranged around a large cylindrical common atrial opening extending above test surface as

I pipe. highly variable, from dirty white to yellow, light green, lavender,

delicate-walled short Color of heads

brown or reddish, due to pigment cells in test; peduncle often grey. Test with thin, very tough outer layer.

Zooids 2-3 mm long due to contraction, but if expanded, may be much longer. Body with an expanded

I thorax; by very narrow rows stigmata present, each row crossed at its

abdomen connected neck. Four of

middle by slender parastiglnatic transverse vessel; 12-14 stigmata per row on each side. Stomach wall

smooth externally but with irregular internal ridges giving it a reticulated appearance.

I Ovary posterior to intestinal loop and partially to testis; incubatory pouch rarely containing up to 4

embryos, extending into test from posterior end of thorax by long slender, easily breakable neck.

I flattenedRupp emarekss.urThis face,species the incubator is msty easily pouch,ideand ntifiedthe bypartheastigma pedutnicultarante ssvhape erse ofvetsheselscolonies in the with branchial teir

basket. The species occurs in quiet waters on marina floats, and may be very abundant during summer

I months iDistribution. n Washington. Puget Sound, San Juan Archipelago, on rocks usually in shallow water; southern

California (San Diego); Santa Mafia Basin, 87 m; ? British Columbia.

I Eudistoma carolinense (?) Van Name, 1945

I Figure 11.4 B-C

Eudistoma carolinense V an Name 1945:123-124, fig. 58.--Plough 1978:59, 61, fig. 26e.

i Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA-13, 86-98 m,

sample 4 (1 small colony, USNM).

I Description. From Van Name (1945). Colony composed of irregularly tapering clavate heads over a narrowed neck region, arising from an expanded, encrusting base; 20-30 mm in height, 3-10 mm wide at

top;specimens may be flattened laterally, usually curved andcrooked. The heads not all arising from expanded

I bwith ase of the colony, but in many cases from lower part of another head or neck. Test densely impregnated sand. Contracted zooids only about 2 mm long, with anterior ends withdrawn 4-5 mm below colony

surface. Each separate head in colony narrow, small, containing from 1-6 zooids, each with its oral and

I aptrial er rowsiphons on eachopening side ofsepar branchial ately tosac.suOrrfaalce,tentacl as tyepical s nume forrougens,us.of Thr 2 sizeeesrow. Stomach s of stigmata, rounded, about smooth 10 stigamta walled;

with gland surrounding intestine consisting of very few tubules swollen at ends. Testes pyriform, 6-7.

I Strong longitudinal muscle bands extend from thorax far down sides of abdomen.

I 271
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p._[. I

V°d

C |

A ,i

Figure 11.4. Distaplia occidentalis, A, zooid, ×42 (b.s. - branchial siphon, e - esophagus, end. - endostyle, i.p. - m

incubatory pouch, o. - ovary, ov.d. - oviduct, p.gl. - pyloric gland, s. - stomach, t. - testis, v.cL - vas i

deferens) (from Ritter and Forsyth, 1917); Eudistoma carolinense, B, colony, x1.3 (fxom Van

Name, 1945); C, zooid, left and right sides of body, x3.2 (from Plough, 1978).

!

m

Remarks. The two small colonies in this collection resemble inalmost every respect E. carolinense I[

Van Name, 1945, but that species has been recorded only from the Atlantic, in warm shallow water from the

m

southeast U. S. coast. Thus, it may not be appropriate to establish a new species name for these colonies

when they agree so closely with Van Name's description. The colonies in this collection are about 10 mm i

tall. The distal and basal portions of the voucher specimen are expanded and about 5 mm wide, the middle

w

portion a constricted neck; the zooids are 5-6 mm long. The zooids are slightly longer than indicated by Van

Name (1945) and Plough (1978), but the species in this genus are all capable of extreme contraction, so

lengths of preserved specimens can be very variable. I

Distribution. Coastal SE United States, down to 30 m; California, Santa Mafia Basin. 86-98 m. !

I

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N Family Synoicidae

I Aplidium eah'fornicum (Ritter and Forsyth, 1917)

Figure 11.5 A

n Amaroucium californicum Ritter and Forsyth, 1917:483, pI. 46, fig. 72.--Van Name, 1945:47-48, fig. 12.

Aplidium (=Amaroucium) californicum Abbott, 1975:648, 650.--Abbott and Newberry, 1980:182-183, fig.

i 12.1.--Lambert et al., 1987:473, fig. 23.3.

Material Examined, California, off Point Arguello, Sta. BRA-6, sample 2, 61 m (1 small colony,

I USNM).Description. From Van Name (1945). Colonies extremely variable in size, shape and color. Often

flat and cake-like; sometimes attaining large size. Surface usually smooth, without embedded sand grains;

N test gelatinous, somewhat translucent, zooids visible within. Surface of test may be tougher than interior softer regions. Color of test ranges from opalescent white to yellowish, yellowish-brown or reddish brown,

or mottled. Zooids also variable in color, from yellow or orange in some colonies to bright red in others.

Zooids usually 6 mm or less in length. Branchial aperture with 6 lobes, atrial smooth and with long undivided

Ii atrial languet. Eight to 12 rows of stigmata, with about 12 stigmata per row on each side.

Stomach short, rounded, with about 15-20 regular longitudinal folds (the number varies from 12-

I this 23). variable Ovary loincatedlength. in anterior region of post-abdomen; double row of testes occupy most of post-abdomen,

Remarks. The genusAmaroucium was formally synonymized withAplidium by Nishikawa (1991),

I thus making official a long-standing informal lumping of these two genera.Aplidium californicum is probably the most common compound ascidian on the California coast, especially in shallow waters. The great

variability between colonies can make this species difficult to identify, however. The zooids can be removed

I very collection easilyabout from2preserved mm colonies; they may in fact fall out when the colony is dissected. Zooids in this long, bright orange-red, with I0 rows of stigmata. Tokioka (1967) tentatively grouped

this species with the apparently identical A. muln'plicatum from Palau and the Gilbert Islands, though he

I remarkedDistri that the localities are very far apart. bution. Pacific coast of North and South America from British Columbia to Gulf of California,

on rocks and man-made pilings and floats; rare in waters below 85 m. Santa Mafia Basin, 61 m.

!

Aplidium spauldingi (Ritter, 1907)

I Figure 11.5 B

Psammaplidiura spauldingi Ritter 1907:41, pl. 3, fig. 39.

I Amaroucium spauldingi Hartmeyer, 1909-1911:1471 .--Van Name, 1945:53.

Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA- 16, sample 4, 96

I m (1 piece of colony, USNM).

Description. From Ritter (1907). Top of colony smooth, hard from great quantity of sand uniformly

i distributed through test. Numerous small zooids evenly distributed, systems not obvious. Each branchial opening marked on colony surface by small papilla. Color uniform grey due to embedded sand. Zooids

long, slender, nearly straight; thorax about equal in diameter to abdomen and post-abdomen. Thorax about

3 mm long, abdomen about 2 mm long, and post-abdomen several times as long as thorax and abdomen

N combined. Mantle with few slender muscle far down into longitudinal bands, extending post-abdomen.

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A B I

I

Figure 11.5. Aplidium californicum, A, zooid, x22 (fp - fecal pellet, sd - sperm duct, st - stomach) (from Van a

Name, 1945, adapted from Ritter and Forsyth, 1917); Aplidium spauldingi, B, zooid, x8 (al. - atrial •

languet, ov. - ovary, t. - testis) (from Ritter, 1907). U

Branchial opening on prominent 6-lobed siphon, atrial siphon situated well back from end of branchial !

siphon about equal in thickness to thorax. Atrial languet long, and either undivided or bifid at tip. About 18

rows of stigmata. Endostyle large and tortuous. Branchial tentacles about 10, of unequal length. Esophagus •

about equal in length to stomach, this with 3-4 large longitudinal folds or lobes. A distinctly set off enlarged I

section of intestine follows pyloric section. Gonads in post-abdomen, testes a number of spherical lobes.

Remarks. The zooids in this collection are from a very small torn piece of colony, and their postabdomens are entirely or partially missing. Nevertheless, because their morphology agrees in all respects J

with the characters listed by Ritter (1907), they are referred to this species with confidence. These characters

include the unusual placement and bifurcated atrial languet, coupled with the large number of stigmatal

rows, very few longitudinal stomach folds, heavily sand-encrusted test, and location of collection. II

Distribution. Type locality, 61 m, 3.8 miles off the east point of San Nicolas Island, California, in

f'me grey sand; Santa Maria Basin, 96 m. •

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I Euherdmania claviformis (Ritter, 1903)

Figure 11.6

! tIerdmania claviformis Ritter, 1903:239-261, pls. 18, 19.

i Euherdmania claviformis Ritter, 1904:650.--Johnson and Snook, 1927:597, fig. 698.--Van Name, 1945:76-

78, fig. 32 (synonymy).--Abbott, 1975:647.--Abbott and Newberry, 1980:186-187, fig. 12.10.

Material Examined. California, Santa Mafia Basin, north of Point Buchon, sample 1, 116 m (1

i colony, USNM).

Description. From Van Name (1945). Zooids in a group, slender, club-shaped, each with its own

i covering of test for entire length, arising from an interwoven mass of branching stolons. Zooids often thickly encrusted with firmly adhering sand except at or near anterior end; test where free of sand translucent,

slightly greenish in life, and tough. Zooids 2-5 cm in length (occasionally longer), 2-4 mm in greatest

diameter, at the level of thorax. Thorax short in proportion to rest of body, separated from stomach and

I gonads by very long slender neck. Gonads posterior to stomach, but without clearly defined post-abdomen.

About 12 rows of stigmata preseent, with about 40 stigmata per row on each side of thorax. Thirty to 40

simple branchial tentacles and additional smaller ones. The transverse vessels each with membrane projecting

I prominently into branchial cavity, and with dorsal languet present where each vessel crosses median dorsal vessel.

i NumerousStomach small, with 6 unequal longitudinal plications and a longitudinally furrowed typhlosole. small oval testes present; ovary usually small because eggs pass into oviduct when still small and

develop in anterior region of oviduct. Developing embryos form longitudinal series in oviduct, with most

i advancedRebeing marks.closest The tozooids anteriorin the end.colony Larvae examined may be 1-are1.2 15mmmmin body long,length. a tittle shorter than Van Name

(1945) lists; this may be a young colony, although Ritter and Forsyth (1917) state that "zooids are somewhat

I smaller This species in souisthern distinguished than in northern from CIavelina colonies." huntsmani It is not as sand-e by thencrusted zooids being in thesmaller, posteriormore portioslender, n as is usual often.

bent or crooked and sand-encrnsted.

I southernDCalifornia istribution.(common), North American on rocksPacific and among coast fralgal om Sanholdfasts Juan Archipel intertidally ago inandPuge in tshallow Sound subtidal; (rare) to

Santa Mafia Basin, 116 m.

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FiN 11.6. Euherdmania claviformis. A, young colony; B, older colony, ×2; 12,zooid with embryos and larvae

in oviduct, x9; D, cross section of stomach; E, part of branchial sac with dorsal languets and adjacent 1

stigmata, greatly enlarged. (A, C, D, E from Van Name, 1945, after Ritter, 1903; B from Johnson

and Snook, 1927).

I

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I Order Phlebobranchia

i Family Rhodosomatidae

Chelyosoma columbianum Huntsman, 1912

I Figure 11.7 A-C

Chelyosoma columbianum Huntsman, 1912:114, 124; 1912a: 126, pl. 11, figs. 2,3, pl. 16, figs. 3,5,6.--Van

I Name, 1945:208-209, fig. 121 (synonymy).--Lambert et al., 1987:468, fig. 23.9.

Material Examined. California, Santa Mafia Basin, north of Point Buchon, Sta_BRA-27, sample

I Point, 1, 96-126Sta.m, BRA-21, (1); offsample Port San1,Luis, 75-90Sta. m (1); BRA-25, same transect, sample 2,sample 64.5-722 (1); m (3,Sta.SBBRA-20, MNH voucher); sample 2, off90-13-0. Purisima5

m (1); Sta. BRA-16, sample 2, 91.5-123 m (2); same station, sample 4 (4); Sta. BRC-14, sample 1,105-117

I mstation, (6); same samplestati4on(1); , sample Sta. BRC-13, 2 (t); same sample statio1,n,88.5-100.5 sample 3 (2); m (2); Sta.same BRA-station, 13, sample sample1, 92-10 2 (1);0same m (3);station, same

sample 3 (2); off Point Arguello, Sta. BRA-6, sample 2, 54-63 m (5); same station, sample 4 (1); Sta. BRAI 4, sample sample2, 2,110-126 168-237m(4,(2).--Western USNM voucherSanta ); same Barbara station, Channel, sampleSta. 4 (I)BRA-1, .----Off Point sampleConception, 2, 69-73.5Sta. m (2); BRA-2, Sta.

BRC-1, sample 2, 73.5-78 m (1).

I Description. Adapted from Huntsman (1912a). Surface of test smooth, yellowish-brown or occasionally clear and transparent. Branchial and atrial apertures situated in a horny disk up to 15 mm in

largest diameter and composed of a number of plates. Body usually much flattened and depressed, elongated

I in direction of about 45 ° with disk, and attached by broad area on side opposite disk. Margin of disk sharper than in Chelyosoma productum, not raised above level of disk in contracted

individuals. Disk somewhat elliptical, usually broader behind atrial opening. Apertures nearer to right side

I of disk than left. Plates typically with 2 ventral, 2 intermediate on left side, and 12 marginal; sometimes with additional central plate, 1 or several intermediate plates on fight side, and a larger or smaller number of

marginal plates; plates not showing growth lines in this species. Siphonal and marginal muscles as in C.

I productum, but with additional short thick strands crossing lines located at some distance from margin.

Usually with 2 stigmata coiled together forming each small infundibulum, but these sometimes

broken into many short ones. Fifty to 100 oral tentacles, 12-22 languets. Stomach narrow, with diameter

I slightly than that of intestine; stomach wall with folds. Gonads for

greater irregular longitudinal typical

genus: ovary branched, located in intestinal loop; testes extensive, spreading over part of intestine anterior

and ventral to stomach.

i Remarks. Chelyosoma columbianum does not attain the size of C. productum. It reaches maturity

at a smaller size, is more flattened, the surface is smoother, the margin of the disk is not definitely raised,

i and the disk is not symmetrical. It can also be distinguished from C. productum by the presence of numerous short muscle strands, usuaUy visible through the transparent plates, connecting the two central plates as well

as most of the other plates, the presence of two intermediate plates on the left side and differently arranged

marginal plates, and the absence of growth lines on the plates. In Station BRA-25 (SBMNH, voucher), one

I specimen a plate largest specimen no plates but, nonetheless,

has third central and the has intermediate

appears to belong to this species.

Type Locality. Stony and shelly bottoms in 18-37 m off British Columbia (Departure Bay,

I Northumberland Straits, and Burrard Inlet).

Distribution. Washington, near entrance of Strait of Juan de Fuca: Albatross Station 2876, off

I Cape Santa Fl Barbara attery, 108 Channel m, annorth d Station to Point 2866,Buchon 313 m (Ritter, in Santa19Maria 13).--Basin, California, on rocks, numerous 54-237statio m. ns from western

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l_re 11.7. Chelyosoma col_ian_. A, whole animal, x2.4 (redrawn from Huntsman, 1912a); B, plates of

disk surrounding apertures, showing connecting muscles, x3 (from Van Name, 1945, after Huntsman, I

1912a); C, plates of disk fi'om another animal, with a slightly different pattern of plates, x2.2 (redrawn

from Huntsman, 1912a); Molgula regularis. D, left side of body; E, right side; x5. (a - atrial siphon,

b - branchial siphon, k - kidney) (fi'om Van Name, 1945). I

Order Stolidobranchia I

Family Molgulidae

Molgula regula_ Ritter, 1907 I

Figure 11.7 D-E

I

Molgula regularis Fitter, 1907:8, pl. 1, figs. 7,8.--Hnntsman, 1912a:135-136.--Van Name, 1945:415-416,

fig. 305.

m

Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA-14, sample 2,

105-117 m (1 voucher, SBMNH).

Description. From Fitter (1907). Body ellipsoid, very regular in outline, surface entirely covered !

with foraminiferous shells and sand particles clinging to numerous filiform processes of test. No definite

area of attachment. Test thin and leathery, semi-transparent; no siphons visible externally. Up to 4 cm long m

by 2-3 cm wide. Large muscle bands of uniform length on mantle, terminating abruptly, extending I

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I longitudinally, radiating from apertures; also with a narrow zone of circular muscles around each aperture.

Rest of mantle with meshwork of fine fibers.

I Branchial opening far forward and 6-lobed; atrial opening 4-lobed near middle of body; branchial tentacles about 10, very branched, with smaller, simpler tentacles between larger ones. Dorsal lamina a

plain-edged rather broad membrane. Branchial sac with 6 prominent folds, 5-7 longitudinal vessels on each

I side large,oflittle eachcurved fold andexcept aboutextending equally spaced. around Infundibula infundibula.large, extending full size to edge of folds; stigmata

Stomach on left side at extreme posterior end of animal; nearly twice as long as broad, smoothI walled or with few slight folds, bearing dark hepatic organ proximally. Intestine very long, thin, of uniform diameter throughout; rectal half forming wide semi-circle; anus without lobes. Kidney a large, brown,

regular elongate central part surrounded by larger clear part and located on right side of body, close to

I posteri along dorsal or endto, of and endobending style. Ovaries down inS-shaped, front of one intesotninal eachloopside, onlong, left side thinand , sinuously kidney oncurved right.tubes Posterior extending ends

bending dorsally, ending near atrial aperture. Testes lobed or branched, may be voluminously developed.

I Remarks. The single specimen in this collection is small, measuring 8 mm dorsoventrally, but has

a long slender ovary on each side. There is no indication that this species is viviparous.

Distribution. California: Corona Del Mar, about 10 m; off Point Loma, 122-134 m; Santa Cruz

I Island, 37 m; Santa Maria Basin, 105-117 m, apparently adhering to a rock; usual habitat, soft substrate.

I Molgula napifermis Lambert, 1993

Figures 11.8, 11.9

I Molgula napiformis Lambert, 1993:111-113, figs. 1-2.

Material Examined. California, Santa Mafia Basin, MMS Phase II Program: Off Point San Luis,

I Sta. R-l, Jan 1988, 91 m (3 paratypes, USNM 35608); off Point Sal, Sta. R-8, June 1987, 90

m (paratype,

CASIZ 086764); off Purisima Point, Sta. R-4, May 1988, 92 m (holotype, USNM 20008) and (5 paratypes,

USNM). Sta. R-1 (3); Sta. R-4 (6); Sta. R-8 (1).

I Description. Animals 2-3 mm in size, with length and width about equal. Tunic clear, transparent,

covered with short hairs to which numerous sand grains firmly adhere. An extremely long branched rootlike

posterior process with long hairs projecting from posterior end of tunic; additional long hairs present or

I process. One specimen with process or stolon about 4 times longer than animal and

absent around main

greatly branched, gradually tapering to very fine hair at apex.

i Siphons short, widely separated; with branchial siphon more noticeable on tunic surface than atrial siphon, produced into 6 pointed processes; atrial siphon square-shaped. All 12 specimens examined with

identical location of siphons; atrial siphon anteriormost, at opposite end of animal from stolon; branchial

i siphon about halfway back, opening laterally. Dorsal tubercle with narrow oval opening. Largest specimen (3 mm in diameter) with 7 branchial folds on left side of branchial sac, and 6 on right side; some folds not

distinct in smallest specimens; with 3 longitudinal vessels per fold. Infundibula large, with only a single row

between each rudimentary branchial fold. All specimens examined more or less immature, precluding

I determination of adult number of infundibular spirals. Dorsal lamina smooth membrane. Largest specimen

with 8 large branchial tentacles, each long, slender, variably branched; additional small tentacles of various

sizes occur between large ones, these difficult to count.

i Mantle with widely spaced longitudinal muscles radiating fi'om siphons over body, overlapping

slightly; circular muscles well developed only on siphons. Intestine and lobed stomach located on left side

i of eachbody. ovary; Theleft kidney gonadnotsecondary developed.inGonads intestinalovalloop. in outline, with branched or unbranched testes surrounding

i 279
I

a I

__ '

_=/_'-_ B m

, I

!

!

!

Figure 11.8. Molgula napiformis. A-B, whole animals, x13 (a - atrial siphon, b - branchial siphon); C, dorsal

tubercle; D, three branchial tentacles; E, detail of branchial sac. C-E greatly enlarged: (fi-omLambert,

I993). I

|

Remarks. This species resembles Molgula pugetiensis in many of its characteristics but differs, (1) w

in possessing a very long' branched tunic stolon and secondary processes in addition to the numerous short Ilia

tunic hairs, (2) in the siphons being farther apart and shorter, and (3) in the body being attached ventrally 1

which results in the endostyle appearing to be posterior. lJ

Distribution. California, Santa Maria Basin, soft substrate, 90-92m. ==

|

I

280 I
I

I

II d

I

I

I

A o B o

I

I Figure 11.9. Molgula branchialnapiformis. A, right side of animal removed fromtunic; B,left side. (a - atrial siphon, b - siphon, d - dorsal ganglion, e - endostyle,l - liver, o - ovary, r - rectum,t - testis), x32:

(from Lambert, 1993).

I

I Family Pyuridae

Boitenia polyplacoderma Lambert, 1993

I Figure 11.10

I Boltenia polyplacoderma Lambert, 1993:113-116, figs. 3-4.

Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA-16, sample 1, 98

i m (2paratypes, USNM 20010); Sta. BRA- 14,, sample 2, 104 m (paratype, CASIZ 086766); Sta. BRA- 13, sample 4, 98 m (2 paratyl_S, CASIZ 086765).--Western Santa Barbara Channel, Sta. BRA-2, sample 1,

116 m (holotype, SBMNH 35607).

I polygonalDefiat-topped scription. Body plates,veryachflawith ttenednumerous anteroposteriorly. closely spaced Tunicannulations. thin, composed Platesofstraight-sided, thickened, irregwithular,3-

8 sides of unequal lengths. Each polygon set off from others by narrow groove in tunic; plates abutting

I cfibrous losely, material. especiallyApertures in contracted 4-sided, specime formedns. byInner 4 main surface triangular of tunicplates; coveredwithbysmaller continuplates ous thinforming sheet between of white

each of these at base of each aperture. Mantle muscles very regular, robust, with longitudinally radiating

i ones deeper, Dorsaland circular ones around each aperture more superficial, creating beautiful regular meshwork. tubercle a simple oval tube, opening on right side between peripharyngeal bands. With

about 18 branched oral tentacles, mostly of 2 alternating sizes; larger tentacles alternately branched, with

i branches between branches. containing Tentacles tiny protruberances; possibly more additional branchedveryinsmall largerprojections animals. occurring ontrunk of main tentacles

I

I 281
I

a I

b

c l

I

I

i

• iWIIill '

!

I

!

I

E W |

I

Fi N 11.10. Boltenia polyplacoderma. A, anterior surface of tunic, x7.2; B, detail of brancl_al sac; C, dorsal

tubercle; D, branchial tentacle (B-D greatly enlarged); E, anterior surface of mantle showing the I

siphons and musculature; F, posterior view of body removed from tunic; E-F x9.7.(a - atrial siphon,

b - branchial siphon, e - endostyle, 1 - liver, lv - longitudinal vessels, o - ovary, od - oviduct, s -

stigmata, st - stomach, r - rectum, t - testis): (from Lambert, 1993). I

282 I
!

I Branchial sac with 6 branchial folds on each side, of unequal size; second (counting from dorsal

lamina) rudimentary, visible mainly as region of more closely spaced longitudinal vessels. Longitudinal

I vessels of stigmata numer between ous, withtransverse about 5vessels. betweenStibranchial gmata arranged folds anwith d abolong ut 8-12 axis orunning n each fdorso-ventraUy old; with about in10branchial -12 rows

sac, characteristic for genus. Stigmata fairly short, broad, with each crossed at midpoint by very thin

i parastigmatic internal longitudinal vessel. Dorsal lamina cleft into series of closely spaced languets. An

unidentified notodelphyid copepod found in branchial sac.

Stomach long, tubular, smooth without plications. Small lobes of the hepatic gland (liver) visible at

I anteri presentor onendeach of stside, omach. withIntestine left onelocompletely ng, tubular,within recurved; gut loop. with margin Ech ovary of anuslong,slightly slender,lobed. withOne numerous gonad

irregularly lobular testes arranged alongside most of length of left ovary and along entire length of right

I ovary. Sizes of specimens examined range from 3-10 mm wide as measured across siphons. All specimens

very flattened, no more than 4 mrn thick. All immature, but with gonads fairly well developed in larger

I specimens. Rem_,rks. The most striking difference between Boltenia polypIacoderma and the other BoItenia

species is the large number of plates on the anterior and lateral regions of the tunic. It could be confused

I with Chelyosoma spp. were it not for the square four-sided apertures, branchial folds, branched oral tentacles and dorso-ventrally arranged stigmata. The tunic pattern resembles that of the European ascidian Pyura

tessellata, so a re-examination was made of several specimens kindly provided by Dr. Ib Svane of the

I Kristineberg Marine Station in Sweden. The stigmata in P. tessellata are oriented longitudinally as in the other Pyura spp. Pyura tessellata has only four branchial folds per side in the branchial sac, a much thicker

tunic, and the body is compressed laterally, while Boltenia polyplacoderma has six branchial folds per side

i and is flattened antero-posteriorly.

Distribution. Canada, Barkley Sound, Vancouver Island, Canada, 61-185 m (C.M. Young, pers.

comm.). California, Santa Maria Basin, 98-116 m. !

Haloeynthia igaboja Oka, 1906

I Figure 11.11

Halocynthia igaboja Oka, 1906:45.--Van Name, 1945:362-363 (synonymy).--Abbott, 1975:644.--Abbott

I and 1980:212-21 12.40a-b.--Lambert 1987:469. Newberry, 3, figs. et aI.,

Material Examined. California, Santa Maria Basin, north of Point Buchon, Sta. 6, 109 m, 2

I specimens; Sta. BRA-27, sample 2, 96-126 m (1); off Purisima Point, Sta. BRA-21, sample 2, 75-90 m (2); Sta. BRA-20, sample 2, 90-130.5 m (1); Sta. BRA-16, sample 2, 91.5-123 m (2, vouchers, USNM), ( 3

vouchers, SBMNH); Sta. BRC-13, sample 1, 92-100 m (3); Sta. BRA-13, sample 1 (1); off Point Arguello,

I Sta. BRA-4, sample 2, 168-237 m (4).

Description. From Van Name (1945). Body oval or globular, attached at posterior end, covered

densely with non-calcareous flexible spines 10-12 mm, each long, straight, tapering, and thornlike with

I recurved branches along sides and circle of obliquely extending straight branches at tips. Between longer spines, many small, more-or-less branched spines extending out from test surface. Body up to 8-10 cm long,

4.5-6 cm wide. Branchial aperture terminal, with atrial openings located dorsally and one third to one half

I bomatter. dy length back. Apertures reddish during life; rest of body brownish due to adherent mud and other

I

I 283
!

!

!

lr

!

" B C l

!

II

E I

Figure 11.11. Halocynthia igaboja. A, whole animal, x2.3 (redrawn from Abbott and Newberry, 1980); B, a large 1

tunic spine, x4; C, a smaller spine without terminal whorl of spinelets, x4; D, two of the short l

whorled spines, x27; E, a single branched oral tentacle, greatly enlarged. (B-E from Ritter, 1907).

I

|

Mantle thin, transparent, with large muscle bands extending nearly entire body length from bases of i

siphons and crossing obliquely. Oral tentacles 12-50, with larger ones pinnately branched. Branchial sac

with 9-10 folds per side and sometimes with poorly developed eleventh fold on one or both sides. Folds with i

20 or more longitudinal vessels, but only 1 vessel on each interval between folds. I

Two to 16 gonads on fight side, 5-14 on the left, each consisting of an elongate, tubular, sinuous m

ovary with numerous small testes. Gonads positioned antero-postefiorly, with anterior ends extending across I

1

intestinal loop.

Remarks. The test spines form a mat that traps sediment, in some cases so much that the siphonal I

openings are hidden and the ascidian resembles a fuzzy ball of mud. This species is distinguished from l

another spiny solitary ascidian, Boltenia echinata, by the longitudinal stigmata, the larger number of branchial

folds, and the 4-7 gonads per side (B. echinata has just one on each side). The species is more common in l

colder, well-circulated waters. I

Distribution. Japan.--North America, Alaska to southern California; low intertidal to 165 rn, on

rock, shelly and gravelly bottoms; Santa Mafia Basin, 75-237 m. •

l

!

I
!

I Pyura haustor (Stimpson, 1864)

Figures 11.2, 11.12

I Cynthia haustor Stimpson, 1864:159.

i Pyura haustor (synonymy).--Abbott, Hartmeyer, 1909-1911:1340.--Van 1975:646, 653.--AbbottName, and Newberry, 1945:338-340, 1980:210, pl. 31fig. figs.12.373,4,a-b.--Lambert text fig. 223

et al., 1987:472, fig. 23.5.

I Material Examined. California, Santa Maria Basin, north of Point Sta.

Buchon, BRA-27, sample

1, 96-126m (1); off Purisima Point, Sta. BRA-20, sample 4, 90-130.5 m (1); Sta. BRA-16, sample 4, 91.5-

123 m (2 vouchers, SBMNH), sample 1 (5); Sta. BRA-13, 1 sample, 92-100 m (1); Sta. BRC-13, 88.5-

I 100.5m (voucher, USNM).

Description. From Van Name (1945). Body taller than wide, 5 cm long or more. Siphons usually

i conspicuous tubes, with branchial tubes larger, longer, directed obliquely forward or upward. Atrial tube arising near anterior end. Tubes vary greatly in length, from very short to longer than body width, sometimes

crooked. Test red or reddish orange, tough, leathery, often discolored with mud and completely encrusted

i with debris or other organisms except near ends of siphons. Test coarsely wrinkled, often elevated into high, sharply defined ridges that meet and cross each other in various directions or sometimes run roughly parallel

and which may have irregular tubercles, giving test appearance of coverinig by network of irregular polygonal

meshes. Surface smoother near ends of siphons, covered with minute slender spines 0.25-0.5 mm in length,

I lacking on body.

elsewhere

On mantle, prominent muscle bands extend down sides of body from bases of siphons, these

i irregularly crossed by many narrow, superficial bands; each siphon with strong ring muscles at base.

I a !!_

!

!

!

! B

!

Figure 11.12. Pyura haustor. A, whole animal, life size (redrawn from Kozloff, 1990) (a - atrial siphon, b -

i branchialsiphon); B, aclumpof animals, x0.4 (from Herdman, 1898).

I 285
!

Oral tentacles 15-30, typically 21, in species from British Columbia and Puget Sound, 42 in specimens i

from south of Point Conception, irregular in size and distribution, compound pinnate and sometimes

extensively branched. Branchial sac with 6 rather wide folds on each side, each with many closely spaced •

longitudinal vessels; transverse vessels numerous, of different sizes. U

Digestive tract forming an elongate, widely open loop. Hepatic organ or liver large, with several

extensively branched glands arising separately from stomach. Individual sacs of gonads irregular in size •

and shape; when numerous may be crowded out of normal serial position along common duct, forming g

more than 2 rows or appearing irregularly distributed on inner surface of mantle; large specimens with 50 or

more gonads on right side, fewer on left. •

g Distribution. British Columbia south to the Mexican border. Abundant intertidally and in shallow

subtidal, often growing in huge masses on rocks, pilings and floats. Subtidally occurring rarely as deep as

279 m; Santa Mafia Basin 88-I23 m, on hard substrate. I

Family Styelida i

Styela gibbsii (Stimpson, 1864)

Ill

Figure 11.13 A-C B

Cynthia gibbsii Stimpson, 1864:159. •

Styela gibbsii Huntsman, 1912a:149, pl. 12, fig. 5, pl. 19, figs. 10, 11.--Van Name, 1945:313-314, fig. 208

(synonymy).--Abbott, 1975:646.--Lambert et al., I987:472.

mi

Material Examined. California, Santa Mafia Basin, off Point Arguello, Sta. BRA-6, sample 2, 61 I

m (1 voucher, USNM).

Description. From Van Name (1945). Body elongate-cylindrical, both apertures on short siphons I1

located close together on anterior end, atrial aperture slightly posterior to branchial aperture. Attachment at

posterior end, somewhat narrow, with few rootlike processes. Test tough, with outer surface bearing numerous

wrinkles; wrinkles mainly longitudinal or oblique, generally crossed by less prominent circular wrinkles on •

some parts of body. At anterior end and on bases of siphons, wrinkles break up into tubercles. Color brown

or brownish yellow, more-or-less discolored with mud, becoming reddish around apertures. Size up to 6 cm

long, 1.5-2 cm wide; smaller specimens may be proportionately wider. Often grows in clusters, n

Mantle thin, with inconspicuous musculature. Branchial tentacles 25-40, unbranched. Branchial

sac with 4 well-developed folds on each side, with first (most dorsal) usually highest, third next highest.

Intemal longitudinal vessels numerous. Transverse vessels quite regular and of about 4 orders. B

g

Stomach elliptical, about twice as long as wide, with up to 30 longitudinal folds. Intestine bending

parallel to stomach, then bending forward again forming rectum. Margin of anus lobed. m

Two gonads per side; ovaries each long, moderately straight and tubular; testes numerous small, 1

somewhat elongate and irregularly shaped along posterior third or half of ovary; ovaries longer on right side

of body, extending for most of body length; on left side, ovaries shorter, more anteriorly located, especially

ventral one. I

Distribution. British Columbia to southern California, usually in shallow water in northern part of

range but deeper in southern part of range, to 91 m, on rocks; Santa Mafia Basin, 61 m. I

i

286 I
!

I Styela milled Ritter, 1907

Figure 11.13 D-E

!

Styela miIleri Ritter, 1907:21.--Van Name, 1945:308-309, fig. 204.

i Tethyum milleri Hartmeyer, 1912:374, 378.

Material Examined. California, Santa Maria Basin, off Point San Luis, Sta. BRA-25, sample 2,

64-5-72 m (2); off Purisima Point, Sta. BRA-20, sample 3, 90-130.5 m (1, SBMNH); Sta. BRA-14, sample

I 232, 96-105 (1 voucher, USNM); Sta. BRC-13, 3, 88.5-100.5 Santa Barbara

m sample m (1).--Western

Channel, Sta. BRC-1, sample 2, 69-73.5 m (2).

i with Description. From Ritter (1907) and Van Name (1945). Body cylindrical to ovoid or dome-shaped;

surface broken by few irregular low wrinkles anteriorly andlarge patches of warty thickenings posteriorly;

scattered minute spines may be present. Color yellowish brown, warty areas darker. Test very thin, parchmentlike, readily detachable from mantle, dull white on inner surface. Posterior anchoring part of test spreading

I out in thin border onto attaching surface; area of attachment with many short filamentous processes. Mantle

thin, with longitudinal and circular muscle fibers uniformly distributed, not in definite bands. Size small:

Ritter's single type specimen 22 mm long (incorrectly given as 22 cm), 12 mm wide.

I No siphons present, but apertures distinct, both 4-lobed, at anterior end. About 30 oral tentacles

present, of several lengths. Four branchial folds per side, all broad, more like undulations except one next to

• endostyle on fight side; 2 folds on each side next to endostyle drawn out anteriorly into long processes.

|

Internal longitudinal vessels numerous, 140 or more per side in adults, closely crowded on and between

folds (18-20 vessels on largest folds). Dorsal lamina broad thin membrane; endostyle heavy, irregularly

i tortuous. Stomach elongate, melon-shaped, with numerous ridges or folds; proximal part of intestine short, bent closely back on stomach; longer rectal section extending forward dorsally, almost to atrial aperture.

Ovary a single elongated cylindrical mass on each side, extending anterodorsatly to posteroventrally. Testes

i in crowdedRemarks. group around Specimens poteriinothis r endcollection of ovaries. are all smaller than Ritter's (1907) holotype and differently

shaped, being flattened anteroposteriorly, the largest one 13 mm wide and 5 mm tall; they correspond more

I thas o VanlongName siphons; (1945:thisFig. might 204).indicate No siphthat onsthey are visible were retracted externally, whenbutpreserved. when remoThe ved test fromisthesandtestencmsted, the body

although this character was not listed by either Ritter or Van Name. The test is thin and parchment-like,

corresponding to the species description. Gonads are very large relative to the small body size, especially

I the testes.

Distribution. This is an abyssal Pacific species, recorded from red or gray mud in depths of 825-

I 4074 this species m off sowas uthern collected California, from the the Bay SantaofMaria PanamaBasin , Peru, andandWestern possiblySanta southern BarbaraChilChannel, e. In the current 64-130 survey m.

!

!

!

!

I 287
i

I

I

I

I

I

,

,

I

Figure 11,13. Styela gibbsii. A, whole animal, xl.4 (redrawn from Huntsman, 1912a); B, piece of branchial sac,

x3Na1me, ; C,1945). alimentary D, body canalouand tline,gonx2; ads,E,xterminal 1.6 (B anpart d Coffromgonad, Herdman x6. (a-1898); atrial Sty siphon, ela mill Iveri

- longitudinal (from Van I

vessels, o - ovary, r - rectum, s - stomach, sd - sperm duct, t - testis.

I

I

I

I

288 I
!

I Styela tesseris Lambert, 1993

Figures 11.14, 11.15

I

Styela tesseris Lambert, 1993:116-118, figs. 5-6.

I Material Examined. California, Santa Maria Basin, off Purisima Point, Sta. BRA-21, sample 2, 75-90 m (holotype, USNM 20011); off Point ArgueUo, Sta. BRA-6, sample 2, 54-63 m (paratype, USNM

20012).

I Description. Holotype and both immature; 2 mm long, 15 long, 8

paratype paratype holotype

ram wide across siphons. Body flattened laterally; tunic with distinctive, irregularly tessellate pattern, with

flat-topped plates raised and separated from one another by furrows over entire tunic except adjacent to

I siphonal openings, where tunic smooth. Pattern resembles mosaic tilework with narrow between

groove

tiles. Tunic between tessetlations flexible, changing shape as tunic bends or moves. Platelike tunic thickenings

may overlap at edges in contracted areas of tunic. Furrow between each raised area presumably allowing for

I great expansion and contraction. Tunic thick, with smooth inner white layer; with outer tessellations not

penetrating to white layer. At posterior end of both animals, tunic forms wide, short, laterally flattened

stalk; body parts not extending into this region; with small scattered sand grains embedded superficially in

I tunic. Tessellations largest, most pronounced anteriorly; posteriorly becoming progressively smaller, flatter,

less distinct; absent on stalk region; tesselations running in more-or-less horizontal pattern across tunic

dorsoventrally, especially in more posterior region.

I siphons square openings, siphon siphon slightly posterior

Both with branchial terminal, atrial to

branchiae. On mantle, numerous very fine longitudinal muscles extend full length of body, with only few on

siphons. Deeper circular muscles predominate on siphons. Opening of dorsal tubercle simple oval slit. With

I about 40 long slender unbranched oral tentacles, mostly of 2 alternating sizes, and a large number of

very

tiny simple atrial tentacles. Dorsal lamina a wide flat membrane. Branchial sac with 4 folds per side, with

following numbers of longitudinal vessels (numbers in parentheses referring to vessels on folds): Right

I side, counting from the dorsal lamina to the ventral endostyle: 6 (6) 3 (6) 3 (6) 5 (5) 3; left side, starting from dorsal lamina: 2 (6) 3 (6) 3 (6) 3 (6) 3. On flat areas of sac, 4 stigmata between internal longitudinal vessels;

each stigma crossed at mid-point by very thin parastigmatic transverse vessel.

I Stomach elongate, with about 23 longitudinal folds or ridges, part of intestine folding

Proximal

back alongside stomach; distal portion curving anteriorly toward atrial siphon. Hepatic gland absent. In

larger animal 2 thin-walled gastro-intestinal connections (pyloric ducts) present within first intestinal curve;

I much wider than second. With duct in smaller animal.

one single pyloric

Holotype with 2 long, tubular ovaries on each side, with few irregularly lobed testes grouped around

i posterior end of left anterior ovary, which is the most fully developed. Iindividual sperm ducts forming common sperm duct running along surface of ovary to end near mouth of oviduct and base of atrial siphon;

right anterior ovary least developed, without eggs. Gonads not developed in smallest specimen. Numerous

endocarps of various sizes attached to inner side of body wall around posterior regions of gonads and

I stomach.

Remarks. The most obvious way in which Styela tesseris differs from all the described NE Pacific

i Sdeeper tyeIa swater, pecies but is thenever tessell recorded ated pafromttern the of thePacific. tunic.SThi tyelas s

tesseris pecies most differsclosel fromy reS.satlantica embles S.inatIant the tessellated ica, from

pattern on the tunic, the fewer internal longitudinal vessels on and between the folds in the branchial sac,

and the rather wide dorsal lamina even in these immature specimens. In addition, the dorsal tubercle is an

I oval slit without inrolled horns.

Distribution. California, Santa Maria Basin, 54-90 m, hard substrate.

I

i 289
I

Figure 11.14. Styela tesseris. A, whole animal, x7; B, detail of tunic, x23; C, dorsal tubercle; D, small piece of •

branchial sac (C-D greatly enlarged); E, body removed from tunic, right side, x7. (a - atrial siphon,

II

b - branchial siphon, Iv - longitudinal vessel, s - stigmata): (from Lambert, 1993).

I

I

290 I
i

I

m b_ a

at

!

II

, ii ,

I

i

I B °

I

c

I

I Figure 11.15. Styela tesseris. A, animal opened to show internal anatomy, x9; B, enlarged detail of gonad; C,

detail of stomach and anterior intestine showing the delicate connection between them. (a - atrial

siphon, at - atrial tentacles, b - branchial siphon, bt - branchial tentacles, e - endocarps, es - esophagus,

I i - intestine, o - ovary, pd - pyloric duct, s - stomach, sd - sperm duct, t - testis): (from Lambert, 1993).

I

I

I

i 291
!

Literature Cited I

Abbott, D.P. 1975. Phylum Chordata: introduction and Urochordata. Pp. 638-655, In: R.I. Smith and J.T. 1

Carlton (eds.), Light's Manual: Intertidal Invertebrates of the Central California Coast, 3rd ed.

University of California Press, Berkeley.

I

Abbott, D.P. and A.T. Newberry. 1980. Urochordata: the tunicates. Pp. 177-226, Chapter 12, In: Morris, 1

R.H., Abbott, D.P., and E.C. I-Iaderlie(eds.), Intertidal Invertebrates of California. Stanford University

Press, California.

Bancroft, F.W. 1899. Ovogenesis in Distaplia occidentalis Ritter (M.S.), with remarks on other species. I[

Bulletin of the Museum of Comparative Zoology 45:59-112.

Hartmeyer, R. 1909-1911. Ascidien. In: Bronn, H.G., Klassen und Ordnungen des Tierreichs 3:1281-1773. I

Hartmeyer, R. 1912. Zur Verbreitung nordischer Ascidien. Sitzungs Berichte der Gesellschaft der

Naturforschend der Freunde zu Berlin, ann. 1911, pp. 439-440. 1

I

Herdman, W.A. 1898. Descriptions of some simple ascidians collected in Puget Sound, Pacific coast.

Proceedings and Transactions of the Liverpool Biological Society 12:248-267. •

Huntsman, A.G. 1912. Ascidians from the coasts of Canada. Transactions of the Canadian Institute, ann. I

1911:111-148.

Huntsman, A.G. 1912a. Holosomatous ascidians from the coast of western Canada. Contributions to Canadian 1

Biology, Ottawa, pp. 103-185.

Ill

Johnson, M.E. and H.J. Snook. 1927. Seashore Animals of the Pacific Coast. Dover Publications, Inc., New II

York, 659 pp.

Kozloff, E.N. 1990. Invertebrates. Saunders Cotlege Publishing, 866 pp.

Lambert, C.C., G. Lambert and E.N. Kozloff. t987. Phylum Urochordata, Class Ascidiacea. Pp. 467-476, 1

In: Kozloff, E.N. (ed.), Marine Invertebrates of the Pacific Northwest. University of Washington

Press. I

Lambert, G. 1979. Early post-metamorphic growth, budding and spicule formation in the compound ascidian,

Cystodytes lobatus. Biological Bulletin 157:464-477.

Lambert, G. 1992. Ultrastructural aspects of spicule formation in the solitary ascidian Herdmania momus B

(Urochordata, Ascidiacea). Acta Zoologica 73:237-245.

Lambert, G. 1993. Three new species of stolidobranch ascidians (Chordata: Ascidiacea) from the California 1

i

continental shelf. Proceedings of the California Academy of Sciences 48(4): 109-118.

Lewin, R.A. and L. Chang (eds.). 1989. Prochloron, a Microbial Enigma. Chapman and Hall, New York. I_

196 pp.

Monniot, C., F. Monniot and P. Laboute. 1991. Coral Reef Ascidians of New Caledonia. Orstom Editions,

Paris. 247 pp. i

Monniot, C., F. Monniot and F. Gaill. 1975. Les Sorberacea: une nouvelle classe de tuniciers. Archives de

Zoologic Exprrimentale et grnrrale 116:77- t22. 1

Oka, A. 1906. Notizen uber japanische Ascidien I. Annotationes Zoologicae Japonenses 6:37-52.

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I Ooishi, S. 1991. North Pacific copepods (Cyclopoida: Ascidicolidae) associated mostly with compound

ascidians. Proceedings of the Fourth International Conference on Copepoda. Bulletin of the Plankton

I Society of Japan, special volume, pp. 49-68. Plough, H.H. 1978. Sea Squirts of the Atlantic Continental Shelf from Maine to Texas. Johns Hopkins

University Press, Baltimore, 118 pp.

I Ritter, W.E. 1903. The structure and affinities of Herdmania claviformis, the type of a new genus and

family of ascidians. Mark Anniversary Volume, pp. 237-261.

I Ritter, W.E. 1907. The ascidians collected by the United States Fisheries Bureau steamer Albatross on the

coast of California during the summer of 1904. University of California Publications in Zoology

4:1-52.

I Ritter, W.E. and R.A. Forsyth. 1917. Ascidians of the littoral zone of southern California. University of

California Publications in Zoology 16:439-512.

I Stimpson, W. 1864. Descriptions of new species of marine Invertebrata from Puget Sound collected by the naturalists of the Northwest Boundary Commission. Proceedings of the Academy of Natural Sciences

Philadelphia 16:153-161.

I Van Name, W.G. 1945. The North and South American Ascidians. Bulletin of the American Museum of

Natural History 84:1-476, 31 pls.

I Young, C.M. and E. Vazquez, 1995. Morphology, larval development, and distribution of Bathyperafeminalba

n. sp. (Ascidiacea: Pyuidae), a deep-water ascidian from the fjords and sounds of British Colombia.

Invertebrate Biology 114:89-106.

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I 293
IIlIIlIIIIiIlIlIIl
Appendix

Lists and Maps of Stations

Table A.L Position of soft-subslrate stalions taken during the Phase I Reconnaissance.

Station Latitude Longitude Depth

(m)

1 35°27.86'N 121°05.33'W 98

2 35°27.70eN 121°06.5 2'W 200

3 35°27.07'N 121° 10.20'W 291

4 35%6.56'N 121o| 4.93'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 121°18.65'W 591

11 35°17.80'N 121°22.13'W 690

12 35°15.03"N 120057.3 I'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 I21°16.29'W 591

17 35°11.61'N 121°22.55'W 654

18 35°09.08'N 120°56.55'W 197

19 35°08.93"N 120°59.66"W 296

20 35°15.72"N 121°04.68'W 396

21 35°06.11 _N 120°44.82"W 49

22 35°05.85'N 120°50.23'W 99

23 35°05.60'N 120°55.18"W 195

25 35°05.07'N 121°00.75"W 390

26 35°04.38'N 121°15.99'W 590

27 35°04.30'N 12l°19.27'W 611

28 35°04.22'N 121°19.65'3,.V 603

30 34"54.19'N 120°47.07"W 98

31 34°53,76'N 120°52.96"W 200

32 34°53.56"N 120°56.81'W 297

33 34°53.43'N 120°59.66"W 396

34 34°53.15'N 121°04.40"W 492

35 34°52.96'N 121°10.30"W 548

36 34°52.77'N 12i °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 34049.24'N 121°00.8 I'W 392

41 34°48.35'N 12I°19.14'W 495

42 34048.04'N 120o47.5 0'W 100

43 34°46.59"N 120°52.92"W 197

45 34044.91 'N 120°59.59"W 395

46 34°41.22"N 121°13.56"W 597

47 34°41.99"N 121°10.81"W 378

48 34045.1 I'N 120°52.85"W 196

49 34°45.03'N 120°56.3 I'W 290

50 34°37.80'N 121°01.66"W 591

52 34°39.56'N 120°47.64'W 98

53 34°37.69'N 120°50.38'W 196

54 34°36.57'N 120°52.02"W 396

55 34°33.66'N 120056.3 I'W 590

56 34°30.32'N 121°01.02"W 900

295
|

Table A.1 (Continued)

m

Station Latitude Longitude Depth m

(m)

m

58 34°34.35"N 120°45.18'W 99 •

59

60

34°33.65

34o33.25

'

'

N

N 120°48.34

120°47.18

'

'

W

W 216

275

m

61 34o33.01'N 120°48.89"W 345

62 34o30.46"N 120°52.13'W 582 •

63 34°26.29'N 120°58.08'W 930

64 34°33.15'N 120°40.90'W 59

65 34°31.27'N 120°43.27'W 107 m

66 34°30.46'N 120°44.55'W 201 •

67 34°30.29'N 120°45.50'W 282

68 34°29.24'N 120°45.99'W 390

69 34°22.88'N 120°54.20'W 927 am

70 34°29.67'N 120°43.70'W 200

71 34°29.04"N 120°44.01'W 306 l

72 34°28.4 I'N 120°44.76'W 401

73 34°28.2 I'N 120°36.80"W 98 •

74 34o26.84"N 120°38.61"W 20l l

75 34°26.08"N 120°39.65'W 293

76 34o25.59"N 120°40.98"W 387

77 34o22.62"N 120°44.02"W 578 m

78 34°18.78'N 120°49.30'w 762

79 34°24.12'N 120°28.32'W 98

80 34o22.86"N 120°28.34'W 196 am

81 34°21.26'N 120°28.83"W 294 •

82 34°18.7 I'N 120°29.55'W 394

83 34o17.20"N 120°30.2ffW 444

84 34°13.54'N 120°31.19'W 394 •

85

86

34

34°24.45

°25.88'

'

N

N

12

12

0

0

°

°

1

17.

6.3

02

I

'

'

W

W 113

197 m

87 34°21.60'N 120°17.11"W 299

88 34o17.89'N 120°16.86"W 393 •

89 34° 13.79'N 120°16.56'W 471 l

90 34°09.44'N 120°16.30"W 375

91 34 °11.73'N 120°07.43'W 540

92 34°08.70"N 120°07.50'W 444 m

93 34o07.63'N 120°07.5 l'W 357

96 34°22.9 I'N 120°05.42'W 296

94 34°24.54'N 120°05.47'W 96 m

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 56 l

99 34 °11.22"N 120°05.86"W 540 m

100

101

34o08

34°07.5

.67

I

,

'

N

N

120o05.50

120°05.65'

"

W

w 4

357

43 m

102 34°59.7 I'N 120°48.22'W 99

103 34o59.63"N 120°53.56"W 197 •

104 34o59.45'N 120°56.49'W 294 l

105 34o59.23'N 120°59.60"W 392

106 34°58.95'N 121°04.42'W 492

107 34o58.65'N 12t°15.08"W 573 m

108 34o58.21'N 121° 17.88'w 492

Note: Sample labels from the Soft-substrate stations have several identification codes which include a station number, sample m

type, replicate number, and analysis type. These are as follows: 00l to 200 = the range of station numbers; BSS = Benthic •

Sediment Single (i.e., a non-replicatod station); BSR = Benthic Sediment Replicate (three replicates taken at this station); BSV =

Benthic Sediment Variance (subsamples); 01-09 = replicate numbers; TX = a taxonomy sample. Sample labels having the

designation BRA, represents a sample from rocks taken as part of the hard bottom survey. •

|

296 m
I

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!

li_dre A.1. Map showing location of soft-substrate stations from the Phase I Reconnaissanceand Phase II

i MonitoringPrograms.

I

i 297
!

Table AS. Location of soff-subs_a_ stations _ken during _e Phase N Momtofing Program. I

Station Latitude Longitude Depth g

(m)

I

R-1 35°05.83'N 120°49.16"W 91 •

R-2 35°05.5_N 120°53.40'W 161 II

R-3 35°05.3_N 121°00.9_W 409

R-4 34°43.01_N 120°47.39%V 92 I

R-5 34°42.69_N 120°50.83%V 154 []

R-6 34°41.4_N 120°57.90%V 410 U

R-7 34°52.9_N 121°10.30%V 565

R-8 34°55.30_N 120°45.87'W 90 1

R-9 34°53.68_q 120059.12%V 410 l

PJ-I 34°55.79qq 120°49.91_V 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 1

PJ-5 34°55.32'N 120°50.24'W ! 52

PJ-6 34°54.71'N 120°49.91'W 148 I

PJ-7 34°55.79"N 120°48.60"W 123 []

PJ-8 34°56.87'N 120°49.91"W 142

PJ-9 34°55.79"N 120°51.23'W 169

PJ-10 34°53.63"N 120°49.91"W 147 1

PJ- 11 34°57.95"N 120°49.91"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 1

PJ- 15 34°55.79'N 120°50.57%V 155 |

PJ- 16 34°55.03'N 120°48.99%V 130

PJ- 17 34056.56'N 120°48.98"W 126

PJ- 18 34°56.56'N 120°50.84'W 158 []

PJ- 19 34055.03'N 120050.84_V 167 I!

PJ-20 34°50.38q_ 120°49.91"W 148

PJ-21 35°01.23q_ 120°51.15_V 143

PJ-22 34°55.25qq 120°49.93%V 143 g :

PJ-23 34°56.33'3I 120°49.90%V 143 •

!

Table A.3. Sampling dates of MMS Phase II Monitoring Program. I

Cnfise Date I

1-1 October 1986

1-2 January 1987

1-3 May 1987 •

2-1 July 1987

2-3 October 1987 l

2-4 January 1988

2-5 May 1988 ms

3-1 October 1988

3-4 May t989 l

I

298 1
Table A.4. MMS Phase I - Locations of hard-substrate Iransects.

Station Beginning End Depth

Latitude Longitude Latitude Longitude (m)

I A/B 34°24.454"N 120°01.876"W 34°24.464'N 120°00.878'W 69-73.5

1C/D 34°24.076'N 120°00.443"W 34°24.184'N 120°01.480'W " 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°t0.984'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°314.315"W 54-63

13 A/B 34°42".570'N 120°4'7.899'W 34°42.107'N 120°48.253'W 92-100

13 C/D 34°42.556'N 120°48.147'W 34°42.974'N 120°47.424'W 88.5-100.5

14 A/B 34°43.589'N 120°49.093"W 34°42.826"N 120°48.370'W 96-105

14 C/D 34°43.244"N 120°49.406'W 34°42.893"N 120°48.822'W 105-117

16 A/B 34°46.544'N 120°50.197'W 34°45.912'N 120°49.726'W 91.5-123

I7 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 I20°45.903"W 34°47.548"N 120°46.123'W 75-90

22A/B 34°50.365'N 120°48.22 I'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°I 1.586'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 I20°59.64 I'W 35021.867"N 120°59.299'W 96-105

29 A/B 35°27.864'N 121°05.331'W 35027.805'N 121°05.277"W 102-106.5

Table A.5. MMS Phase II - Locations of hard-subslrate photosurvey stations.

Station Latitude Longitude Depth

(m)

PH-E 34°30.26'N 120°42.76"W 119

PH-F 34°30.8 I'N 120042.36"W 105

PH-I 34°29.96'N 120041.68"W 107

PH-J 34029.82'N 120o41.82"W 117

PH-K 34°29.37"N 120°42.26"W 160

PH-N 34029.2 I'N 120°42.05'W 166

PH-R 34029.1 I'N 120°42.67'W 213

PH-U 34°31.4 8"N 120°43.5 I'W 113

PH-W 34°31.52'N 120°45.86'W 195
!

•, _,!_'_no _,: , J , : ...... _ _ 1 _ I I

, • -__ i__:!iii;_:__:_!_i ] > : i _ i_:!__!:_:!_ _i?!1

,, _ii_::_:i:!_, ?!_:i ],ii;_!_;,i_i _i/!:i:i_ _ b__iii__:1

. . ___ _ c_o_,_ __, I

:

'-'_ _°,o-. HI

I

_ ,,_ i ! ,:' 23 " "

i,, _ "1 /

i..le e,, 2.2 I I

i "1" ') /

m-w • I1"; / ...... • 2 .......... /

.o_o., _.-Jw 1t/ / /

P.-K• II: / / II

I !

34°28'N 120 °44'W

I

Figure A.2. Map showing location of hard-substrate stations from the Phase I Reconnaissance and Phase II

Monitoring Programs. Phase II stations are indicated in the inset. I
I

I

I

i

I

I •

I

I LJ

I

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 cultural values of our national parks and historical places; and providing for the enjoyment of life through outdoor recreation. The Department assesses

our energy and mineral resources and works to ensure that their development is in the best

interests of all our people by encouraging stewardship and citizen participation in their care.

I The Department also has a majorresponsibility for American Indian reservation communities and for people who live in islandterritories under U.S. administration.

The Minerals Management Service Mission

I _=_ As a bureau of the Department of the Interior, the Minerals Management Service's (MMS)

' primary responsibilities are to manage the mineral resources located on the Nation's Outer

Continental Shelf (OCS), collect revenue from the Federal OCS and onshore Federal and Indian

I lands, Moreover, and distribute those revenues. in working to meet its responsibilities,the Offshore Minerals Management Program

administers the OCS competitive leasing program and oversees the safe and environmentally

sound exploration and production of our Nation's offshore natural gas, oil and other mineral

I reffi esources. cient, timely The MMS and accurate Roylty cManagement ollection andProgram disbursementesofitsrevenue responsibi fromlitimineral es by ensuring _easingthe and

production due to Indian tribes and allottees, States and the U.S. Treasury.

I The responsive MMS strives to the topublic's fulfill itsconresponsibiliti cerns and interests esthroughby the maintaining general guiding a dialogueprinciples with alof: l potenti (1) being ally

affected parties and (2) carrying out its programs with an emphasis on working to enhance the

quality of life for all Americans by lending MMS assistance and expertise to economic

'BB development and environmental protection.

il

,