I

I OCS Study

MMS 96-0029

I

I

I TAXONOMIC ATLAS OF THE BENTmC FAUNA OF

THE SANTA MARIA BASIN AND

i WESTERN SANTA BARBARA CHANNEL

I

FINAL REPORT Volume 11of 14

!

The Crustacea Part 2 --The Isopoda, Cumacea, and Tanaidacea

I

Edited by:

I James A. Blake

Paul H. Scott

I Submitted by:

Science Applications International Corporation

10260 Campus Point Drive San Diego, California 92121

I For:

U.S. Department of the Interior

Minerals Management Service

I Pacific OCS Region 770 Paseo Camadlto

Carnarillo, California 93010

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

I

Minerals Management Service

I Pacific OCS Region July 1996

I
!

!

DISCLAIMER I

This Service, reportU.S. hasDepartment been reviewedof theby theInterior PacificandOuter approved Continental for publication. Shelf Region, TheMinerals opinions, Mangement findings, I

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

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

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

conformity with Minerals Management Service editorial standards.

TAXONOMIC DISCLAIM ER I

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

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

!

!

!

!

!

II

!

i,

!

!

!

Ji

!
!

!

PROJECT ORGANIZATION

!

I PROGRAM MANAGER

I DR. ANDREWL. L[SSNER Science Applications International Corporation

10260 Campus Point Drive

I San Diego, California 92121

I DEPtrrvPROGRAMANAGER

i DR. JAMES A. BLAKE

ENSR Consulting and Engineering

I 89 Water Street

Woods Hole, Massachusetts 02543

!

I PROJECT TECHNICAL OFFICER

MR.FRANKJ. MANA_O

I U.S. Department of the Interior

Minerals Management Service, Pacific OCS Region

I 770 Paseo Camarillo Camarillo, California 93010

!

Co_trr_c AUTHORS

I DR. RICHARD C. BRUSCA

DR. MAsArrmoDoJnu

I Ms. LINDAD. McCA_ DR. JORGEN SIEG

DR. LESWArLING

i Ms. REGINAWETZER

DR. GEORGE D.F. WtLSON

!

i iii
!

ACCESS NUMBER: 30484.11

TECHNICAL SUMMARY I

STUDY TITLE: TaxonomicAtlasofthe Santa MariaBasinandWesternSanta BarbaraChannel,

am

REPORTTITLE: TaxonomicAtlasof the Santa MariaBasinand WesternSanta BarbaraChannel.Volume i

11.The CrustaceaPart2 -- The Isopoda,Cumacea,and Tanaidacea.

an=

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

SPONSORINGOCS REGION: Pacific. i

[] APPLICABLEPLANNINGAREA: SouthernCalifornia.

FISCALYEARS OF PROJECT FUNDING:1990; 1991;1992;1993; 1994;1995. I

COMPLETION DATEOF REPORT: July 1996.

i

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

CUMULATIVEPROJECTCOST: $724,384. a

= PROJECT MANAGER: Andrew L. Lissner.

AFFILIATION: ScienceApplications InternationalCorporation. ii

ADDRESS: 10260 Campus Point Drive,San Diego, California92121.

II

PRINCIPALINVESTIGATORS': JamesA. Blake,Andrew L. Lissner. I

KEY WORDS: Marine invertebrates,Arthropoda, Crustacea, Isopoda, Cumacea, Tanaidacea,taxonomy, II

Catifomia,

SantaMaria Basin,SantaBarbaraChannel,continentalshelf, i

BACKGROUND: The TaxonomicAtlas ofthe Santa MariaBasinand western Santa BarbaraChannel isan

extensionof the benthicreconnaissance(PhaseI)and monitoringprograms(Phase II)that wereconducted i

by the MMS since 1983. The organisms that were collected as partof those programs providethe material n

on whichthe Atlas isdeveloped. Inorder tofully document the faunacollected by those programs,a series

of 14 volumeswill be preparedthat providekeys,descriptions,and illustrationsof the benthicfauna of the •

hard and soft substrateenvironments. A team of 40 experts onthe fauna hasbeen assembledto carryout =

this work and their contributionsare distributedamong the 14volumes.

OBJECTIVES: The objectivesof Volume11areto continuecoverageofthe Crustaceathat was initiatedin n

Volume12. Thisvolumetreatstheasellotandnon-asellotIsopoda,the Cumacea andtheTanaidacea. II

DESCRIPTION:Volume11includesatreatmentofthreeimportantcrustaceangroups,the isopods,cumacean i

and tanaids. The 3 chaptersare organizedinto sectionsthat includethe morphology,taxonomichistory,

biology,keysof species,and descriptionsof generaandspecies.Eachspeciesisfullyillustratedwithrelevant

characteristicslabeledandidentified.Detailsby groupinclude-- Isopoda:28 genera,37species,14 newto []

science, none described in chapter.-- Cumacea:10 genera, 16species, 10 new to science, all described =

here, onenew genus, one new subgenus.- Tanaidacea:14genera, 18 identifiedspecies, 3 indeterminate

species, 14 new to science, describedhere; 2 described earlier; 2 new genera, 1 described here; another

namedin an earlier paper. •

[]

i
I

ACCESS NUMBER: 30484.11

I SIGNIFICANT CONCLUSIONS: The presentvolumeincludesthreeimportantcrustaceanorders:Isopoda,

Cumacea,andTanaidacea,Amongthetsopoda,atotalof28generaand37speciesaretreated,ofwhich14

species,mostlyasellotesare newto science, NoneofthesenewtaxaareformallydescribedintheAtlas,but

I are designatedwith provisionalletteredcategories. The Cumacea are reported with 10 genera and 16 species, 10of which are new to scienceand alldescribedhere. One genusand one subgenusare alsonew

to science. All cumaceans areillustratedwith scanningelectron micrographsand line drawings. Amongthe

I Tathesnaeidspecies acea,1ar4egenneweratoascience, nd 18 spewithcie12saredescribed described, herewand ith a2nnamed other3inspeancies earlinierdetermi paper.nate Two.genera Fourteenwereof

found that were new to science, one is namedhere, anotherwas namedearlier. All three chapters include

i extensiveintroductorysectionswithdetailsofmorphologyandtaxonomichistories.Bibliographiesareextensive

and each chapter representsa major contribution to the systematics of benthic Crustacea in the eastern

Pacific.

I STUDY RESULTS: The fauna of the SantaMaria Basinand westernSanta BarbaraChannelis evaluated from samplesthat were taken fromdepthsof about50 to 1,000 m, Organismswere collectedfrom soft

sedimentsusingboxcoresandfromrocksusingmanipulatorarmsofsubmersiblesand remotelyoperated

i vavailable ehicles.toThtheecotelamlectofionsinvestigators. are organizeAdditional dinto setsmaterial of voucfrom hersfrtheombulk Phacollections se I and II now that haarchived ve beewith n made the

Natural HistoryMuseum of LosAngelesCounty isalsobeing examined. Thetotal numberof species treated

in the entire TaxonomicAtlas (14volumes) may exceed 1,300species.

I Atotalof52generaand74speciesarereportedforthecrustaceanordersIsopoda,Cumacea,andTanaidacea.

Ofthese, 38species were determinedto be new to scienceof which24 aredescribed as partofthis volume

I adescrib nd oneecompa dherein) nioandnpaper onenew (14ssubg peciesenus ofisodescpedribed stohere be dearescralso ibedinc insleudpaerd.atEeapchapchapt ers). Therincludes ree newgen exteensive ra(2

illustratedreviewsof the morphologyofthesecrustaceantaxaas wellas glossariesand keys. Eachauthor

I has withprrelat oviededdspextecies ensiasveliwell terature asexisting reviewsproblems. and synonymiesof knowntaxa. Discussionsprovidecomparisons

I STUDY PRODUCT: Blake,J.A.and P.H.Scott(Editors).1996. TaxonomicAtlasof the BenthicFauna ofthe Santa Maria Basinand Western Santa BarbaraChannel. Volume11. The CrustaceaPart 2--1sopoda,

CumaceaandTanaidacea.A finalreportpreparedbyScienceApplicationsInternationalCorporation,forthe

U. Study S. DMMS epartm96-0029. entof theContract Interior,No. Min14-35-0001-30484. eralsManagementService, PacificOCS Region,Camarillo,CA. OCS

!

I

I

I

I

I "P.l'saffiliationmaybedifferentthanthatlistedforProjectManager(s).

I

_o-z_-19_

RBPORTDOCUMENTATIONz.RrpomNO, . MMS 96-0029 2" 3. n.cu_e_sdk:c-._ No.

PAGE

4. Title aml $ul_itle 5. Repm,t Dm

TaxonomicAtlasoftheSantaMariaBasinandWesternSantaBarbaraChannel.Volume July 1996

11.The CrustaceaPart2 -- The Isopoda,Cumacea,andTanaidacea. 9

7. Author(s) 8, PerforminE Orlr.anization Rept, No.

JamesA. Blakeand PaulH. Scott (editors)

9. Performing Orpnizselon Name and Address 10. Pm_ct/Tamk/Work Unit No.

ScienceApplicationsInternationalCorporation ::. coof,,c,c)°, omof(G_.)

10260Campus PointDrive

San Diego,California92121 (c) 14-35-001-30484

(G)

12. Sponsori_ Orllnization Name ind Address 1,3. Type of Report & Period Cove_ed

Final 11 of 14volumes,

MineralManagementService,U. S. Department ofthe Interior, 9t89to 7/96 PacificOCS Region,770 Paseo Camarillo

Camarillo, California 93010 _4.

IS. SUpl_emeofory Notes

Contributingauthors: Dr. Richard C. Brusca, Dr. Masahiro Dojiri, Ms.Linda D. McCann,Dr. J0rgen Sieg, Dr,LesWatling,

Ms. Regina Wetzer,and Dr. George D.F. Wilson

IlL Abstract (Limit: 200 words_

The crustacean Orders Isopoda, Cumacea, and Tanaidacea are described with keys and illustraUons. A total of 52

generaand 74species are included inthis volume. For the Isopoda, a total of 28genera and 37 species aretreated,

of which 14species, mostly asellotes are new to science. None ofthese new taxa are formallydescribed in the Atlas,

butare designatedwith provisional letteredcategories. The Cumacea are reported with 10generaand 16species, 10

of which are new to science and all described here. One genus and one subgenus are also new to science. All

cumaceansare illustratedwith scanning electron micrographsand line drawings. Among the Tanaidacea, 14genera

and 18species are described, with another 3 species indeterminate. Fourteen of these species are new to science,

with 12 describedhere and 2 named in an earlier paper. Two genera were foundthat were new to science, one is

named here, another was named earlier. All three chapters include extensive introductory sections with details of

morphologyandtaxonomic histories. Bibliographies areextensiveand eachchapter representsa majorcontributionto

the systematicsof benthic Crustacea inthe eastern Pacific.

17. Document AmDlysis -. Descriptors

Marine invertebrates,Arthropoda,Crustacea,Isopoda,Cumacea,Tanaidacea,taxonomy,Califomia,Santa Maria

Basin, Santa BarbaraChannel,continental shelf.

b. rdtofi_rs/Ol:_m-EnckKI Trams

TaxonomicAtlas,Keysto species,faunal guide, marineecology, crustaceans,marinebenthos.

¢. COSATI Field/Group

1L Aveillbillty Statement - lg. Security Class (This Report) 21. No. of Pal[e$

Unlimited Unclassified 268

20. Security CIm (This Patie| 22. Price

See ANSI-,Z39.I8) See Instructions on Reverse OPTIONAL FORM 272 {4-l/_

(Formorly NTIS-3S)

I Ol_pll_me_ of Commerce
!

Table of Contents

!

1.0 THE ORDERISOPODA I

1.1 INTROI)UCTmNTOTHE_t__ARINEISO]PODA

(by Regina Wetzer, Richard C. Brusca and George D. F. Wilson) 1

MII

Introduction ..................................................................................................................................... 1

Summary of Isopod Research in Northeast Pacific Region ............................................................ 2 am

General Isopod Anatomy ................................................................................................................. 2 I!

Glossary of Technical Terms .......................................................................................................... 5

Phyletic Key to the Suborders of Isopod (Adults) .......................................................................... 7 ]1_

II 1.2 DESCRIPTIONS OF THE SPECIES OF THE SUBORDERS ANTmrSmF_, EPICARIDEA_ FLABELLIFERA_

GNATHIIDEA,ANDVALVIFERA(by Regina Wetzer and Richard C. Brusca)

Key to the Families of Anthuridea Known from California Waters ................... 10 !

Family Anthuridae Leach, 1814 ....................................................................................... 10

Key to the Genera of Anthuridae Collected as Part of the MMS Surveys ......... 11 I

Amakusanthura californiensis (Schultz, 1964) ...................................... 13 |

Cyathura munda Menzies, 1951 ............................................................ 13

Haliophasma geminatum Menzies and Barnard, 1959 .......................... 15 !

Family Hyssuridae W_igele, 1981 .................................................................................... 18 l

Kupellonura sp. A .................................................................................. 19

Family Paranthuridae Menzies and Glynn, 1968 ............................................................. 21 Is

Paranthura elegans Menzies, 1951 ....................................................... 22 Jl

Suborder Epicaridea Latreille, 1831 .............................................................................................. 22

lid

Family Bopyridae Rafmesque, 1815 ................................................................................ 24 m

Munidion pleuroncodis Markham, 1975 ................................................ 25 i

Suborder Flabellifera G.O. Sars, 1882 .......................................................................................... 28

Key to the Families of Flabellifera Known from California Waters .................. 28 J_

Family Aegidae Dana, 1853 ............................................................................................. 30 I

Key to California Species of Rocinela ................................................................ 30 I

Rocinela angustata Richardson, 1904 ................................................... 31 I

Family Cirolanidae Dana, 1853 ....................................................................................... 33 il

Metacirolanajoanneae (Schultz, 1966) ................................................ 34

Family Serolidae Leach, 1814 .......................................................................................... 34

SeroIis carinata Lockington, 1877 ......................................................... 36 i

i

Family Tridentellidae Bruce, 1984 .................................................................................. 36 !

Tridentella glutacantha Delaney and Brusca, 1985 .............................. 38

Suborder Gnathiidea Leach, 1814 ................................................................................................. 41 Ill

Key to the California Species of Gnathia (adult males) ..................................... 42 g

Gnathia crenulatifrons Monod, 1926 .................................................... 43

Gnathia productatridens Menzies and Barnard, 1959 ........................... 43 ]l

Gnathia sanctaecrucis Schultz, 1972 ..................................................... 43 al

Gnathia tridens Menzies and Barnard, 1959 ......................................... 47

!

°.°

VIII I
!

I Suborder Valvifera G.O. Sars, 1882 .............................................................................................. 47

Key to the California Families of Valvifera ....................................................... 47

I Family Arcturidae G.O. Sars, 1897 .................................................................................. 50 ldarcturus allelomorphus Menzies and Barnard, 1959 ......................... 50

Family Idoteidae Fabricius, 1798 ..................................................................................... 53

Key to the Genera of Idoteidae Collected as part of the MMS Surveys ............. 53

I Idotea (Idotea) rufescens Fee, 1926 ....................................................... 54

Key to Species of Synidotea Collected as Part of the MMS Surveys ........................................... 54

Synidotea calcarea Schultz, 1966 .......................................................... 56

I Synidotea Iverson, ..............................................................

media 1972 56

1.3 THE SUBORDERASELLOTA(by George D. F. Wilson)

I Introduction 59

General Morphology ..................................................................................................................... 60

Collection and Preservation .......................................................................................................... 60

i Laboratory ......................................................................................................................

Methods 61

Glossary and Terminology ............................................................................................................ 61

Key to the Families of the Asellota and the Species ..................................................................... 62

I Recovered from the Santa Maria Basin 62 Description of Asellotan Species .................................................................................................. 68

Family Stenetriidae Hansen, 1905 ................................................................................... 68

I Stenetrium A 69

sp.

Family Munnidae Sars, 1897 ........................................................................................... 71

Munna sp. A ........................................................................................... 71

Family Paramunnidae Munnogonium VanhCJffen, 1914 .......................................................................... 73 cf tillerae (Menzies and Barnard, 1959) ....................... 73

Pleurogonium G.O.Sars, 1883 ............................................................... 76

II Pleurogonium californiense Menzies, 1951 .......................................... 76

II Pleurogonium sp. A ............................................................................... 78

Family Janiridae Sars, 1897 ............................................................................................. 80

I Janiralata sp. A ...................................................................................... 80 sp. B ...................................................................................... 82

Janiralata sp. C ...................................................................................... 84

I Family Joeropsididae Janiralata Nordenstam, sp. D (cf.1933 rajata......................................................................... ) ................................................................... 8686

Joeropsis concava (Schultz, 1966) ........................................................ 88

l[ Joeropsis sp. A ....................................................................................... 90

Family Munnopsidae Sars, 1869 ...................................................................................... 90

Subfamily Eurycopinae Hansen, 191 .................................................................. 90

I Eurycope Belonectes californiensis sp. A ..................................................................................... Schultz, 1966 .................................................. 9294

Subfamily Ilyarachninae Hansen, 1916 ........................................................................... 96

I Subfamily incertaeIlyarachna sedis .................................................................................................. acarina Menzies and Barnard, 1959 ................................... 9698

Munnopsurus sp. A ................................................................................ 98

i Family NannonisMunnopsurus cidze Hansen,sp.191B ............................................................................................................................................................. 100 102

Nannonisconus latipleonus Schultz, 1966 ........................................... 102

I
!

Family Desmosomatidae Sars, 1897 .............................................................................. 104 I

Momedossa symmetrica (Schultz, 1966) ............................................. 104

Desmosoma sp. A ................................................................................. 106 _ll

Prochelator sp. A ................................................................................. 108 ii

Literature Cited ........................................................................................................................... 110

2. Cu]v_c_ (by Les Wafting and Linda D. McCann) i

Introduction ................................................................................................................................. 121

Cumacean Morphology ............................................................................................................... 121 I

Classification ............................................................................................................................... 127 i

m

Key to the Families of Cumacea with Species in the Santa Maria Basin Region ....................... 128

Family Diastylidae ......................................................................................................... 128 I

Key the Diastylidae from the Santa Maria Basin Region ............................. 128 If to

Diastylis sentosa Wafting and McCann, new species .......................... 130

Diastytis crenellata Wafting and McCann, new species ..................... 131 m

DiastyIis quadripIicata Wafting and McCann, new species ................ 133 m

Diastylis santamariensis Wafting and McCann, new species ............. 135

Diastylis californica Zimmer, 1936 ..................................................... 135

Diastylis pellucida Hart, 1930 ............................................................. 138 I

Leptostylis calva Wafting and McCann, new species .......................... 139 m

Leptostylis abditis Wading and McCann, new species ........................ 140 Jam

Family Lampropidae ...................................................................................................... 142 l[

Hemilamprops californicus Zimmer, 1936 .......................................... 142

Family Leuconidae ......................................................................................................... 144

Key to the Leuconidae of the Santa Mafia Basin Region ................................. 144 li

Jill

Alloeoleucon santamariensis Wafting and McCann, new species ....... 145 ll

Leucon (Diaphonoleucon) declivis Wading and McCann, new species .. 147

Leucon (Crymoleucon) bishopi Bacescu, 1988 .................................... 149 1

Leucon (Leucon)falcicosta Watling and McCann, new species ......... 151 I

Leucon (Leucon) armatus Given, 1961 ................................................ 151

Leucon (Leucon) magnadentata Given, 1961 ...................................... 153 m

Eudorella pacifica Hart, 1930 .............................................................. 155 I

Eudorella redacticruris Watling and McCann, new species ............... 156

Eudorella truncatula Bate, 1856 .......................................................... 156 I[

Eudorellopsis longirostris Given, 1961 ............................................... 159 II

Family Nannastacidae .................................................................................................... 159

Key to the Nannastacidae of the Santa Maria Basin ......................................... 160 m

Campylaspis canaliculata Zimmer, 1936 ............................................ 161 an

Campylaspis rufa Hart, 1930 ............................................................... 163

Campylaspis rubromaculata Lie, 1971 ................................................ 164 I_

Campylaspis maculinodulosa Wafting and McCann, new species ...... 164

Campylaspis hartae Lie, 1969 ............................................................. 167

Campylaspis blakei Watling and McCann, new species ..................... 168 m

Campylaspis biplicata Watling and McCann, new species ................. 170 l

ProcampyIaspis caenosa Watling and McCann, new species ............. 172

Cumella (Cumella) morion Wading and McCann, new species .......... 174 1

Cumella (Cumella) californica Wading and McCann, new species .... 175 II

l,

x !
!

m Acknowledgements ..................................................................................................................... 176

Literature Cited ........................................................................................................................... 177

n Attachment I ................................................................................................................................ 179 II ............................................................................................................................... 180

3. THE TANAmac_ (by Mas Dojiri and Jiirgen Sieg)

I Introduction ................................................................................................................................. 181

Taxonomic History ...................................................................................................................... 181

m External Morphology .................................................................................................................. 182

Internal Anatomy ......................................................................................................................... 195

Development ............................................................................................................................... 195

Ecology and Biology ................................................................................................................... 196

I Zoogeographic Distribution ........................................................................................................ 196

Depth Distribution ....................................................................................................................... 197

Materials and Methods ................................................................................................................ 197

I Aknowledgements .......................................................................................................................

198

Abbreviations Used in Keys and Figures .................................................................................... 198

Glossary ....................................................................................................................................... 198

i List of Species 202

: Key to Families ........................................................................................................................... 203

•m Key to Genera and Species .......................................................................................................... 204

i I Species Descriptions ................................................................................................................... 207

i Suborder Apseudomorpha Sieg, 1980 ......................................................................................... 207

Family Apseudidae Leach, 1814 .................................................................................... 207

I Carpoapseudes caraspinosus Dojiri and Sieg, new species 207

Suborder Tanaidomorpha Sieg, 1980 .......................................................................................... 210

Family Tanaidae Dana, 1849 ......................................................................................... 210

U Zeuxo maledivensis Sieg, 1980 ............................................................ 210 Family Leptocheliidae Lang, 1973 ................................................................................ 213

Leptochelia dubia (KrOyer, 1842) ........................................................ 213

I Family Paratanaldae Lang, 1949 .................................................................................... 218 Paratanais intermedius Dojiri and Sieg, new species ......................... 218

Family Anarthruridae Lang, 1971 .................................................................................. 220

I SubfamilySiphonolabrum Anarthrurinae californiensis Lang, 1971 ............................................................... Dojifi and Sieg, new species ............... 222200

Subfamily Akanthophoreinae Sieg, 1986 ......................................................... 224

i PaParalep raleptogna tognathia thia cbisetulosa _.gracilis Dojiri (Kr0yerand, 1842) Sieg, ...................................... new species ................. 224 226

Scoloura phillipsi Sieg and Dojiri, 1991 .............................................. 228

i AChauli raphuopleona ra breviaria dentataDojiri Dojiri andandSiegSieg, , newnespecie w specs ies............................... .......................... 231 234

Araphura cuspirostris Dojiri and Sieg, new species ........................... 236

U Tanaella Male specpropinquus ies 1 ...................................................................................... Dojiri and Sieg, new species ............................. 239 242

Male species 2 ...................................................................................... 244

,n SubfamilyMaleLeptspecies ognathiinae 3 ...................................................................................... Sieg, 1973 ............................................................. 244 247

Leptognathia cf. breviremis (Lilljeborg, 1864) .................................... 247

i Tanaopsis cadieni Sieg and Dojiri, 1991 ............................................. 249
!

Family Typhlotanaidae Sieg, 1984 ................................................................................ 253 I

Typhlotanais wiUiamsae Dojiri and Sieg, new species ....................... 253

Typhlotanais crassus Dojiri and Sieg, new species ............................. 256 i

Family Pseudotanaidae Sieg, 1973 ................................................................................ 258

Pseudotanais makrothrix Dojiri and Sieg, new species ....................... 258

Pseudotanais californiensis Dojiri and Sieg, new species ................... 260 I

Literature Cited ........................................................................................................................... 265

APPENDIX..................................................................................................................................... 269

!

!

I

i

I

I

!

I

!

!

!

I

!

I
!

I List of Figures

I Figure 1.1. Basic isopod morphology. Dorsal aspect of a female TridentelIa glutacantha, a flabelliferan

(left), and Ianiropsis tridens, an asellotan (right) .............................................................. 3

i Figure 1.2. Nome (B, C,nF, clature H); Cirolanidae of isopod cephalon (D, E, G,(A). 1) ................................................................................. Examples of isopod mouth appendages: Idoteidae4

Figure 1.3. Apanthura californiensis Schultz, 1964. Holotype LACM 55-35.1 (female). California,

I Los Angeles Co., off Santa Monica Pier, mud, 73 m. 06 February 1955. Coll. R/V "Velero

IV", AHF 2988-55 ............................................................................................................ 12

I Figure 1.4. Cyathura material LACM munda 48-62.3 Menzies,(mancas). 1951. BasedManca onMen paratype zies' holotype material drawi fromnCalifornia, g (male) andMarin paratype Co.,

Tomales Bluff, intertidal. 23 May 1948. Coll. R. J. Menzies, AHF 48-621 .................. 14

1.5. Menzies and 1959. Female. California, San Figure Haliophasma geminata Bamard, Diego Co.,

Oceanside. 20 February 1957. Coll. R/V "Velero IV", AHF 4868-57 .......................... 16

Figure 1.6. Haliophasma geminata Menzies and Bamard, 1959. Male. California, San Diego Co.,

II Oceanside. 20 February 1957. Coll. R/V "Velero IV", AHF 4868-57 .......................... 17

Figure 1.7. Hyssura sp. A. 014-BRC-03-TX, USNM Phase 1primary voucher collection ............. 20

i Figure 1.8. Paranthura elegans Menzies, 1951. Paratype LACM 48-62.2. California, Matin Co.,

Tomales Bluff, intertidal. 23 May 1948. Coil. R. J. Menzies ........................................ 23

I Figure 1.9. Munidion pleuroncodis Markham, 1975. Holotype USNM 141597 (female). Mexico, off

Baja California, 26°22'N, 115°05'W. 19-21 November 1966. Coll. W. C. Klawe, R/V

"David Start Jordan", Sta. 69 ........................................................................................... 26

I Figure 1.10. Munidion pleuroncodis Markham, 1975. Allotype USNM 141598 (male). Mexico, off

Baja California, 26°22'N, 115°05'W. 19-21 November 1966. Coll. W. C. Klawe, R/V

I "David Starr Jordan", Sta. 69 ........................................................................................... 27

Figure 1.11. Rocinela angustata Richardson, 1904. Syntype USNM 22710 (female). Alaska, Bering

Sea, N. W. of Unimak Island ........................................................................................... 32

I Figure 1.12. Metacirolanajoannae (Schultz, 1966) ............................................................................ 35

Ilr Figure 1.13. Serolis carinata Lockington, 1877. California, San Diego Co., Pt. Loma, 47 m. 20 July

1 1989. Coll. Pt. Loma Biology Lab, Sta. A-14, SDNHM A.0014 .................................... 37

Figure 1.14. Tridentella glutacantha Delaney and Brusca, 1985. Holotype CAS IZ025948 (male).

i California, 1953. Coll.LOs R/VAngeles "Velero Co., IV", oSta. ff LOs 2413-53 Angeles.................................................................... breakwater light, loose rock, 320-360 m. 39

Figure 1.15. Tridentella glutacantha Delaney and Brusca, 1985. Allotype LACM 53-113.1 (female).

I California, LOsAngeles Co., Santa Catalina Island, loose rock bottom, 304-310 m. 18 May

1941. Coll. R/V"Velero III", Sta. 1323-41 ..................................................................... 40

I Figure 1.16. Gnathia outfall. 05crenulatifrons October 1989. Monod, Coll. Pt. 1926. LomaMBiology ale. Californi Lab, aSta., SanA-5,Diego, SDNHMCo.,A.0114 off Pt. ..... Loma44

I
!

Figure 1.17. Gnathia productatridens Menzies and Barnard, 1959. Holotype AHF 5712 (male). i

California, Santa Barbara Co., off Santa Barbara Pt. Light, green silt, 89 m. 03 July 1957.

Coll. R/V "Velero IV", Sta. 5173-57 .............................................................................. 45

i

Figure 1.18. Gnathia sanctaecrucis Schultz, 1972. Holotype AHF 5927 (male). California, Channel

Islands, Santa Cruz Channel, green sand, 201 m. 22 December 1959. Coll. R/V "Velero

IV", Sta. 6805-59 ............................................................................................................. 46 H!11

Figure 1.19. Gnathia tridens Menzies and Bamard, 1959. Holotype AHF 5711 (male). California,

Santa Barbara Co., off Santa Barbara Point Light, 16 m. 17 January 1957. Coll. R/V m

"Velero IV", Sta. 4822-57 ................................................................................................ 48 !

Figure 1.20. Gnathia tridens Menzies and Barnard, 1959. Paratype AHF 5711 (female). California,

Santa Barbara Co., off Santa Barbara Point Light, 16 m. 17 January 1957. Coll. RN I

"Velero IV", Sta. 5164-57 ................................................................................................ 49 g

Figure 1.21. Idarcturus aIlelomorphus Menzies and Barnard, 1959. Holotype AHF 5713. California,

Santa Barbara Co., off Goleta, medium-coarse gray sand, 17 m. 09 April 1957. Coll. R/V |

"Velero IV", Sta. 4938-57 ................................................................................................ 51

Figure 1.22. Idarcturus allelomorphus Menzies and Barnard, 1959. California, San Diego Co., off Bird 'i

Rock, 32°49.25'N, 117°19.60'W, sandy silt, 60 m. 19 October 1989. Coll. Pt. Loma m

Biology Lab, Sta. B-5, SDNHM A.0014 ......................................................................... 52

i

Figure 1.23. ldotea rufescens Fee, 1926. Female. Mexico, Baja California, Coronado Is. 08 November B

1946. Coll. C. C. Hubbs, H46-118. LACM 46-57.1 ...................................................... 55

Figure 1.24. Synidotea calcarea Schultz, 1966. Holotype AHF 6833-60 (female). California, Channel i

Islands, Tanner Canyon, San Clemente Island, green muddy sand, 792 m. 19 January 1960. I

Coll. R/V "Velero IV", Sta. 6833-60 ............................................................................... 57

Figure 1.25. Synidotea media Iverson, 1972. Holotype CAS 563 (female). California, Monterey Co., i

offPt. Soberanes, 183 m. July 1971. Coll. R/V "Searcher". ......................................... 58

Figure 1.26. Stenetrium sp. A. Copulatory male from Phase I station 020-BRA. A, dorsal view, scal bar

= lmm. B, right pereopod I. C, right pleopod II ............................................................ 70 !11

Figure 1.27. Munna sp. A. A-B, brooding female from Phase I station 014-BRC, lateral and dorsal i

views, scale bar = 1 ram. C, fight uropod, lateral view, brooding female. D, pleopod I, |

ventral view, male from Sta. 004-BRA ............................................................................ 72

Figure 1.28. Munnogonium cf. tillerae. A & C, male (from Phase II, Cr.2-3, sta.R-4). A, dorsal view, I

scale bar = 0.5 mm. B, left pereopod I, brooding female (LACM cat. no. 56-26.2). C, fight iN

pereopod II ....................................................................................................................... 74

Figure 1.29. Pleurogonium caIiforniense Menzies, 1951. A, Dorsal view of hermaphroditic brooding D

female, Phase II station R-4, scale bar = 0.5 mm. B, left antennula. C, left mandible, dorsal

view .................................................................................................................................. 77

n

Figure 1.30. Pleurogonium sp. A. A-B, Dorsal view of hermaphroditic brooding female, Phase II station N

R-4, scale bar = 0.5 mm. C, left antennula. D, left mandible, dorsal view ..................... 79

Figure 1.31. Janiralata sp. A., preparatory female, MMS Phase I station 017-BRA. A, dorsal body R

view, scale bar = 1 mm. B, left antennula. C, left pereopod I, with enlargement of opposing w

margins of propodus and carpus ...................................................................................... 81

i

xiv I
!

I Figure 1.32. Janiralata sp. B, brooding female, MMS Phase I station 004-BRA. A, dorsal view, scale

bar = 1 mm. B, pereopod II with enlargement of proximoventral denticles on propodus ..

I .......................................................................................................................................... 83 Figure 1.33. Janiralata sp. C. A, dorsal view, preparatory female (MMS Phase I station 016-BRA),

pleotelson slightly damaged, scale bar = 1 ram. B-D, copulatory male (MMS Phase I

I station 016-BRA). B, pleotelson outline. C, right pereopod I, with enlargement of proximoventral spines on propodus. D, pleopod I .......................................................... 85

i Figure 1.34. Janiralata sp. D (cf. rajata), MMS Phase I station 016-BRA, preparatory female. A, dorsal

view of body, scale bar = 1 mm. B, fight pereopod I with enlargement of proximoventral

denticles on propodus, lateral view .................................................................................. 87

I Figure 1.35. Joeropsis concava Schultz, 1966, male from Phase I station 016-BRA. A, body dorsal view, scale bar = 1 mm. B, antennula. C, male pleopod I with enlargement of distal tip.

........................................................................................ , ................................................. 89

i Figure 1.36. Joeropsis sp. A. A-B, brooding female from Phase I station 006-BRA. A, body dorsal

view, scale bar = 1 mm. B, antennula. C, male pleopod I with enlargement of distal tip,

i Figure 1.37. Eurycope male specimen californiensis from sameSchultz, station1966. .................................................................................... A-C, copulatory male (Phase I sta. 50-BSS), dorsal 91

lateral views of body (scale bar = 1 mm), and enlargement of antennular basal article. D,

I preparatory female (Phase II Cruise 3-4, station R7), left uropod, ventral view ............. 93

Figure 1.38. Belonectes sp. A, preparatory female, Phase I station 072-BSS. A-B, body, dorsal and

lateral views, scale bar = 1ram. C, cephalon, lateral oblique view. D, right uropod, ventral

I view .................................................................................................................................. 95

Figure 1.39. llyarachna acarina Menzies & Bamard, 1959. A-B, body lateral and dorsal views, scale

I bar pleopod = 1 mm, II ventral preparatory view andfemale distalfromtip, MMS male fromPhasesame 11,Cruise station3-4, ....................................... Station R-5. C-D, male97

Figure 1.40. Munnopsurus sp. A. A-B, body dorsal and lateral views, scale bar = 1 mm, copulatory

i male, Phase II, Cruise 1-2, Station R9. C, right antennula. D-E, pleopod I,

male ventral

view with enlargement of distal tip. F, left uropod, ventral view ................................... 99

I Figure 1.41. Munnopsurus damaged, scalesp.barB. =A,1cmm. opulatoB,ry left maleantennula. (Phase I, Sta. C, male 063-BSS), pleopoddorsal I, ventral view, pleside, otelson and

enlargement of distal tip. D, fight uropod, ventral view ............................................... 101

I 1.42. Nannonisconus Schultz, 1966. Manca 3 Phase I station 081- Figure latipleonus stage specimen,

BSS, body length 1.5 mrn, scale bar = 0.5 mm. A, body, dorsal view. B, body, lateral view.

C, enlargement of antennula and base of antenna, in situ on head ................................ 103

I Figure 1.43. Momedossa symmetrica (Schultz, 1966), copulatory male (NAVY 103 station). A-B, body,

dorsal and lateral views, scale bar = 1 ram. C-D, pereopods I and 17Ifight, , lateral view. E,

I uropod, fight, ventral view ............................................................................................. 105

Figure 1.44. Desmosoma sp. A, Brooding female, dorsal view, composite illustration of several specimens

(Phase I station 014-BSS), scale bar = 1ram. B, brooding female, lateral view, brood plates

I and mouth part detail omitted. C, fight antennula, dorsal view. D, fight uropod, ventral view ................................................................................................................................ 107

!

I XV
!

Figure 1.45. Prochelator sp. A. A, brooding female (Phase I station 022-BSS), body dorsal view, scale !

bar = 1 mm. B-C, copulatory male, body lateral and dorsal view. D-E, pereopods I-II,

right, copulatory male (Phase II, Cruise 3-4 Station R-5). F, uropod, tip of pleopod I and lit

right pleotelson posterolateral spine, copulatory male .................................................. 109 I

Figure 2.1. Representative cumaceans from various families likely to encountered in the Santa Maria

Basin Region: A, Lampropidae, mature male with elongate antenna 2; B, Lampropidae, n

female with eggs in brood pouch; C, Lampropidae, mature male with grasping antenna 2; g

D, Diastylidae, subadult female; E, Diastylidae, subadutt male; F, Nannastacidae, subadult

female; G, Nannastacidae, mature male; H, Leuconidae, subadult female ................... 122 D

i

Figure 2.2. Side view of the cumacean body with details of the seven anterior head and thoracic

appendages. A1, antenna 1; A2, antenna 2; Md, mandible; Mxl, maxilla 1 (maxillule); l

Mx2, maxilla 2 (maxilla); Mxpl, maxilliped 1; Mxp2, maxilliped 2; P1, pereopod 1; P5, I

pereopod 5; PL1, pleonite 1; PL6, pleonlte 6; T4, thoracic somite 4 (=pereonite 1); T8,

thoracic somite 8 (=pereonite 5); af, accessory flagellum; an, antennal notch; b, basis; br, II

branchial apparatus; c, carapace; co, coxa; ex, exopod; fl, frontal lobe; ip, incisor process; i

lm, lacinia mobilis; mf, main flagellum; mp, molar process; o, oostegite; pa, palp; pl,

i

pseudorostral lobe; s, siphon; t, telson; u, uropod (from Watling, 1979) ...................... 123 i

Figure 2.3. Detailed views of selected cumacean head and anterior thoracic appendages. A, antenna 1 i

from female Leuconidae; B, antennae 1 and 2 from mature male Diastylidae (note enlarged

peduncle article 5 and large setal brush on antenna 2); C, antenna 2 modified for grasping, i

from male of Lampropidae; D, antennae 1 and 2 from mature male of Leptostylis (note !1

enlarged antenna 1and setal brush on peduncle article 3); E, antenna 2 of female Diastylidae;

F, mandible typical of most families; G, mandible from Campylaspis; H, a typical maxilla

1; I, a typical maxilla 2; J, maxilliped 1 with details of gill lobules from Eudorella; K, II

endopod of maxilliped 1; L, maxilliped 2 of Diastylis; M, maxilliped 3 of Diastylis; N,

maxilliped 3 of Campylaspis .......................................................................................... 124

II

Figure 2.4. Detailed views of pereopods, uropods, and telons. A, pereopod 1 from Diastylidae; B,

pereopod 2 from Diastytidae; C, pereopod 3 from Diastylidae; D, uropod and telson from

Lampropidae; E, uropod and telson from Diastylidae; F, uropod from Nannastacidae (telson i

fused to pleonite 6) ......................................................................................................... 126 1l

Figure 2.5. Diastylis sentosa Watling and McCann, new species. Female: A, carapace and pereonites, i

lateral view; B, body, lateral view; C, carapace and pereonltes, dorsal view; D, telson and II

uropods. Scale, A-C = 1 mm, D = 0.5 mm .................................................................... 131

Figure 2.6. Diastylis creneUata Watling and McCann, new species. Female: A, carapace and pereonites, H

lateral view; B, same, dorsal view; C, maxilliped 3; D, pereopod I; E, telson and uropods. i

Juvenile: F, telson and uropods. Scale, A,B = 1 mm, C-F = 0.5 mm............................ 132 i

Figure 2.7. Diastylis quadriplicata Watling and McCann, new species. Subadult male: A,carapace and D

pereonites, lateral view; B, body, lateral view; C, carapace and pereonites, dorsal view; D,

telson and uropod. Scale, A-C = 1 mm, D = 0.5 mm..................................................... 134

Figure 2.8. Diastylis santamariensis Watling and McCann, new species. Female: A, carapace, lateral m

view; B, body, lateral view; C, carapace and pereonites 1-2, dorsal view; D, E, telson and

uropods. Scale, A-D = 1 mm, E = 0.5 mm ..................................................................... 136 i

!1

!

xvi i
!

I Figure 2.9. DiastyIis californica Zimmer 1936. Female: A, carapace and pereonites, dorsal view; B,

same, lateral view; E, antenna 1; F, maxilliped 3; H, telson and uropods. Juvenile: C, carapace,

i dorsal view. Male: D, body, lateral view; G, telson and uropod. From Zimmer (1936); no scale provided ................................................................................................................. 137

Figure 2.10. Diastylis pellucida, Hart 1930. Male: A, carapace and pereonites, lateral view; B, body,

I lateral view; C, carapace and pereortites, dorsal view. Scale = 1 mm ........................... 138

Figure 2.11. Leptostylis calva Watling and McCann, new species. Female: A, body, lateral view; B,

i carapace and pereonites, lateral view; C, same, dorsal view; D, teslon and uropods. Male: E, telson and uropods. Scale, A-C = 1 rnm, D,E = 0.5 mm ........................................... 140

Figure 2.12. Leptostylis abditis Watling and McCann, new species. Female: A, body, lateral view; B,

I carapace and anterior pereonites, lateral view; D, carapace and pereonites, dorsal view; E, telson and uropods. Male: C, carapace and antenna 1, dorsal view. Scale, A = 0.5 mm, B-D

= 0.2 ram, E = 0.5 mm ................................................................................................... 141

m Figure 2.13. HemilampropscalifornicusZimmer 1936. Female: A, carapace and pereonites, lateral view;

B, telson and uropods. Scale, A = 1 mm, B = 0.5 mm ................................................... 143

I Figure 2.14. Alloeoleucon view; B, antennae santamariensis 1 and 2; C,Watling maxilliped and McCann, 3; D, pereopod new species. 1; E, pereopod Male: A,2;body, F, uropods. lateral

Female: G, antenna 1; H, maxilliped 3; I, pereopod 1; J, uropods. Scale, A, C-J = 0.5 ram,

I B=0.1 mm ..................................................................................................................... 146

Figure 2.15. Leucon (Diaphonoleucon) declivis Wading and McCann, new species. Subadult male: A,

body, lateral view; B, antenna 1; C, pleonites 1 and 2 with reduced pleopods; D, uropods; E,

I first pleopod pair. Scale, A = 1.0 mm, B-D = 0.5 ram, E = 50 pm ................................ 148

Figure 2.16. Leucon (Crymoleucon) bishopi Bacescu, 1988. Subadult male: A, pereopod 1; B, pereopod

2; C, uropods. Scale, A-C = 0.5 mm .............................................................................. 150

Figure 2.17. Leucon (Leucon)falcicosta Wafting and McCann, new species. Male: A, carapace, lateral

view; B, body, lateral view; C, carapace and pereonites, dorsal view. Female: D, antenna 1;

I E, maxilliped 3; F, pereopod 1; G, pereopod 2; H, uropod. Scale, = ram, B-C = 1 ram,

A 0.1

D,E = 0.25 rmn, F-H, 0.5 mm ........................................................................................ 152

I Figure 2.18. Leucon view; B,(Leucon) body, latearmatus ral view.Given, Scale =1961. 1 mmFemale: .................................................................... A, carapace and pereonites 1-3, lateral 153

Figure 2.19. Leucon (Leucon) magnadentata Given, 1961. Female: A, carapace and anterior pereonites,

I lateral view; B, lateral view; C, and anterior dorsotateral view.

body, carapace pereonites,

Scale = 1 mm .................................................................................................................. 154

I Figure 2.20. Eudorella carapace and pacifica anteriHart, or pereonites, 1930. Female: dorsalA,view; carapace, D, calateral rapceview; and B, pereonites, body, lateral lateral view; view. C,

Scale A, C = 1 mrn, B,D = 1 mm .................................................................................. 155

I Figure 2.21. Eudorella redacticruris Watling and McCann, new species. Female: A, and carapace

pereonites, lateral view; B, antenna 1; C, pereopod 2; D, uropods. Male: E, pereonite 5 with

penial lobes and pleonites 1 and 2 with appendages; F, ventral view of penial lobes. Scale,

I A= 1 mm, B-E = 0.5 mm, F=0.1mm 157

!

I' I xvii
!

Figure 2.22. Eudorella truncatula Bate, 1856. Female: A, carapace and pereonites 1-4, lateral view; B, i

body, lateral view; C, D, uropods. Scale, A, B = 1 mm, scale C = 0.1 mm, D = 0.5 mm ...

........................................................................................................................................ 158 I

Figure 2.23. Eudorellopsis longirostris Given, 1961. Subadult male: A, carapace and pereonites, lateral

view; B, body, lateral view; C, carapace, dorsal view. Subadult female: D, body, lateral

view. Scale A, B, D = 1 mm, C = 0.1 mm ..................................................................... 159 R

Figure 2.24. Campylaspis canalicuIata Zimmer, 1936. Male: A, carapace and pereonites 1-2, lateral

view; B, body, lateral view; H, uropods. Female: C, body, lateral view; D, maxilliped 3; E, iI

pereopod 1; F, pereopod 2; G, uropods. Scale A, B = 1 mm, G, H = 0.5 mm (C-F from |

Zimmer, 1936; no scale provided) ................................................................................. 162

Figure 2.25. Campylaspis rufa Hart, 1930. Female: A, carapace and pereonites, lateral view; B, body, i

lateral view; C, carapace, dorsal view; D, uropods. Scale A-C = lmm, D = 0.1 mm ... 163 I

Figure 2.26. Campylaspis rubromaculata Lie, 1971. Subadult female: A, body, lateral view; B, carapace II

and anterior pereonites, lateral view; C, carapace, dorsal view; E, uropod. Scale, A-C, 1.0 il

ram, D =0.5 mm ............................................................................................................ 165

Figure 2.27. Campylaspis maculinodulosa Watling and McCann, new species. Male: A, body, lateral I

view; B, carapace and pereonites, lateral view; F, uropods. Female: C, carapace and i

pereonites, lateral view; D, maxilliped 3; E, uropods. Scale, A= 0.1 mm, B,C = 0.2 mm, E

= 0.5 mm ........................................................................................................................ 166 i

g

Figure 2.28. CampyIaspis hartae Lie, 1969. Female: A, carapace and pereonites, lateral view; B, body,

lateral view; C, carapace, dorsal view; D, uropods. Scale, A-C = 1 mm, D = 0.5 mm. 167 ii

il Figure 2.29. Campylaspis blakei Wafting and McCann, new species. Female: A, carapace and pereonites,

lateral view; B, body, lateral view; C, uropods; D, carapace, dorsal view ; E, pereonites 1-

5 and pleonites 1-4, dorsal view; F, uropod. Scale, A-E = 1 mm, F = 0.5 mm ............. 169 H

g

Figure 2.30a. CampyIaspis bipIicata Wading and McCann, new species. Female: A, carapace and

pereonites, lateral view; B, body, lateral view. Male: C, body, lateral view; D, carapace, n 

dorsal view; E, same, lateral view. Scale, A = 0.2 mn_ C-E = 0.3 ram, B = 0.5 mm ... 171 i

Figure 2.30b. Campylaspis bipIicata Watling and McCann, new species. Female: F, maxilliped 3; G,

pereopod 2; H, uropod. Male: I, uropod. Scale, H, I = 0.5 mm .................................... 172 B

i

Figure 2.31. Procampylaspis caenosa Wafting and McCann, new species. Female: A, Carapace and

anterior pereonites, lateral view; B, body, lateral view; C, carapace, dorsal view; D, maxilliped IBm

2; E, uropods. Scale, A-C = 1 mm, E = 0.5 mm ............................................................ 173 B

Figure 2.32. Cumella (Cumella) morion Watling and McCann, new species. Female: A, carapace,

pereonites, and anterior pleonites, lateral view; B, body, lateral view; C, carapace,

anterodorsal view; D, uropods. Scale, A,C, D = 0.2 mm, B = 1 mm ............................. 175 II

Figure 2.33. Cumella (Cumella) californicaWatling and McCann, new species. Female: A, carapace i

and pereonites, dorsolateral view; B, pereonites and pereopods 1-3; C, uropods. Scale, A = !

1 mm, B,C = 0.2 mm ...................................................................................................... 176

w

Figure 3.1. Generalized habitus of a tanaidomorphan: top, dorsal view; bottom, lateral view ........ 183 i

I

Figure 3.2. Generalized f'rrst antenna of female and male tanaidomorphan ..................................... 185

I

xviii i
!

I Figure 3.3. Generalized tanaidomorphan second antenna, labmm, fight and left mandible, first maxilla,

and second maxilla ......................................................................................................... 187

I Figure 3.4. Generalized tanaidomorphan labium, maxilliped, and epignath ................................... 189

Figure 3.5. Generalized tanaidomorphan cheliped and pereopod .................................................... 191

I Figure 3.6. Generalized tanaidomorphan pleopod and representative examples of uropods ........... 194

Figure 3.7. Carpoapseudes caraspinosus Dojiri and Sieg, new species female: Q, general habitus of

I female; A. 1,fLrst antenna; Mxp, maxilliped; Che, cheliped; P.1, pereopod 1 ............... 209

Figure 3.8. Zeuxo maledivensis Sieg, 1980 male: C_, general habitus of male; Md, mandible; Mxp,

maxilliped; O_ Che, cheliped; P. 1, pereopod 1;P.6, pereopod 6; P1,pleopod; Uro Of, umpod

i of male; neuter, general habitus of neuter; Uro, uropod of neuter ................................. 212

Figure 3.9. Leptochelia dubia (Kr0yer, 1842) female: _, general habitus of female; A. 1, first antenna;

I A.2, second antenna; Md, mandible; Mxp, maxilliped; Che, cheliped; P. 1, pereopod 1; P.6, pereopod 6; Uro, uropod ................................................................................................ 215

II Figure 3.10. Leptochelia dubia (Kr_yer, 1842) male: _, general habitus of male; A.1, first antenna;

II C_1, flagellar segments of first antenna (variant); A.2, second antenna; Mxp, maxilliped;

Che, cheliped; P.1, pereopod 1; P.6, pereopod 6; Uro, uropod ...................................... 216

I Figure 3.11. Paratanais intermedius Dojiri and Sieg, new species, female: Q, general habitus of female; A. 1, fwst antenna; A.2, second antenna; Md, mandible; Mxp, maxilliped; Che, cheliped;

P.1, pereopod 1; P.6, pereopod 6; Uro, uropod .............................................................. 219

I Figure 3.12. Siphonolabrum californiensis Dojiri and Sieg, new species, female: Q, general habitus of

female; A.1, first antenna; L, labrum; Md, mandible; Mxp, maxilliped; Che, cheliped; P.1,

I pereopod 1; P.6 pereopod 6; Uro, uropod ...................................................................... 222

Figure 3.13. Siphonolabrum californiensis Dojiri and Sieg, new species, male: C_, general habitus of

male; A.1, first antenna; Mxp, maxilliped; Che, cheliped; P. 1, pereopod 1; P.6, pereopod 6;

I PI, pleopod; Pit, pteotelson; Uro, uropod 223

Figure 3.14. Paraleptognathia cf. gracilis (Krcyer, 1842), neuter: neuter, general habitus of neuter; A. 1,

I first Uro,antenna; uropod .................................................................................................................... Md, mandible; Che, cheliped; P.1, pereopod 1; P.6, pereopod 6; Pit, pleotelson; 225

Figure 3.15. Paraleptognathia bisetulosa Dojiri and Sieg, new species, female: Q, general habitus of

I female; A. first mandible; Che, P. 1 P.6, 6;

1, antenna; Md, cheliped; 1, pereopod pereopod

Pit, pleotelson; Uro, uropod ........................................................................................... 227

I Figure 3.16. Scoloura antenna; Md, phillipsi mandible; Sieg andChe, Dojiri, cheliped; 1991,P.1, female: pereopod Q, general 1; P.6,habitus pereopodof female; 6; Uro,A.uropod. 1, first

Neuter: N, general habitus of neuter .............................................................................. 230

I Figure 3.17. Chauliopleona dentata Dojiri and Sieg, new species, female: (_, general habitus of female;

Q A. 1,fast antenna of female; Md, mandible; Che, cheliped; P.1, pereopod 1; 9 1, terminal

I segment general habitus of pereof opodsubadult 1 of nemale; uterssubadult and some@A. variant 1, fin-st females; antennaUro, of subadult uropod. male Subadul .......t 232 C_,

!

|
BiIIIlIiIiI!!!iiIIi
!

I 1.0 THEORDERISOPODA

i 1.1 INTRODUCTION TO THE MARINE ISOPODA

by

I Regina Wetzer 1,Richard C. Brusca l,

and George D.F. Wilson 2

l

i Introduction

The order [sopoda is one of nine orders in the crustacean superorder Peracarida. Peracarids are

I dcombination istinguishedofromcharacters: other eumalacostracan absence of caudalsuperor rami donersthe(Hoplocarida, telson; maxillipedal Syncaridbasis a, Eucafid typicallya) byproduced the follointowinang

anteriorly directed, bladelike endite; mandibles with an articulated accessory process in adults, between the

molar and incisor processes (the lacinia mobilis); carapace, when present, not fused with posterior pereonites

I and usually reduced in size; females with unique thoracic coxal endites, calledoostegites, that form a ventral

brood pouch, or marsupium (except in Thermosbaenacea); and, in most groups, young hatching as mancas (a

I pre-juvenile hence true larval stageforms lackingdotnot he last occur pairinofthisthoracopods). superorder. All About peracarids 12,000uspecies ndergo ofdirect peracarids developmen havetbeen withdescribed. brooding,

Although we have tried to keep the amount of terminology in this chapter to a minimum, use of certain

I destandard tailed dcrustacean escription ofterminol isopodogymorispholog unavoidable. y can be Mfound ost oinf tBru hesescaterms and are Wilsdefined on (1991). below. An extensive and

Synonymies are not provided, but they can be found in the key references listed for all taxa. Several

I general guides to marine isopods of North and middle America have been published. These are: Richardson, 1905 (still a highly valuable reference, although obviously rather out-of-date), Menzies and Frankenberg,

1966, Schultz, 1969 (covers all of North America; many errors - use with caution), Miller, 1975 (good key to

I common intertidal California species), Brusca, 1980 (keys to Gulf of California fauna), Brusca and Iverson,

I985 (only summary treatment available for the tropical eastern Pacific region), Kensley and Schotte, 1989

(excellent, with keys and figures of Caribbean and many Gulf of Mexico species, supersedes Menzies and

I Glynn, northwest196Pacific 8). Twodeep-water other general monograph works are also useful. The first is an English translation of Birstein's (Birstein, 1973). The second is a three volume compendium on

temperate northern hemisphere marine isopods by Kussakin (1979, 1982a, 1988). These latter works are in

i Russian and compiled entirely from the literature. The California marine isopod fauna numbers about 176 valid named species, representing 30 families

in 8 suborders. The U. S. Department of the Interior, Minerals Management Service (MMS), soft-bottom

I benthic macroinvertebrate faunal surveys in the Santa Mafia Basin and the Western Santa Barbara Channel

• (hereafter referred to as the "MMS survey") recovered 37 species, 14 of which are undescribed. Where

possible, type material was examined and illustrated for the present study (indicated in the figure legends).

!

I _Grice Marine Biological Laboratory, University of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412

2Division of Invertebrate Zoology, Australian Museum, 6 College Street, Sydney South, New South Wales 20130,Ausmdia

,II
1

Illustrations were prepared by John Simpson, Frances Runyan, and the authors. This work was accomplished I

with the assistance of funding from the MMS, and grants to R. C. Brusca and G. D. Wilson from the National

Science Foundation. i

Summary of Isopod Research in Northeast Pacific Region I

The earliest studies on northeast Pacific marine isopods are recorded in the publications of Dana

(1853, 1854), Lockington (1876, 1877), Benedict (1897, 1898a-b), and Boone (1918, 1923). However, itwas i

the pioneering 25-year systematic research program of Harriet Richardson that laid the foundation of our I

modem knowledge regarding this fauna (Richardson, 1897, 1898, 1899a-b, 1900, 1904a-b, 1905a-b, 1906,

1908, 1909). Miller, Schultz, and especially Menzies built on Richardson's foundation through the 1950's, 1

1960's, and early 1970's (Miller, 1940, 1941, 1968, 1975; Miller and Lee, 1970; Miller and Menzies, 1952; i

Menzies 1950a-b, 1951a-e, 1952a-b, 1954a-b, 1958, 196I, 1962; Menzies and Bamard, 1951, 1959; Menzies

and Bowman, 1956; Menzies and Miller, 1972; Menzies and Mohr, 1952, 1962; Menzies, Mohr, and Wakeman, I

1963; Menzies and Waidzunas, 1948; Menzies and Widrig, 1955; Schultz, 1964, 1966, 1972, 1973, 1977; m

Bowman and Schultz, 1974; Sassaman, Schultz, and Garthwaite, 1984). Research on northeast Pacific isopods

by R. Brusca and G. D. Wilson began in the mid- 1970's and continues today (Brusca, 1977, 1978a-b, 1980, _l

1981, 1983a-b, 1984, 1987; Brusca and France, 1992; Brusca and Gilligan, 1983; Brusca and Iverson, 1985; U

Brusca and Ninos, 1978; Brusca and Wallerstein, 1977, 1979a-b; Brusca and Weinberg, 1987; Brusca and

Wilson, 1991; Brusca et al., 1995: Bruce, et aI., 1982; Cadien and Brusca, 1993; Delaney and Brusca, 1985; Ii

Guzman, et aI., 1988; Perry and Brusca, 1989; Stepien and Brusca, 1985; Thun and Brusca, 1980; Wallerstein II

and Brusca, 1982; Wetzer, etal., 1987; Wilson, 1976, 1982a; Wilson, etal., 1989).

!

General Isopod Anatomy ilil

The order Isopoda Latreille, 1817, is distinguished from the other eight ordersof Peracaridaby the

following combination of characters (see Figures 1.1 and 1.2): l

First thoracomere fused to cephalon, andrarely the second as well. Without acarapace (i.e. with only I

a cephalic shield). Body usually dorsoventrally depressed, cylindrical (tubular)in some suborders. Antennules

and antennae uniramous, but with a minute scale ("squama") in a few taxa. Eyes unstaLked but on lobes in I

some Asellota, Gnathiidea and Valvifera. Mandible usually with a 1- to 3-articulate palp and a multidentate II

incisor process; left and right lacinia often differ; molar process highly variable. Maxillule and maxilla both

without palps. Thoracic appendages uniramous (without exopods). First thoracopods modified as maxillipeds; I

with a short coxa and usually a short anteriorly-directed lamellar epipod; basis flattened and produced into II

bladelike anteriorly-directed endite, often with coupling spines; palp of up to 5 articles. Second thoracopods

modified as maxillipeds (pylopods) only in Gnathiidea. Pereopodal coxae small and simple, or more-or-less l

fused with body somites and forming lateral extensions on pereonites (coxal plates); line of dorsal fusion I

usually demarcated on pereonites ]/-VII (occasionally pereonite I). Pleon short, in many with various segments

fused; telson always fused with last pleonite to form a pleotelson. Pleopods biramous, flattened, specialized am

for respiration and/or swimming; endopods of second pair typically with stylets in males ("appendix I

masculinae"). Uropodal rami uniarticulate. Heart located primarily in pleon; usually with 2 pairs of ostia and

5pairs of lateral arteries. Maxillary glands usually present in adults. Young leave marsupium before appearance II

of last pair of pereopods (as "manca"). Isopods exhibit biphasic molting. I

I
1

1

I antennule _I

1

I 3
I

I

__,+ I

t

._ . / |

i.., ,,. I

coupling \,,,

t some l . i

J m_i,,_,_ \ \.. mood+,e

. -_ .... _-,_3_

_-f .._j_ coopl,°°+oe ""J II

• v ?S_.._. incisorprocess tacinia mob]lis _ palp,_/,.'_ II

_ <c ,,+ @-+\ cf

."" '_'\ / t \ maxilliped {.--( I

(•__.... _/" ,, \'.,+... "Q---J_){ =xa cexa molar process/ / 1

... ! ........}, _

( I f /

" El i •

I

""t maxi la,/ mandible ./

II

!l / '"

I

li_re 1.7.. Nomenclature of isopodcephalon (A). Examples of isopod mouth appendages: Idoteidae (B,C, F,

H); Cirolanidae (D, E, G, I). I

Brusca and Wilson (1991) recognize two basic isopod morphologies, "long-tailed" and "short-tailed". i

Long-tailed isopods have the telsonic region of the pleotelson greatly elongated, thus the anus and origin of the II

uropods are positioned basally on the pleotelson (Flabetlifera, Anthuridea, Gnathiidea, Epicafidea, Valvifera).

In the short-tailed isopods, the telsonic region is greatly reduced or vestigial, positioning the anus and uropods •

terminally or subterminally on the pleotelson (Phreatoicidea, Asellota, Microcerberidea, Oniscidea, I

Calabozoidea).

Isopods can be sexed in several ways. If oostegites, or marsupium, are present, one is obviously Ii

looking at a female. The openings of the oviducts in females (near the base of the legs on the 6th thoracomere) I

are extremely difficult to observe. If oostegites are absent, males can be distinguished by the presence of

paired penes on the sternum of pereonite VII (or pleonite 1) and/or the presence of appendix masculinae on the

endopods of the second pleopods. Absence of penes, appendix masculinae, and oostegites indicates the specimen I

is either a female or a juvenile that has not yet developed secondary sexual features.

I

I

4 I
!

I Glossary of Technical Terms

I aesthetase. Thin-wailed chemosensory seta usually distal. Situated away from the base or point of found on antennular flagellum, origin or attachment.

antennules. Antennae 1or first antennae; endite. A lobe on the inner (medial) margin of the

I anteriormost paired appendages of head (= protopod of an arthropod limb; enlarged

antennula), anteriorly directed lobe of the basis of an

I antennpaired ae. Antennae appendages 2 or seco of head. nd antennae; second endopod. isopodInner maxilliped. (medial) ramus of an appendage.

appendix maseulina. Copulatory stylet arising exopod. Outer (lateral) ramus of an appendage.

I from medial of mate 2

margin pleopod

endopod, used for transfer of spermatophores flagellum. Narrow distal part of antenna or

in at least some species. (P1.appendices antennule, usually multiarticulate,

I masculinae) occ without asionaintrinsic lly reducemusculature. d to one or a few articles,

biramous. Composed of two rami or branches.

frontal margin. Separates the upper surface of the

I earina. A keel, acute from the frontal lamina; anterior

or ridge. cephalon

margin of cephalon.

cephalon. Head; strictly speaking the cephalon of

I anfuseisdopowithd isthea cephal first thoothracoroaxmere , as (iant isdalways also frontalbases laminaof.theA antennae sternal plaantedarisi pronbably g between the

with the second thoracomere in the homologous to the epistome of other

I Gnathiidea). arthropo lamina may ds; inextend manyanteriorly isopods theto befrovisible ntal in

chelate. Having a chela; the propodus and dactylus the dorsal aspect; in many Valvifera the

i flatter ormingarticulates a pincerlike submedially structure onwherein the former the umay pperbeandseparately lower regio elevated ns of this and visible structurein the

to produce a "moveable and fixed finger" dorsal aspect as two distinct structures

arrangement. (True chelae are extremely rare (frontal lamina 1 and frontal lamina 2).

I in isopods.) frontal process. The lower region of the epistomal

coupling spine(s). When present, coupling spines plate or sclerite, which in some Valvifera

I (oandr hpleopodal ooks) occurpeduncles, on the maxandilli

serve pedaltoendites lock projects outward to become visible in the dorsal aspect as a protuberance lying above

the opposing appendages together, allowing the frontal lamina.

them to function as a single unit.

I gnathopod. A loosely used term usually referring

eoxa. Basal article of an appendage, to chelate or subchelate pereopods associated

i eoxal plat expande es. Cdoxae into offlattened the perelamellar opods laterally structures whandling. ith the heaThis d regtermionisandrarely usedused for f

for ood

extending freely (as "plates") to overhang the isopods.

i coxa-basis hinge of the leg. incisor process. Grasping, piercing or slicing elypeus. In most (but not all) isopods the labrum structure arising apically on body of

consists of two pieces; the anteriormost mandible.

I (proximal) piece is referred to as the clypeus,

labium. The posterior border of the buccal field;

the posteriormost (distal) piece as the labrum the "lower lip" (see paragnath).

proper.

!

i '
1

E

labium. Flaplike structure posterior to and borne palp. Articulated ramus consisting of one to three Ii

upon the frontal lamina or clypeus, usually articles on the mandible, and up to five

free but occasionally fused to head; the articles on the maxilliped. (Palps do not •

"upper lip" (see clypeus), occur on the maxillules or maxillae of I

isopods.)

laeinia mobilis. Small, usually toothed, process

articulating at base of incisor of left or both paragnath. In isopods, the labium is usually i[

mandibles; present in most isopod groups, produced and cleft into a large bilobed w

structure, and hence more commonly referred

lamina dentata. Serrate platelike structure on the I

mandible of anthurideans, presumed to have to

metastome,

as the paragnath hypopharynx). (= hypostome, 1

been formed by the fusion of spines of the

spine-row, peduncle. Enlarged proximal, or basalmost region !

of an isopod antennule, antenna, pleopod, or I

manea. Young of most peracarids (including uropod; contains intrinsic musculature.

isopods), lacking last (eighth) thoracic

appendages at time of release from penes. Paired (occasionally fused) submedian

broodpouch, processes from vas deferens on sternum of II

male pereonite VII or pleonite 1.(Sing.

mandible. Third pair of head appendages; first penis) I

(anteriormost) pair of mouthparts

I

functioning asjaws and typically sclerotized, pereon. Those thoracic segments not fused to the

marsupium. Structure in which eggs and embryos cephalon. _l

are retained and brooded by female; the pereonite. A segment of the pereon (= pereomere).

broodpouch. Isopod marsupia are typically

formed by overlapping medial plates pereopods. The paired legs of each pereonite. •

(oostegites) arising from certain pereonal pleon. Abdomen.

coxae in females; in a few groups the

oostegites have been reduced or lost in lieu of pleonite. A segment of the pleon (= pleomere). I

internal brooding, pleopods. The biramous, paired, lamellar II

maxillules. Fourth pair of head appendages, appendages of each pleonite. lit

functioning as mouthparts, immediately pleotelson. In all isopods the sixth pleonite is fused •

posterior to mandibles (= first maxillae), to the telson to form a pteotelson. In

maxillae. Fifth pair of head appendages, anthurideans the line of fusion is often visible l.

functioning as mouthparts, immediately as a deep dorsal groove or fold. I

posterior to maxillules (= second maxillae), praniza. Juvenile, immature stage of gnathiideans.

maxillipeds. Modified first pair of thoracopods, prehensile. Appendages (usually pereopods) 1

functioning as mouthparts, adapted for holding or clinging, the dactyl is

El

molar process. Grinding, or piercing or slicing as long or longer than the propodus, strongly 1

structure, arising mid-basally on body of developed and recurved. I

mandible; pars molaris, protandrous, in sequential hermaphroditic forms,

oostegite. Thin ventromedial plates, born on coxae becoming a functional male (producing D

of some pereopods in female peracarids spermatozoa) before becoming a functional I

forming the marsupium, or broodpouch, female (producing eggs).

I

6 !
!

I protogynous. Insequential hermaphroditic forms, statocyst. Small saclike sensory organ, usually

becoming a functionalfemale (producing containing a granule(s), used to indicate to

i eggs) before becoming a functional male the animal its orientation; present singly or (producing spermatozoa), paired on the pleotelson of some

anthurideans.

proximal. Situated near the base or point of

i attachment, subehelate. Having a subchela; forming a pincerlike structure by the dactylus folding

pylopods. Second pair of maxilhpeds in gnathiids back on the propodus.

i (appendages of the second thoracomere). rostrum. Anteromedial projection of frontal margin tergite. Dorsal sclerite of exoskeleton on arthropods.

of cephaton.

I scale. A small articulate piece occurring on the

uropods. The paired biramous appendages of the

isopod pleotelson, representing the

antennutes or antennae in some crustaceans, appendages of the fused sixth pleonite.

i thought to be a remnant of the second ramus (presumably the exopod). Scales rarely occur

in isopods.

I spine rothew.mola Spinose r andlobe incisonor the procmandible, esses. between

I Phyletic Key to the Suborders of Isopod (Adults)

i

I 1A. With only 5 pairs of pereopods (thoracomere 2 entirely fused to cephalon, with its appendages

[pylopods] functioning as a second pair of maxiUipeds; thoracomere 8 reduced, without legs); adult

i males with mandibles grossly enlarged, forceps-like, projecting in front of head; adultfemales without mandibles ......................................................................................................................... Cmath[idea

lB. With 7 pairs of pereopods (thoracomere 2 not fused to cephalon, with 1 pair of maxillipeds and 7

l pairs ofpereopods); males without projecting, forceps-like mandibles; females with mandibles .... 2

2A. Adults obligate parasites on other crustaceans; bilateral symmetry reduced or lost in females; male a

I small bilaterally symmetrical symbiont living on the body of the female; antennae vestigial; antennules

Ill

reduced to 3 or fewer articles; without maxiUules ............................................................ Epical-idea

2B. Not obligate parasites on other crustaceans; bilateral symmetry retained in both sexes; male not as

I above; antennae not vestigial; variable; usually ....................................

antennules with maxillules 3

I 3A. Body like) duecylindri to ventrally cal or tubular elongatedin cross-secti abdominalopleura; n, but with oftendistinct appearirowng lofaterally filter setae compressed along medial (amphipod- margin

of maxilla; penes located on coxae of male pereopod VII; apex of pleotelson curves dorsally; pleonite

I 5India) elongate, ............................................................................................................................. markedly longer than any other pleonites (known only from southern hemisphere Phreatoicidea and

3B. Body variable, but not appearing laterally compressed as above; without row of filter setae along

I medial of pleotelson margindoes of maxill not curve a; pedorsally; nes on sternum pleoniteof5male rarelypereonite elongateVII (markedly (or on sternum longer than of pleor other Litepleonites 1); apex

only in Limnoriidae) ......................................................................................................................... 4

,!
!

4A. Anus and articulating base of uropods positioned terminally (or subterminally) on pleotelson; uropods I

styliform (in the oniscidean family Tytidae the uropods are modified into flattened ventral plates that

cover the anal chamber) ................................................................................................................... 5 •

II 4B. Anus and articulating base of uropods positioned at base of pleotelson; uropods flattened ..............

................................................ * ................................................................................ , ....................... 8

!

5A. With lateral coxal plates; antenna peduncle 5-articulate; antennule reduced to 5 or fewer articles;

maxillipeds without coupling spines; penes of male arise from articulation between pereonite VII and m

pleonite 1; mandible without palp; pleopodal exopods broad and opercular to the thick tumescent I

endopods; female pleopod 1 present ................................................................................................. 6

5B. Without lateral coxal plates (pereopodal coxae small); antenna peduncle 6-articulate; antennule reduced •

or not reduced; maxillipeds with or without coupling spines; penes of male arise on sternum of I

pereonite VII; mandible with palp; pleopods not as above; female pleopod 1 absent ...................... 7

Ill

6A. Exoskeleton of pleonites 1 and 2 reduced to only sternal plates; uropodal rami fused to peduncle; I

blind groundwater animals (known only from ground waters in Venezuela) ................ Calabozoidea

6B. Pleonites 1 and 2 not reduced as above; one or both uropodal rami articulate on peduncle; with or i

without eyes ........................................................................................................................ Oniscidea

m

7A. Minute, usually less than 3 mm long; long and slender, length about 6 times width; antennal peduncle I

without a scale; antennule reduced, peduncle indistinguishable from flagellum; maxilliped without

coupling spines on endite; female pleopod 2 biramous; male pleopod 2 endopod not geniculate; J

interstitial ................................................................................................................. Microcerberidea

7B. Rarely minute, usually greater than 4 mm long; body not elongate (length less than 6 times width);

antenna] peduncle usually with a scale; antennule rarely reduced, peduncle and flagellum distinct;

maxilliped almost always with coupling spines on endite; female pleopod 2 uniramous; male pleopod II

2 endopod large and geniculate; rarely interstitial ................................................................. Asellota

i

8A. Body elongate, length usually more than 6 times width; uropodal exopod curving dorsally over

pleotelson; co×ae of maxillipeds fused to head (i.e. not freely articulating); mandible with unique E..

lamina dentata (in lieu of spine row and lacinia mobilis; lamina dentata, spine row and lacinia mobilis I

lacking in Paranthuridae); maxillule an elongate stylet with apical hooks or serrate margin; maxilla

vestigial and fused with paragnath (or absent) .................................................................. Anthuridea II

8B. Body not markedly elongate, length usually less than 4 times width; uropodal exopod not as above; I

coxae of maxillipeds not fused to head; mandible without lamina dentata; maxillule variable; maxilla

well developed, never fused with paragnath ..................................................................................... 9 I

9A. Uropods modified as a pair of ventral opereula covering the entire pleopodal chamber; males with

penes arising on sternum of pleonite 1, or on articulation between pereonite VII and pleonite 1;

mandibular molar process a stout, flattened grinding structure ........................................... Valvifera I

9B. Uropods not modified as ventral opercula covering pleopods; males with penes arising on sternum of

pereonite VII; mandibular molar process usually a thin, bladelike, cutting structure, or absent (flattened

only in Sphaeromatidae) .................................................................................................. Flabetlifera

I
i

!

I 1.2 DESCRIPTIONS OF THE SPECIES OF THE SUBORDERS

ANTHURIDEA_ EPICARIDEA_ FLABELLIFERA,

I GNATHHDEA, AND VALVIFERA

m by

I Regina Wetzer 3, and Richard C. Brusca 3

i Suborder Anthuridea Leach, 1814

Description. Body long, slender, subcylindrical; circular in cross-section; length 6-I5 times width.

I Antennules short except in males of some species, uniramous (without scale), usually with 3 peduncular • articles; some species with only 3 antennular articles, setation suggesting loss of 1peduncular article as well

as most flagellar articles. Antennae short, uniramous (without scale), peduncle 5-articulate, with few flagellar

I artic (the les. "lamina Manddentata"); ible without palpdistin of 1-3 ct lacinia articles,moorbilisabsent. or spiMaxillule ne row, instea withdinner usuallyramus withreduced, a dentateouter lobe ramus or platea

slender stylet with terminal spines (spines often reduced to a row of short dentictes). Maxillae rudimentary,

i ucoupling sually fused with paragnath. Maxillipeds more-or-less fused to head, usually with small endite; without

spines; patp of 1-5 articles; apex of palp acute or rounded. Pereonites mostly longer than wide (in

contrast to most isopods, in which the reverse is true); dorsum often with distinct ridges, grooves or

i chromatophore patterns; distinct coxal plates rarely evident. Pereopods I-III (especially pereopod I) tend

towards subchelate form; pereopods IV-VII generally ambulatory. Pleonites 1-5 free or fused; pleonite 6

fused with telson, but often demarcated from the telson by a deep dorsal groove or fluting; pleopods 1-5

similar, or pleopod 1 modified as an operculum. Uropods attached laterally at base of pleotelson, but exopods

I curve dorsally to arch over pleotelson; often with one or two telsonic statocysts.

Remarks. In all isopods the sixth pleonite is fused to the telson to form a pleotelson. In anthurideans

i the freesixth pleonites. pleoniteHowever, is oftenhistological distinctly demstudies arcatedhavebysoa dofarrs

failed al grootovereveal givinganytheanthuridean illusion of anspecies articuinlatwhich ion andthe6

telson freely articulates on the 6th pleonite.

I mouthpartsFourandfamilies pleon: Anthuridae of AnthurideaLeach, are currently 1814; Paranthuridae recognized,Medistinguished nzies and Glynn, prima1968; rily byHyssuridae charactersW_igele, of the

1981; and, Antheluridae Poore and Lew Ton, 1988. The family Anthuridae possess 1or 2 telsonic statocysts

I have (or noa statoeysts reduced number ), operculate of articles. first The pteopods, family and Paranthuridae broad maxilhas lipedal 0 or 1ptelsoni alps thcatstatocysts tend to beandopercu operculate liform f'rrst and

pleopods. Species of Hyssuridae have very long bodies (about 15times longer than wide), no telsonic statocysts,

I maxillipeda long free plleonites endite, and andlapalp ck operculif (W_igeleorm1981). pleopods. Anthuridae Antheluridae and Paraarenthuridae def'med have by thebeexencepti previously onal widthreported of the

from California waters. The MMS project also collected a new species of Hyssuridae.

!

I 3Grice Marine Biological Laboratory, University of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412

I 9
!

Anthurideans appear to be primarily carnivores, feeding on small worms and arthropods of various I

kinds. Most inhabit littoral or shallow shelf environments, although some deep benthic (and some freshwater)

species are also known. Many species are known to be protogynous, and males have not yet been reported for •

several of these. Fewer than 200 species of anthurideans have been named, but many more remain to be l

described.

Key References. Richardson, 1905; Miller, 1975; Poore, 1980; W_igele, 1981a-b; Poore and Lew I

Ton, 1988a-c; Kensley and Schotte, 1989; Cadien and Brusca, 1993. U

!

Key to the Families of Anthuridea Known from California Waters

1A. Mouthparts adapted for piercing and sucking, together forming a conelike structure; mandible usually I

with an untoothed styliform incisor, but lacking molar process and lamina dentata .... Paranthuridae

lB. Mouthparts adapted for cutting and chewing; mandible usually with a molar process and lamina I

dentata, incisor usually toothed ........................................................................................................ 2 |

m

2A. At most, pereopod I subchelate, with propodus expanded; pleonites generally fused, if free, much •

shorter than wide; f'n-stpleopods operculate; with 0, 1, or 2 telsonic statocysts ............... Anthuridae J

2B. Pereopods I-III subchelate, subsimilar; pleonites free, often as long as wide; first pleopods not m

operculate; never with telsonic statocysts ......................................................................... Hyssuridae |

Family Anthuridae Leach, 1814 N

Description. Mandibular palp apically rounded, incisor usually toothed; lamina dentata and molar U

process usually present. Maxilliped no more than 5-articulate, with palp as broad as basis and last article

smaller than preceding articles; palp often broad and somewhat operculiform to other mouthparts. Pereopods II

I-VII with very few sensory setae, usually only 1 distal spine on carpus and propodus; pereopod I usually l

subchelate, with propodus inflated; pereopods II and III never subchelate. Pleon short; pleonites free or fused

(at least dorsally). Pleopod 1 larger than pleopods 2-5, with operculiform exopod and smaller endopod. !

Uropodal endopod usually shorter than basis; with a pair of telsonic statocysts, or single medial statocyst, or !

lacking statocysts. Sexual dimorphism is common and males often have longer pleonites than females.

Remarks. Five species, representing 5 of the 33 described genera of Anthuridae, are known from •

California waters. The MMS survey recovered 3 of these 5 species; Cadien and Brusca, 1993. |

Key References. Menzies, 1951a; Menzies and Barnard, 1959; Schultz, 1977; W_igele, 1979, 1981;

Kensley, 1982; Poore and Lew Ton, 1986; Kensley and Schotte, 1989. •

II

!

!

.
!

I Key to the Genera of Anthuridae Collected as Part of the MMS Surveys

I 1A. Pleopodal endopod 1 with marginal setae ........................................................................................ 2

lB. Pleopodal endopod 1 without marginal setae ...................................................................... Cyathura

i 2A. Pereopods IV-VII with triangular carpus; maxilliped 5-articulate, article 3 wider than long; endite

usually reduced or absent; mandibles symmetrical ................................................... Amakusanthura

I 2B. Pereopods IV-VII with rectangular carpus; maxilliped 4-articulate, article 4 smaller than 3; mandibles asymmetrical, left molar with small tooth which is absent on right molar .................... Haliophasma

I Genus Amakusanthura Nunomura, 1977

I Synonymy. Apanthuretta Wiigele, 1981b.

Description. Dorsum smooth, sometimes pigmented; eyes present, or not visible. Antenmdar flagellum

I short, of 3 articles, the last short andbearing aesthethascs, flagellum short,

3 Anteimal of 2-4 articles. Mandibles

symmetrical, not sexually dimorphic; incisor, lamina dentata and blunt molar process present; palp 3-articulate,

article 3 one-third length of article 2 and with 3-4 terminal setae. Maxilliped bearing an acute filamentous

I endite with a terminal seta; palp of 3 articles, article 1 wider than long; article 2 usually a row

with of mesial

setae; article 3 oblique, subterminal, much smaller than 2, with 4-5 apical setae. Pereopod 1, propodal palm

stepped (sometimes weakly toothed), with stout mesial setae. Pereopods 2 and 3 with propodus only a little

I more on posterior pereopods. Pereopods triangular-trapeziform carpus,

robust than 4-7 with with free anterior

margin. Pleon longer than wide; pleonites fused or often pleonites 1-4 separated dorsally by shallow integumental

grooves, 4-5 fused dorsally. Uropodal endopod as long as peduncle, its margins setose; exopod narrow and

I with a sinuous dorsal margin, or with an obscure dorsal lobe.

Males. Male head smaller than in juvenile, with broadened flattened rostrum, antenna 1 flagellum

i with more than 10 isometric articles each beating numerous aesthetascs, much longer than head. Male pereopod 1 not grossly modified. Pleotelson narrower, with mid-dorsal longitudinal depression.

Remarks. Amakusanthura can be distinguished from the very similarApanthura by a more elongate

I pleon and by a long male antennal flagellum. For recent reviews of Amakusanthura see Poore and Lew Ton

II (1985 and 1988d). The latter paper provides keys to the Australian species in this and related genera, and

reassigns several species from Apanthura to Amakusanthura, including Apanthura californiensis Schultz,

1964.

I Key References. Nunomura, 1977; Poore and Lew Ton, 1985, 1988d.

!

!

|
!

Figure 1.3. Amakusanthura catiforniensis (Schultz, 1964). Holotype LACM 55-

35.1 (female). California, Los Angeles Co., off Santa Monica Pier,

mud, 73 m., 06 February 1955, coll. R/V Velero IV, AHF 2988-55.

!
!

I Amakusanthura cah'forniensis (Schultz, 1964)

Figure 1.3

I Synonymy. Apanthura californiensis Schultz, 1964.

I Description. Length to 11mm. Eyes present, but small. Frontal margin of head forms acute medial projection, but not extending as far forward as anterolateral projections. Antennules and antennae (of males)

each 8-articulate. Maxilliped with 5 (4 free) articles; endite broad and lobelike. Pereon without dorsal coloration,

I pits, of short or keels. stoutPereoni spinelikete VII setaeabout alonghalfinferior length ofmargin; pereonipropodus te VI; pereopod with 1 VII largepropodus spine atanddistal dactyinferior lus withangle. row

Pleonites 1-5 distinct laterally, but fused medially. Uropodal endopod longer than pleotelson; pleotelson and

I both arisesuro11poda 5distance l ramifromwith base row oofendopod. f many setae along minutely serrated margins. Appendix masculinum of male

Remarks. Schultz's (1964) original description had several errors: pleonites 1-5 are fused medially

I (based on examination of the holotype), the maxillipedal endite is broad and lobelike (not acute), and the apex of the maxillule is truncate (not angular).

Distribution. Known only from type-locality (off Santa Monica, California, 80 m), and from the

I present study.

Key References. Schultz, 1964.

I Genus Cyathura Norman and Stebbing, 1886

i Description. Dorsum often pigmented. Eyes present or absent. Antennular flagellum 1-3 articulate in females, up to 5-articulate in males. Antennal flagellum short, 1-3 articulate. Mandibles symmetrical; with

incisor, lamina dentata, molar process, and 3-articulate palp. Maxilliped of 4 articles (3 free); endite absent or

i reduced. Pereopod I subchelate, propodus inflated, with a tooth on the palm. Pereopods IV-VII with triangular • carpi. Pleon short, about aslong as pereonite VII; pleonites 1-5 short, fused; fusion of pleonite 5 to pleonite 6

and to telson often dorsally demarked. Pleopod 1 exopod operculiform, endopod without marginal setae.

Pleopods 2-5 with endopods each bearing 1 seta. Uropodal endopod short, more or less square or triangular.

I Telson with 2 basal with but without dorsal statocysts; long apical setae, long setae.

Remarks. Cyathura contains more species than any other New World anthuridean genus. Many

i Cyathura occur in thearePacific blind hy(Cp.ogean mundainters Menzies, titial specie 1951sand . OfCthe. guaroensis 8 species noBrusca w known and from Iverson, the 1985). New World C. munda , only is2

known only from California, and C. guaroensis is known only from Pacific Costa Rica.

I Key References. Richardson, 1905; Wfigele, 1979; Kensley, 1982; Brusca and Iverson, 1985; Kensley and Schotte, 1989.

I Cyathura munda Menzies, 1951

Figure 1.4

I Description. Body length exceeding 9 times width. Most specimens white with scattered patches of black chromatophores. Frontal margin of cephalon produced as short subacute rostrum, projecting forward

about same distance as anterolateral angles. Eyes small. Antennular peduncle 3-articulate; male with 4-

I articulate flagellum with long brushlike setae; female with 2- to 3-articulate flagellum with cluster of apical setae on terminal article; basalmost flagellar article of female minute. Antennal peduncle 5-articulate; flagellum

of 2-4 minute setigerous articles. Maxillipedal palp 3-articulate, broad, rounded, middle article longest, apical

!
I

I

I

i I

!_ I!

I

I

I

I

I

I

I

I

I

Figure 1.4. Cyathura munda Menzies, 1951. Based on Menzies' holotype drawing i

(male) and paratype material LACM 48-62.3 (mantas). Manca

paratype material from California, Matin Co., Tomales Bluff, intertidal,

23 May 1948, coll. R. J. Menzies, AHF 48-621. I

14 I
I

I article small. Pereonites I-VI similar in length; pereonite VII stightly shorter than pereonite VI; pereonites 17

and HI each with transverse anterior dorsal groove for reception of posterior margin of preceding pereonite.

i Ple creating onitesillusion 1-5 entire of freely ly fused, articulating without pleonites; obvious lapleonite teral incisions; 6 with pronounced musculaturetransverse of pleonitesdorsal visibridges le through demarcating cuticle

lines of fusion with pleonite 5 and telson. Telson lateral margins smooth, posterior margin slightly concave;

i paired statocysts Remarks.present Cyathura anteriorly. munda closely resembles C. guaroensis (known only from Pacific Costa Rica).

The latter can be distinguished by its dorsal pigment pattern, the setal pattern and lack of a tooth on the

I beyond inferior themargin posteoriforthemargin propodus of theofpleotelson. pereopod I, and the possession of large uropodal endopods extending

Distribution. Matin County, California to the Mexican border and Gulf of California, intertidal (on

I Egregia and Laminaria holdfasts) to 132 m. Key References. Menzies, 1951a; Menzies and Barnard, 1959; Miller, 1975.

I Genus Haliophasma Haswell, 1881

i Synonym. Silophasma Schultz, 1977.

Description. Eyes well-developed. Antennule with 3-articulate flagellum, basal flagellar article minute.

I Antenna absent onwith 4- to 7-articulate flagellum. Mandibles asymmetrical, left molar with small tooth which is right molar; palp 3-articulate; palp article 3 with 1 seta, or with transverse or oblique row of 2 to

many setae. Maxilliped 4-articulate, article 4 smaller than 3. Pereon with dorsolateral grooves and sometimes

i additional very broad;pitting pereopods or sculpII tuandre;IIIpereonites with propodus IV-VI elongate with dorsal andpits. subrectangular; Pereopod I stout, pereopods propodus IV-VII expanded with carpi and

subrectangular and not underriding propodus, carpus about half as long as propodus. Pleonites 1-5 fused at

i least medially; pleonite 6 demarcated from telson by transverse dorsal ridge. Pleopod 1exopod more or less indurate, operculiform. Uropodal endopod shorter than telson, exopod folding alongside or over telson. Telson

thick, more or less indurate and usually dorsally sculptured; pair of statocysts present (but not always obvious).

I multiarticulate Male characterized highly setosebyflagellum, more elongate larger body eyes, form, and elongate less pronounced pereopods,dorsal telsonsculpture, and uropods. antennule Appendix with

masculinum simple.

I from NorthRemarks. AmericaHaliophasma north of Mexico. is primarily Femalesa bearing tropical oogenus, stegitesandhave onlynot a single been reported species for hasthis beengenus. reported

Key References. Schultz, 1977; Poore 1975; W_igele, 1984; Poore and Lew Ton, 1988b.

!

Haliophasma geminatum Menzies and Barnard, 1959

I Figures 1.5 and 1.6

i Silophasma geminatum of Schultz, 1977. Description of female. Cephalon smooth, slightly longer than wide, rectangular in outline, widest at

position of eyes. Rounded rostrum as long as anterolateral lobes. Pereonal tergites rectangular, lateral body

I walls visible in dorsal aspect. Lateral body margins ornamented with pits and depressions. Bases of pereopods I-VI fit into lateral grooves in body wall; grooves directed posteriorly on pereonite PHI, directed anteriorly on

pereonites IV-VI. Pereonite I anterior margin medially scalloped, lateral margins extend anteriorly to acute

I points, posterior margin straight; anterior margin of pereonites II and HI deeply excavated to receive preceding

| "
!

|

Figure 1.5. Haliophasma geminatum Menzies and Barnard, 1959. Female. •

California, San Diego Co., Oceanside, 20 February 1957, coll. R/V Velero

/17,AHF 4868-57. 1_

|

_6 !
I Figure 1.6. Haliophasma geminatum Menzies and Barnard, 1959. Male. California,

San Diego Co., Oceanside, 20 February 1957, coll. R/V Velero IV, AHF

I 4868 -57.

i 17
I

pereonites; pereonites IV-VII weakly excavated; pereonites II-V each with 1 posterior transverse groove N

containing deep medial pit; depth of transverse groove and medial pit gradually decreases posteriorly; pereonites

IV-VH with medial oblong (sometimes keyhole-shaped) pit located on anterior third of pereonite (barely •

visible on pereonite VII); pereonite VII half as long as pereonite VI. Pleon with 4 lateral sutures, scarcely |

visible medially, indicating 5 fused segments; short sixth pleonite demarcated dorsally. Pleopod 1 exopod

operculiform and pitted. Telson spatulate, sculptured with a long median carina and 2 shorter more pronounced It

lateral carinae; all 3 carinae fuse to form an elevated plate anteriorly, the anterior margin of which bears a pit U

that aligns with the posterior notch of pleonite 6. Paired statocysts near base of telson. Uropodal endopod

extending nearly to posterior margin of telson and longer than wide, apex blunt, both margins distinctly It

serrated; exopod pyriform, outer margin sinusoidal, denticulate, slightly shorter than uropodal peduncle, i

Description of male. Antennular peduncle 3-articulate; flagellum very setose extending to posterior

margin ofpereonite IT.Antennal peduncle 4-articulate followed by 4 flagellar articles. Median telsonic carina •

less developed than in females. Appendix masculinum tubular, with simple apex, not extending beyond tip of II

pleopod 2.

Distribution. Reported from Monterey Bay to central west Baja California, Mexico; coastal shelves, •

slopes and submarine canyons, 9 to 512 m. MMS survey voucher material from stations R-4 and R-8 was

examined.

Key References. Menzies and Barnard, 1959; Menzies, 1962; Iverson, 1974; Poore, 1975; Schultz, 1

1977; Poore and Lew Ton, 1988b.

Family Hyssuridae Wagele, 1981 I

Description. Body small and slender. Mouthparts compact, not piercing. Mandibular lamina dentata I

usually present; molar process forms an acute or blunt spine, or is reduced; palp of 1 or 3 articles. Maxilliped

narrow; endite present, short or reaching to second palp article; palp of 5 free articles or single article.

Pereopods I-ffl subchelate; pereopod II as large as or larger than pereopod I, basis linear, carpus produced 1

posterodistally; pereopods IV-VII carpus triangular (rectangular in Hyssura), with 0-2 posterior carpal spines

and 0-2 posterior propodal spines. Pleonites 1-5 freely articulating, relatively elongate. Pleopods 1-5 of equal

length; pleopod 1not operculiform. Uropodal peduncle short, about one-third total length of uropod; exopod n

attached basally on peduncle. Telsonic statocysts absent. Ii

Remarks. Hyssuridae contains 6 genera. The MMS survey collected one species which best fits into n

the genus Kupellonura. 1

u

Key References. W_igele, 198lab; Negoescu and W_igele, 1984; Poore and Lew Ton, 1988c; Kensley

and Schotte, 1989. 1

Genus Kupellonura Barnard, 1925

Description. Eyes present or absent; enlarged in males. Antennule with brush-like flagellum in male; I

flagellum 4-articulate, with one aesthetasc on terminal article and occasionally one on 2nd article. Antennal

flagellum 8-articulate; peduncle without stout plumose setae. Mouth appendages of the "normal biting type," n

not piercing or formed into a conelike bundle. Mandibular molar process forms a simple blunt tooth (spine).

Maxilliped with endite reaching third palp article; palp 5-articulate. Pereon without dorsolateral keels or

dorsal pits. Pereopod 1 with straight palm; ungui short. Pereopods 2 and 3 with 5th article acutely projecting •

inferiorly; 6th article ovate; palm axial, with marginal spines. Pereopods 4-7 carpus triangular; carpus and U

propodus each with one posterodistal spine. Pleon elongate, segments distinct. Pleopod 1 not larger than

others, and with both mmi equally developed. Most or all pleopodal rami with several marginal setae. Pleotelson N

l
!

I shorter than pleonites 1-5, thin, dorsally flat, not indurated, with small statocysts. Uropods not indurate;

endopod free, longer than peduncle; exopod large, broad, and with lateral margin widely convex or lobed;

i exopods hinged transversely, folding dorsally and overlapping broadly over pleotelson. Pleotelson spatulate, about as long as uropods; without statocysts.

Remarks. Synonyms include Horolanthura (of Kensley, 1975 in part), Kensleyanthura W_igele,

I and 1981,W_igele and Belizanthura (1984), and Kensley, Poore and1982. LewThe Toncomplex (1988c). synonymy This genusiswasdescribed peviouslyin Kensley kown from (1982),theNegoescu Atlantic,

Mediterranean, New Zealand, and Caribbean. Poore and Lew Ton (1988c) list "uropodal exopod with a

I lateral feature lobe" that canas betheused singleto unambiguously synapomorphy distinguish for this genus, it fromandtheweclosely agree related that thisgenera mayHyssura be the single Barnardunique and

Neohyssura Amar. It is most easily confused with Hyssura, from which it can be most quickly distinguished

i bycarpus its wide, lobed uropodal exopods (more narrow, and unlobed in Hyssura) and triangular pereopod IV-VII (rectangular inHyssura). Poore and Lew Ton (1988c) also note that the genus is most easily recognized

by the uropodal exopods, which are "held obliquely erect in preserved material and usually bear a lateral

i lobe." AboKeyut Refer a dozeence n species s. Barnard, are currentl 1925;yW_igele, recognized. 1981; Poore and Lew Ton, 1988c.

i gupeilonura sp. A

i Figure 1.7

Description. Without eyes. Anterolateral lobes of cephalon small, rounded; rostrum not extending

beyond anterolateral lobes. Basal peduncular articles of antennule broad and subquadrate, flagellum 4-articulate.

I Mandibular molar process long and acute, with ridge of large spines. Maxillipedal endite reaches second palp

article. Pereonites H-Ill with anterior depression to receive preceding pereonites I-ll, respectively. Dorsum of

pereonite V slightly raised. Lateral body walls of pereonites III-VI barely visible in dorsal aspect (lateral body

I walls slightly exaggerated in Fig. 10.I.7). Pereonite VII with small dorsal posterior ridge. Pereopods IV-VII

with triangular carpus. Pleonites 1-5 free. Uropodal exopods subrectangular, with distinct lateral lobe, and

overlapping broadly to almost entirely obscure pleotelson; both uropodal rami almost reaching posterior

I of Pleotelson ornamented with two lateral carinae. margin pleotelson. slightly convex dorsally,

Remarks. This problematic species has features of both Kupellonura and Hyssura. We assign it to

i the former genus because of the presence of lobes on the lateral margins of the uropodal exopods, a unique synapomorphy for this genus. It also possess a triangular carpus in pereopods IV-VII, whereas the carpus of

Hyssura species is rectangular in shape. However, the mouth parts are more characteristic of Hyssura in that

i the mandibular molar process is acute (not blunt, as is characteristic of Kupellonura), and the maxillipedal endite is short, reaching only the second palp article (rather than the third article, as is typical of Kupellonura).

One of the two specimens of this species we examined had a 4-articulate flagellum on the left antenna and an

8-articulate flagellum on the fight.

I Distribution. The above description is based upon an examination of MMS survey voucher material

(USNM specimen no. BRC-14; SBMNH specimen no. BRA-14).

I

!

I
I

I

I

I

I

I

I

I

I

I

I

I

I

I

I

I

I

Figure 1.7. Kupellonura sp. A. Sta, BRC-14, USNM Phase I primary voucher •

collection.
!

I Family Paranthuridae Menzies and Glynn, 1968

I and sucking. DescrMandible iption. With usually or without with untoothed, eyes. Mouthparts styliformtoincisor; gether fwithout orm an elongate a molar process cone adapted or lamina for piercing dentata;

palp of 0-3 articles. Maxillules long and slender, with distal barbs or serrations. Maxillae reduced. Maxilliped

I long and tapering, palp with 0-3 articles, basis long and slender, endite longer than one half palp length, or smaller and reduced. Pereopods with several sensory setae on carpus and propodus; pereopods I, or I-1II

subchelate. Pleonites 1-6 usually demarcated dorsally (except in Calathura and Pseudanthura); telson tonguei opereutiform shaped (exceptexopod inParanthura and smallbellicauda endopod. and Pleotelson related species). with 0 orPleopod 1statocyst. 1 larger than pleopods 2-5, with large

Remarks. Five species of Paranthuridae, representing 3 of the 16 genera of the family, occur in

I California waters. One of these species was recovered by the MMS survey. Key References. Poore, 1980, 1984a; W_igele, 1981; Kensley, 1982; Negoescu and W_igele, 1984.

I Genus Paranthura Bate and Westwood, 1868

Description. Eyes present. Antennular flagellum short, never longer than peduncle, of fewer than I0

I articles articles, inshorter female; thanbmshlike peduncle.inMandible male. Antennal with an flagellum acute incisorusually and 3-articulate forms a short palp;flatdistal setosepalpplate articleof with fuseda

comb of about 12 setae. Maxilla forms a sharp, weakly-serrate spine. Maxilliped elongate; endite lacking;

I palp of 1-2 articles, the terminal article minute if present; palp with a proximal seta, a dorsal seta and 12-13 terminal setae; suture between maxillipedal basis and cephalon distinct. Pereon with feeble ornamentation,

otherwise smooth. Pereonites IV-VI without dorsal pits. Pereopod I subchelate, palm axial or moderately

I oblique, pleonite with 6 usually a mesial dorsally cuttingdemarcated edge; pereopods from telson; IV-VIIrarely carpi2-5rectangular. fused dorsally. Pleonites Pleopod usuallyI exopod separateopereuliform, and distinct;

but only slightly indurate. Uropods with narrow or moderately broad exopods folding over telson; uropodal

I endopod without statocyst. usually reaching apex of telson. Telson thin, narrow, not indurate, and with long terminal setae;

Remarks. This is the largest of the paranthurid genera. The 50+ species of Paranthura are all very

I similar, distinctions being made on subtle morphological differences in the shape and proportions of the articles of the pereopods, uropods, and telson. Species of Paranthura are common in shallow temperate and

tropical waters. Two species occur in California waters, P. elegans Menzies, 1951 and P. linearis Boone,

I 1923; the status of the latter is uncertain. Key References. Richardson, 1905; Miller and Menzies, 1952; Poore, 1984a.

!

!

.

!

!
!

Paranthura elegans Menzies, 1951 I

Figure 1.8

Description. Body length exceeding 9 times width, to 9.5 mm in length (to 15 mm in southernmost II

part of range); dorsum mostly unornamented; eyes large. Anterolateral angles of cephalon only slightly longer I

than rostrum. Antennule 8-articulate; article 4 deeply immersed in article 3. Antennal peduncle 5-articulate 1

with first article divided, basal 3 articles partially fused with one another; flagellum composed of a single i

setigerous, clavate article bearing about 3 small indistinct articles distally; flagellum about one half length of

5th peduncular article. Maxilliped 2-articulate, distal article tapering to very narrow point. Pereonite II slightly I

longer than pereonite I; pereonites II and III with dorsal anterior depression to receive posterior margin of I

preceding pereonites; pereonite III slightly wider than long; pereonites IV-V similar in length, roughly one

fourth longer than pereonite HI; pereonite VI with anterior transverse ridge; pereonite VII one half length of II

pereonite VI. Pleonites apparently free, but articulations very faint medially; pleonite 5 three times length of

pleonite 4; pleonite 6 with sinuate posterior margin and pronounced median cleft. Apex of uropodal endopod

extends beyond apex of telson. Telson elongate, with evenly convex setigerous posterior margin and finely 1

serrate posterolateral borders.

Remarks. This is one of the most commonly encountered anthurideans in California waters.

Distribution. Marin County, California to Bahia San Quintin, Baja California, Mexico and into the 1

Gulf of California, intertidal and to 55 m. Found on soft bottom substrates, boat docks, and low rocky

!

intertidal areas with loose sediments. In

Key References. Menzies, 1951a; Menzies and Barnard, 1959. I

Suborder Epicaridea Latreille, 1831 I

I

Description. Ectoparasites of other crustaceans (ostracods, copepods, cirripeds, and matacostracans). •

Eyes usually present in males, typically reduced or absent in females. Antennules very reduced, usually of 2-

w

3 articles; 3-articulate peduncle generally apparent only inlarval stages. Antennae vestigial in adults. Mouthparts Ill

reduced, forming a suctorial cone with a pair of piercing stylets formed from the mandibles; mandibular palp I1

absent. Maxillules and maxillae reduced or lost. Females usually greatly distorted but with oostegites retained; am

body reduced to little more than an unsegmented egg sac in some forms. Males small and not distorted, usually

living upon body of female. 1

w

Remarks. There are no good references on the Epicaridea as a whole, although Strcmberg (1971)

reviews the embryology (including several California species), and Jay (1989) cites several other papers I

containing general information. Overall, the quantity andquality of published work on the Epicaridea lags far II

behind that of the other isopod suborders. Some authors place the 4 recognized families in 2 superfamilies,

Bopyroidea (Bopyridae, Dajidae and Entoniscidae) and Cryptoniscoidea (Cryptoniscidae). The cryptoniscids

are parasites on osWacods, cirripeds, mysidaceans, amphipods, cumaceans and other isopods. They are probably 1

all protandrous hermaphrodites. The Dajidae are parasites of pelagic mysidaceans, euphasiaceans, and caridean

shrimp. The Entoniscidae are internal parasites of various decapods. Bopyridae parasitize a wide range of

decapod crustaceans. 'l

Key References. Richardson, 1905; Nierstrasz and Brender _iBrandis, 1923; StrCmberg, 1971;

Markham, 1975, 1985, 1988; Miller, 1975; Bourdon, 1980; Upton, 1987a-b; Jay 1989; Kensley and Schotte,

1989. I

!
I

I

I

I

I

,

,,

I

s

!

'. I _ :'_

I

I '

! Figure 1,8. Paranthura elegans Menzies, 1951. Paratype LACM 48-62.2.

California, Matin Co., Tomales Bluff, intertidal, 23 May 1948,

I coll. R. J. Menzies.

m 23
!

Family Bopyridae Rafinesque, 1815 I

Description of female. With complete, or at least partial body segmentation. Body slightly to greatly •

asymmetrical, flattened dorsally. Posteroventral margin of head usually with 1-2 lateral projections on each

side. Eyes, if present, are small irregular dorsal pigment spots. Pereon generally with all 7 pereonites distinct,

although first (and sometimes second) pereonite occasionally partially fused to cephalon. Pereopods prehensile •

but reduced, all 7 pairs usually present on at least one side, up to 6 being absent on other side; 5 (rarely 7) I

pairs of oostegites. Pleon of 2-5 free pleomeres, plus pleotelson; sides of pleomeres often produced as large

lateral plates, or epimeres (often resembling pleopodal rami). Pleopods well-developed or rudimentary, but •

usually present, occasionally absent on posteriormost pleonites. Uropods, when present, uniramous or bimmous, I

often resembling pleopods.

Description of male. Very small, at least twice as long as wide, symmetrical and distinctly segmented. •

Cephalon rounded anteriorly, occasionally more-or-less fused with first pereonite. Antennae often prominent.

Eyes as in females. Pereon of 7 distinct pereonites, each usually with a pair of prehensile pereopods. Pleon of

1-5 pleonites, plus pleotelson (if unsegmented, lacking appendages; if multisegmented, often with uniramous •

pleopods or at least with ventral tubercles on all but the last pleonite). Uropods, if present, uniramous. I

Remarks. Adult bopyrids are parasites either on the abdomen or in the branchial chamber of decapod

crustacean hosts. In branchial parasites, the female attaches ventrally to the host's branchiostegite, inducing

a bulge in the host's carapace. Males are much smaller and are usually found on the ventral side of the pleon

of the female isopod. Females brood many small eggs in an oostegial brood pouch, that hatch as a freeswimming epicaridium stage. The epicaridium attaches to an intermediate host, a calanoid copepod. Once on

the copepod, the isopod molts into a microniscus stage, and then into the cryptoniscus stage. The cryptoniscus

detaches from the copepod, is free-swimming, and eventually attaches to the definitive host. Bopyrids are

sequential hermaphrodites. Upon attachment to the definitive host, the cryptoniscus develops into a female; a •

second cryptoniscus settles on the host and develops into a male. The family Bopyridae contains nearly 500

described species in 10 subfamilies world-wide, all but one of which are parasites on decapod crustaceans.

Thirteen species are known to occur on the Pacific coast of North America north of Mexico.

The cyptoniscus stage of the family Bopyridae possess complex antennules of uncertain homologization.

The first article, and often the second, typically bear toothed "gnathobasic margins" that are important in

species-level taxonomy. One to 3 lobes may arise from the third article, each invested with bundles of long

setae. It is these sensory lobes that Bonnier (1900) and Caiman (1909) presumably interpreted as scales, or

vestigial rami or flagella. The antennules of adult bopyrids are greatly reduced or their articulation is obscure,

and "antennular scales" are absent.

Key References. Richardson, 1905; Markham, 1974, 1975.

Genus Munidion Hansen, 1897 I

Description of female. Body smoothly ovate in outline, with 7 free pereonites and 5 free pleonites

(plus the pleotelson); bilateral body axis distortion generally less than 30°. Cephalon subtriangular, medially I

extending deeply into pereonite I; posterolateral border of cephalon with 2 blunt to sharp processes on each I

side, and sometimes with small dentate processes near central point of cephalic processes. Maxilliped without

palp. All pereonites (or only anterior pereonites) with dorsolateral bosses. Coxal plates distinct. Pereopods '1

similar, slightly larger posteriorly; base produced into blunt carina. Pleonites with distinct epimeres, ranging II

from narrow pointed projections to leaflike petiolate expansions. Five pairs of bimmous pleopods, varying

from narrow lanceolate projections to broad foliate structures. Pleotelson forms knoblike or lanceolate process.

Uropods similar to pleopods.

!

24 I
!

i Description of male. Cephalon wider than long, occasionally fused with pereonite I. Eyes absent or

minute. Body much longer than wide. Antennule and antenna each of 3 articles. Pereonites frequently with

i midventral Remarks. tubercle.Munidio All plenonicontains tes fused7into one subtriangular piece; usually without pleopods or uropods. species, all of which are parasitic on galatheid crabs of the genera

Munida and Pleuroncodes.

I Key References. Hansen, 1897; Richardson, 1905; Markham, 1975.

i Munidion pleuroncodis Markham, 1975 Figures 1.9 and 1.10

i

' l Description of female. Length to 9.8 ram; bilateral body axis distortion approximately 18°; body

shape subpyriform.Withouteyes. Cephalon much broaderthan long, subpentagonal with broadfrontalmargin;

anterior margin slightly convex; posteroventral border of cephalon with 2 lateral digitate processes and a

I series ofdentateprocesses across the medial margin. Antennules andantennae obscurely 3-articulate,distalmost article minute. Antennae separatedby a "frontal plate". Maxilliped subtriangular,obscurely segmented. Coxal

plates large, well-developed on all pereonites, extending far beyond edges of pereon and overlapping each

I other. Pereonites with lohelike dorsolateral tergalprojections, increasing in size posteriorly. Pleonites distinct, although pleon obscured by large ovate epimeres and by long biramous lanceolate pleopods. Coxal plates and

pleonal epimeres resembling the large thin,leaflike oostegites.

i Remarks. The male is typically found attached ventrally to the pleon or posterior oostegites of the female. M. pleuroncodis is very similar to M. princeps Hansen, 1897, a parasite on Munida refulgens Faxon,

1893, which occurs offCocos Island and the coast of Ecuador. M. pleuroncodis can be most easily distinguished

by the shape of the pleopodal endopods, which are lanceolate in the former but oval in M. princeps, and the coxal plates, which are expanded laterally in the former but pressed against the sides of the pereon in M.

princeps. The cephalon of M. pleuroncodis is fused to the first pereonite in males, but free in M. princeps.

I Distribution. Central California to least) central Mexico, the "red

(at infesting only pelagic galatheid

crab" Pleuroncodisplanipes Stimpson. Markham (1975) described this species from collections made by the

Inter-American Tropical Tuna Commission from the R/V David Starr Jordan off Baja California, Mexico in

i 1966, and from specimens washed ashore at Pacific Grove,California in I973. The host is a member of the tropical west American fauna and records from north of central Baja California presumably correspond to

northward extensions of warm coastal waters, such as occur during E1 Nifio events.

I Key References. Markham, 1975.

|

!

,

!

!

1
I

I

I

I

!

I

I

I

I

I

I

I

I

I

I

Figure 1.9. Munidion Mexico, offpleuroncodis Baja CaliforniMarkh a, 26°22' am,N,1975115. °05Holotype "q¢, 19-21USNM November 141597 1966,(femcoll.ale). W. 'i

C. Klawe, R/V David Starr Jordan, Sta. 69.

I

I
I

I

I

I

I

I

I

"': i'_

I

I

t

I

I

I

I

I Figure 1.10. MMexico, unidionoffpleBaja uroncodis California, Markham, 26°22q_I, 1975. 115°05'W, Allotype 19-21USNovem NM 141598 b (male). er 1966, coll.

W. C. Klawe, R/V David Starr Jordan, St.a.69.

!

I
!

Suborder Flabellifera G.O. Sars, 1882 I

Description. Eyes usually well-developed; reduced or absent in cave anddeep-sea forms. Antennules I

usually uniramous, without a scale (except in the cirolanid genus Bathynomus), peduncle 3-articulate (4- U

articulate in Serolidae and Phorotopodidae); antennule reduction occurs in some families. Antennal peduncle

commonly 5-articulate, (questionably 6-articulate in some). Frontal lamina, clypeus, and labrum usually n

welt-developed; in groups with the antennae set very close together, the fi-ontallamina is reduced to a narrow

ridge (e.g. Anuropidae, some Aegidae) or a minute plate (e.g. Serolidae). Mandibles usually robust, adapted

for cutting and grinding, occasionally for piercing; lacinia mobilis, spine row and molar process usually n

present, but often reduced or modified in various families; usually with a 3-articulate palp. Maxillule biramous, l

sometimes adapted for piercing. Maxilla biramous, outer ramus usually consisting of 2 lobes. Pereopods

generally ambulatory, sometimes prehensile; pereopods I-II subchelate only in Serolidae, some Sphaeromatidae Ii

(Ancininae), and some Cirolanidae; posterior pereopods sometimes secondarily natatory (some cirolanids). !i

Pleon of 0 to 5 free segments, plus pleotelson. Usually 5 pairs of pleopods present. Uropods arise laterally,

usually forming a tailfan with pleotelson (except in Anuropidae). _l

Remarks. The Flabellifera is a diverse and probably non-monophyletic taxon currently containing M

about 3,000 described species in 180 genera and 15 families. Nearly 60 species, in 9 families, have been

reported from California waters. The MMS survey recovered 4 of these species, in 4 families. The largest n

flabelliferan family, Sphaeromatidae (with about 50 species described from North America north of Mexico), III

is largely restricted to shallow water and the littoral region.

llll

Key References. Richardson, 1905; Miller, 1975. 1

I

i

Key to the Families of Flabellifera Known from California Waters U

1A. Uropods directed ventrally, identical to and functioning with pleopods; body greatly inflated and j

globular; parasitic on gelatinous zooplankton; antennules greatly modified, 2-articulate, distal artical

greatly expanded and scalloped ........................................................................................ Anuropidae

lB. Uropods unlike pleopods, associated with pleotelson; body not greatly inflated and globular; not i

parasitic on gelatinous zooplankton; antennules not as described above ......................................... 2

mum

2A. Uropods greatly reduced, with very small, often clawlike exopod; body less than 4 mm long; burrowing n

in wood or algal holdfasts ............................................................................................... Limnoriidae

2B. Uropods not greatly reduced; body rarely less than 3 mm long; rarely burrowing in wood or algae (a 1

few species of Sphaeromatidae burrow into coastal wood structures, but they are large animals) .. 3 []

composed of 3 or less dorsally visible free pleonites, plus the pleotelson .............................. 4 m

3A. Pleon

M

3B. Pteon composed of 4-5 free dorsally visible pleonites, plus the pleotelson ...................................... 5

I

!

!
!

4A. Pleon composed of 3 dorsally visible free (complete) pleonites, plus pleotelson; cephaton fused medially

with pereonite I; body strongly depressed and expanded laterally; pereonite VII tergite incomplete or

i abybsent; pushing antennaside ae setantennal very clobases; se together, pleopods frontal1-3lasmall mina reduced and natatory, to asmall basistriangular elongated; platepleopods visible only 4-5

large, broadly ovate, suboperculiform ................................................................................. Serolidae

i 4B. Pleon with pereonite composedI (except of 1-2 dorsally inAncinus visible andfree Bathycopea (complete) ); body segments, convexplusdorsally, pleotelson; not strongly cephalondepressed; not fused

pereonite VII tergite complete; antennae not set close together, frontal lamina large and distinct;

pleopods subequal, of modest size, basis not elongated; pleopods 4-5 ovate but not operculiform ...

.................................................................................................................................. Sphaeromatidae

mm

i • 5A. All pereopods prehensile (dactyls longer than propodi); antennae reduced, without clear distinction i

between peduncle and flagellum; maxillipedal palp 2-articulate ................................... Cymothoidae

5B. Pereopods W-VII ambulatory (dactyls shorter than propodi); antennae not as above, with clear

I distinction between peduncle flagellum; maxillipedal palp ................................

and of 2-5 articles 6

I 6A. Maxi mandible llipeds, reduced maxillorules, absent; and maxilla maxillaereduced with stoutot,arecurred, single slender apicalstylet spines;................................ lacinia or molar process Aegidaeof

6]3. Maxilliped without stout, recurved spines; mandible with or without lacinia and molar process; maxilla

i not a slender stylet ........................................................................................................................... 7

i 7A. Mandible with distinct lacinia and large "articulated" bladelike molar process; mandibular incisor generally broad, 3-dentate; maxillule lateral (outer) lobe often with several (10-14) stout spines,

never styletlike or falcate; maxillae well-developed; pereopods I-lH not prehensile (dactyls not longer

i 7B. than Mandible propodi) with.................................................................................................................... lacinia and molar process reduced, vestigial, or absent; mandibular incisor Cirolarfidae narrow;

maxillule lateral (outer) lobe styletlike or falcate; maxillae reduced; pereopods I-Ill usually prehensile

i (occasionally ambulatory) ................................................................................................................ 8

8A. Maxillule lateral (outer) lobe slender, styletlike, apex with 3-5 stout, hooked spines, smaller spines

i subapically; maxilliped with conspicuous endite Tridentellidae

8B. Maxillule not as above, simple and falcate; maxiUiped without an endite (the female of at least one

i species, ExcoraUana houstoni has a small endite) ......................................................... Comllanidae

!

!

!

!

II
!

Family Aegidae Dana, 1853 i

Description. Body cirolanid-like. Dorsum evenly vaulted or quite depressed; smooth. Eyes, when l!

present, usually large, not uncommonly nearly contiguous (entirely contiguous in a few species). Both antennules !1

and antennae well-developed, division between peduncle and flagellum distinct, flagellum multi-articulate;

antermule with 3 peduncular articles; antenna with 5 peduncular articles. Maxillipedal palp of 2, 3 or 5 Ill

articles; terminal articles with spines and/or stout setae. Mandible elongate, incisor narrow, with reduced, l

vestigial, or no lacinia mobilis, spine row and molar process; palp of 3 articles. Coxal plates of pereonites IIVI large and distinct. Pereopods I-ffl prehensile (i.e. dactyls as long as, or longer than, propodi and strongly •

recurved); pereopods IV-VII ambulatory (i.e. dactyls shorter than propodi). Pleon with 4-5 free pleonites, l

plus pleotelson. Uropods flattened, forming a tailfan with pleotelson.

Remarks. The family Aegidae comprises 6 genera. All species are temporary parasites on marine •

fishes. Adults engorge themselves with blood from their hosts, then dislodge and sit on the bottom to digest

their meal. Nine species, in 2 genera, have been reported from Pacific North America north of Mexico, 6 of

which inhabit California waters. Only one species was recovered by the MMS survey. _l

Key Referenc II es. Richardson, 1905; Miller, 1975; Brusca,1983; Bruce, 1988; Kensley and Schotte,

1989; Brusca and France, 1992.

Genus Rocinela Leach, 1815 I

Description. Body less compact and more depressed than in Aega. Eyes well-developed. Anterior i

margin of cephalon usually extended to form a short rostrum covering all or part of antennular peduncle. II

Frontal lamina small, narrow, or arrowhead-shaped, indistinctly fused broad flat with clypeus; labmm free.

Antennules much shorter than antennae; peduncular articles of antennules not expanded. Mandibles with i

incisor narrow, not denficulate; palp of 3 articles, middle article subequal or barely longer than first article, II

which is greatly elongated. Maxilliped with 2- or 3-articulate palp, if 3-articulate the terminal article is very

small; endite greatly reduced. Pereopods I-III often with propodi expanded into a spine-bearing lobe. i

Remarks. Four species of Rocinela occur in California waters: R. murilloi Brusca andIverson, 1985,

R. laticauda Hanson, 1897, R. angustata Richardson, 1904,R. belliceps (Stimpson, 1864). Only R. angustata

was collected by the MMS project, but it is highly likely that R. murilloi occurs in the area as it is proving to I

be the most common Rocinela in museum collections. There has never been a world-wide monographic il

treatment of this genus. Species are poorly known and misidentifications are common. The description and

key provided below are based on characters different from those traditionally used, and both are based on I_

observations of type material of all known Pacific species.

Key References. Richardson, 1898, 1905; Brusca and Iverson, 1985; Kensley and Schotte, 1989;

Brusca and France, 1992. i

Key to California Species of Rocinela i

i

1A. Medial process of uropodal peduncle extended greater than 50% length of endopod; spines onmerus

of pereopod I-III blunt; pereopod I11merus with 3 spines, on inferior margin; uropodal endopod •

longer than exopod ................................................................................................ Rocinela belliceps !

lB. Medial process of uropodal peduncle extended less than 33% length of endopod; spines on merus of

pereopod I-IU acute; pereopod I11meres with 4-8 spines on inferior margin; uropodal endopod shorter

than exopod ...................................................................................................................................... 2

i

l
!

I 2A. Propodi of pereopods I-HI with 4 stout recurved acute spines; merus of pereopod m with 5-8 acute

spines (3-5 distal and 2-3 proximal spines) ........................................................ Rocinela angustata

I 2B. Pro andpodi 1proximal of pereosppinodse) I...................................................................................................................... -l/I with 4-6 acute spines; merus of pereopod Ill with 4 acute spines (3 distal3

I 3A. of pereopods I-III with 5 thin straight acute spines; apical article of maxillipedal palp with Propodi

thin, nearly straight, acute spines ................................................................................... R. laticauda

i 3B. Propodi with of pereopods I-IlI with 4-6 stout, recurved, acute spines; apical article of maxillipedal palp stout, recurved, acute spines .......................................................................... Rocinela muriUoi

I Rocinela angustata Richardson, 1904

Figure 1.11

I Description. Body about 2.5 times longer than wide. Cephalon subtriangular, 2.0 to 2.7 times wider

than long. Eyes large, separated by about 1 eye-width. Rostrum truncate, extended well beyond bases of

I antennae. Frontal lamina narrow, not expanded. Antennae extended to, orpast, pereonite II. Maxillule styliform tapered to apical tooth. Mandibular palp article 2 more than twice as long as article 3, with 13 spines and 3

setae. Maxilla with setose medial margin; inner lobe fingerlike with 2 stout, recurved spines, 1 apical and 1

I subbasal; outer lobe broadly rounded with 2 small, recurved spines on disto-medial edge. Pereonite I longest;

pereonite IV or V widest; coxae not visible, or occasionally posteriormost coxae visible, in dorsal aspect.

Pereopod I dactyl greatly elongate, as long as carpus and propodus combined; propodus with expanded palm,

I with 4 stout, acute curved spines (rarely with acute, slender, straight spines); carpus with 1 spine; merus with 3-6 acute spines, 2-5 distal spines set among setae (distalmost spine distinctly longer than others) and 1

proximal spine. Pereopod HI merus with 5-8 acute spines, 3-5 distal spines set among setae (2 distalmost

i spines short dactyls, distinctlymuch longershorter than othe thanrs)propodi; and 2-3 ischiurn, proximal merus, spines, and otherwise carpusaswith perefringe opod I.ofPere longopoacute ds IV-VII spineswithon

distal margin and acute spines along inferior margin. Pleonite 1 covered by pereonite VII; pleonites 2-4

I sposte ubequal rior inmalrength gin ofanpleotetson; d width; pleortites inner angle 1 andof 5peduncle narrowerexthan tendedpereoni lesstthan es. Ur33opods % length extende of endopod; d slightlyendopod beyond

elongate-ovate, terminally truncate, with about 11 spines (7 lateral and 4 on distolateral border); exopod much

I wider andRemslightly longer than endopod. Pleotelson broadly rounded, wider than pleonite 5. arks. In 1898 Richardson published a redescription of Rocinela Iaticauda Hansen, based on

material collected by the U.S. Fish Commission's Albatross expeditions in the north Pacific. Hansen's original

_ll description of this species was still in press at the time. Unfortunately, all of the Albatross material upon

!l which Richardsonbased herredescription eventually proved notto heR./at/cauda. Hansen's type of the latter

came from Acapulco, Mexico; Richardson'scollections were from Alaska, Canada andCalifornia.Recognizing

I her (including mistake,in Richar the typedsosenrieslater one(1904) syntypeestablished collected ina Honshu, new species Japan). for the Albatross material, R. angustata

Rocinela angustata is often misidentified as R. cornuta, R. belliceps orR. laticauda in museum and

I environmental research collections. It is also extremely similar to the recently describedR, murilloi. Although the latter species was described from the tropics (Costa Rica) recent work has shown it to be the most common

species of Rocinela occurring south of 32°N on the Pacific coast of North America.

i Bering Sea, along coast to Island, Baja

Distribution. Alaska south of western North America Cedros

California, Mexico; 30 to 2214 m taken from fishes, or from soft bottom habitats.

i Key References. Richardson, 1904, 1905; Birstein, 1973; Brusca and France, 1992.

|
I

I

I

i

i

I

I

i

I

I

I

... I

,

i

!

F_re 1.11. Rocinela ang_tata Richardson, 1904. Syntype USNM 22710 (female). Alaska,

Bering Sea, N. W. of Unimak Island. I

I

I
!

i! Family Cirolanidae Dana, 1853

I plates onDpereonites escription.II-VII, Bodyseparated sleek andfromsymmetri bodycbyal,distinct 2-6.5 times sutures. longer Frontal thatmargin wide; with of cephalon well-devel evenly opedconvex coxal

or produced into short rostrum; distinct frontal lamina present. Antennular peduncle 3-articulate, but occasionally

I some with tridentate articles mayincisor, coalesce well-developed to produce 2lobe-like free peduncula spinerrow, articles. spinose Antennal bladelike peduncle "articulated" 4- or 5-articulate. molar process, Mandible and

3-articulate palp. Maxillule outer lobe with 10-14 apical spines; inner lobe with 3-4 apical circumplumose

spines. recurvedMaxilla spines;setose, with distinct, bilobed. minute Maxillipedal to large, palpendite. typicallyPereopods 5-articulate, ambulatory; articles never 1-11Itend withtowards hooked ora grasping strongly

form, with dactyls well-developed. Pleon usually with 5 distinct pleonites plus pleotelson, although fusion/

mR reduction of free pleonites occurs in several genera; pleonite 5 often overlapped laterally by pleonite 4; pleopods

I_ membranous; posterior pleopods often lacking PMS on endopods, especially pleopod 5. Uropods usually with

both rami well-developed, lamellar, forming a "tail fan" with pleotelson; exopod absent or reduced in some

I Remarks. The family Cirolanidae is large, comprising about 52 described genera. Surprisingly, of

genera.

the 8 species (in 6 genera) of Cirolanidae known from California waters, the MMS soft-bottom survey recovered

I only oneKey species. References. Richardson, 1905; Miller, 1975; Brusca and Iverson, 1985; Bruce 1986; Brusca et

al., 1995.

I Genus Metacirolana Kussakin, 1979

I Description. Cephalon with small to moderate-sized rostral process. Frontal lamina anteriorly dilated, freely projecting, often visible in dorsal aspect; clypeus with ventrally-projecting triangular blade; labrum

subequal to clypeus in width, but longer than clypeus. Antennule short, not extended beyond pereonite I;

I ped incisor, unclewith3-articu smalllatae;ccessory basal article tooth onnotmedial articulating marginat ofright right angle mandible; to others. molar Manprocess dible with and broad spine rowtridentate welldeveloped. Maxilliped slender, endite with 1-2 coupling spines. Pleon with 5 free somites; lateral margins of

I pleonite subbasally5 not (about overlone-third apped bydistance pleoniteform4. Pleopods base) on pleopod 1-2 similar 2. Uropods to each with other;inner appendix agle ofmasculi peduncle na inserted acutely

produced.

I Cirolanidae Remarks. Species of Metacirolana can be most readily distinguished from other, similar genera of by the freely projecting clypeus, dilated frontal lamina (often visible in dorsal view), pleonal and

mouthpart morphology. The genus contains about 2 dozen species, only one of which is currently known from

I Pacific North America north of Mexico. Key References. Bruce, 1981, 1986; Botosaneanu, et al., 1986; Brusca et al., 1995.

I

!

1

.!
!

Metacirolanajoanneae (Schultz, 1966) I

Figure 1.12

I

Description. Small, adults 3-5 mm long. Eyes moderate-sized. Antennule extending almost to pereonite

II. Antenna extending almost to pereonite IV. Coxal plates well-developed, II-VII visible in dorsal aspect, ,ram

expanded laterally and with acute posterior angle. Pleomeres 2-5 with large, well-developed epimeres, expanded ll

laterally and with acute posterior angles. Pleotelson with strong, medial, longitudinal ridge; margins of pleotelson um

and uropodal rami notched. Uropodal exopod about one half width of endopod; both rami subsimilar in /

length, extending barely beyond posterior margin of pleotelson. !

m

Remarks. The antennular flagellum tends to be slightly longer in males, and males are often more

slender than the females. I

Distribution. Submarine canyons and basins off central and southern California. The northernmost I

published record is from 36°4 I'N, 122°W (off Monterey) at 218 m.

Key References. Schultz, 1966. •

!

Family Serolidae Leach, 1814 I

Description. Body strongly depressed, broad, with large expanded coxal plates. Cephalon deeply

immersed in pereon. Some species quite large (to 80 mm). Eyes present or absent. Antennules with 4 peduncular II

articles. Antennae with 5 peduncular articles. Mandible with 3-articulate palp; incisor process well-developed, J

with 2 subterminal movable spines (presumably 1 representing the tacinia mobilis); molar process absent.

Maxilliped with 3-articulate palp; without coupling spines on endite. Pereonite I fused dorsally (at least I[

medially) with cephalon and encompassing cephalon laterally; pereonite VII tergite indistinct dorsally, shortened HI

and fused to pereonite VI. Pereopod I of both sexes, and also pereopod II of rnost adult males, subchelate, with

dactyl folding back upon an inflated propodus. Pereopods HI-VII ambulatory. Pleon of 3 free pleonites, at B

least first pleonite narrow, not reaching the lateral margins of the body; pleotelson large (pleonites 4-6 fused

with telson). Pleopods 1-3 peduncles elongate, rami subelliptical; exopod of pleopod 4 indurate, operculate,

covering endopod and pleopod 5. Uropods small, rami narrow, peduncle and endopod coalesced in some

species, in which case the exopod may be greatly reduced. m

Remarks. The Serolidae is principally cold-water and southern hemisphere in distribution. Deep-sea

species often have reduced eyes, or are blind. Serolids are carnivores, scavengers, or omnivores. They are

epibenthic, highly motile animals, capable of shallow burrowing. Serolis is the largest of the 21 described

genera, and is the only genus represented in the northern hemisphere.

Key References. Richardson, 1905; Nordenstam, 1933; Sheppard, 1933; Menzies and Barnard, 1

1959; Harrison and Poore, 1984; Brandt, 1988; Wiigele, 1994.

Genus Serolis Leach, 1818 I

Description. Maxillule lateral (outer) lobe a flattened blade with large apical spines, medial (inner) in

lobe a smaller blade with few apical spines. Maxilla lateral (outer) lobe biramous, (medial) inner lobe a 1

flattened blade, both with tong apical setae. Pereonites VI and VII often medially shortened; pereonite VII

(tergite) medially fused to pereonite VI (also, pereonite VI occasionally fused to pereonite V). Coxal plates

well-developed; those of pereonites III-V marked off by distinct sutures. Pereopod II subchelate in males, 1

ambulatory in females. Uropods lateral, lamellar, and usually with articulating rami. Ill

I

34 I

!

Remarks. See Sheppard (1933) for an excellent review of morphology and taxonomy (including a i

key to all species known at the time). A single species of Serolis, S. carinata, is known from Pacific North

America north of Mexico. am

Key References. Richardson, 1905; Sheppard, 1933; Harrison and Poore, t984. n

Serolis carinata Lockington, 1877 II

Figure 1.13 •

Description. Body broadly ovate; males slightly broader than females. Cephalon approximately as n

broad as long, anterior margin excavated at base of antennules, forming small rostrum. Eyes well-developed, II

posterolaterally positioned on cephalon, reniform, with black pigment. Cephalon, pereonites, and pleonites all

with pronounced mediodorsal carina, forming median keel produced posteriorly on each segment as a short U

spine. Pleotelson posterior margin with deep medial notch. Uropodal rami subequal in length and width, U

rounded distally, reaching posterior margin of pleotelson.

Distribution. Southern California to Baja California, Mexico and into the Gulf of California; low i

intertidal to 98 m from soft bottom habitats. Southern records are usually in deeper water, suggesting these

isopods may conform to the latitudinal submersion phenomenon. Specimens collected by the City of San

Diego Ocean Monitoring Program at Pt. Loma have extended this species recorded depth range.

!1 Key References. Lockington, 1877; Richardson, 1905; Sheppard, 1933; Menzies and Barnard, 1959.

!

Family Tridentellidae Bruce, 1984

Description. Eyes well-developed. Body often with dorsal spines, tubercles, or carinae (always better I

developed in the male of the species). Antennular peduncle 3-articulate, basal article not enlarged. Antennal

peduncle 5-articulate, articles 4 and 5 elongate. Frontal lamina narrow, pentagonal; clypeus short, broad, ill

inverted V-shaped, lateral angles produced to, or almost to, base of mandibles; labrum small, partly or largely il

encompassed by clypeus. Mandible with short, acute incisor; molar process vestigial, weakly sclerotized

(often lost in dissection); lacinia absent; palp 3-articulate. Maxillule lateral (outer) lobe styliform, slightly lib

curved, tapering toward apex, with 3-5 stout hooked apical spines and smaller subapical spines; medial l

(inner) lobe simple, greatly reduced. Maxilla uniramous, stout, 2-articulate; distal region of conical second

article with small spines and/or scalelike setae. Maxillipedal palp 5-articulate, middle article not elongate; (IB

endite elongate, with or without coupling spines. Pereopods 1-1IIsubprehensile; pereopods IV-VII ambulatory, i

Pleopods 1-4 peduncles with 4-6 coupling spines on medial margin; rami lamellar, with plumose marginal

setae on all but endopod of pleopod 5. Appendix masculina of male pleopod 2 rodlike, simple, arising from m

proximal medial margin of endopod. !

Remarks. Tridentellidae is a monogeneric family closely related to Corallanidae, Aegidae, Cymothoidae

and Cirolanidae. It ismost often confused with Corallanidae, but can be most easily recognized by the presence U

of a large maxiltipedal endite (lacking in corallanids). il

Key References. Bruce, t984; Delaney and Brusca, 1985.

I

!

!
!

!

!

!

Figure 1.13. Serolis carinata Lockington, 1877. California, San Diego Co., Pt. Loma, 47 m, 20 July 1989, coil.

I Pt. Loma Biology Lab, Sta. A-14, SDNHM A.0014.

!

!
!

Genus Tridentella Richardson, 1905 I

Synonym. Smicrostoma Hale, 1925.

ll

Description. See family description, i

Remarks. Tridentella is a cosmopolitan genus reported from shallow (11 m) to bathyal (935 m)

depths. It is a small genus of only 14 described species. Most occur in temperate waters, but at least three I

occur in tropical seas. The bladelike slicing mandibles, and the hooked spines on the maxillules and maxillae, n

suggest that adults may be predators and/or scavenging carnivores and several species have been reported as

"parasites" on various marine fishes. Two species have been reported from California waters, T. quinicornis I

Delaney and Brusca, 1985 and T. gIutacantha Delaney and Brusca, 1985. Richardson's (1905) record of T. Ii

virginiana Richardson, 1900 from Santa Barbara Island (Albatross Station 4417) almost certainly was based

on T. quinicornis.

i Key References. Richardson, 1905; Menzies, 1962; Schultz, 1969; Kussakin, 1979; Bruce, 1984;

Delaney and Brusca, 1985; Delaney, 1990.

!

Tridentella glutacantha Delaney and Brusca, 1985

Figures 1.14 and 1.15

15

Description of male. Dorsum highly sculptured. Cephalon with frontal margin produced into large

upturned process, and smaller ventrally projecting rostrum that meets the broad frontal lamina. Clypeus short I1

and very broad. Labrum small, partly encompassed by clypeus. Antennular flagellum of 16-17 articles, extending Ii

to middle of pereonite I. Antennal flagellum of 25-28 articles, extending to posterior margin of pereonite IV.

Maxilliped with large endite, extending to apical palp article and bearing 5-6 coupling spines. Dorsum of 1

pereonite I with 3 large processes; all pereonites with numerous dorsal tubercles, increasing in size posteriorly, U

becoming spinelike on posterior pereonites and pleon, and extending onto coxae. Pereonites [II-VII and all

pleonites with row of large tubercles along posterior margin, these also increasing in size posteriorly. Coxal •

plates large, increasing in size posteriorly and extending beyond posterior margins of their respective pereonites; U

coxal plates with 2 oblique carinae, increasing in size posteriorly. Large, unfused penes on sternite of pereonite

VII, extending nearly to pleonite 2. Pleotelson with longitudinal rows of large spinelike tubercles. Uropods lull

extending barely beyond posterior margin of pleotelson; endopod width about 2 times exopod width; endopod II

longer than exopod.

Description of female. The female of this species is generally much tess spinose than the male, lacks I

the pronounced large upturned process of the frontal margin and the horns of the cephalon and pereonite I. U

Remarks. This species appears to inhabit both rock and mud bottoms. T. glutacantha is easily

recognized by its 3 large cephalic horns, and 3 large hornlike tubercles on pereonite I. Only the northwest I_

Pacific species T.cornuta is similarly horned; however, T.cornuta lacks the robust spination ofT. glutacantha.

I

The MMS primary voucher material includes 1 male, 7, females, 1 manca, and 1postmanca. The secondary

voucher specimen is an intersex individual (butnot a mid-molt individual), with a female cephalon and pereonite 1

I, minute penes, well-developed appendix masculina, andtypical male pleon. Z glutacantha can be quickly

distinguished from its California congener T. quinicornis by the dorsal cuticular spines (lacking inthe latter)

and by the dorsal cephalic processes of males (2 posterolateral horns, plus a large rostrum in the former; 5

small cephalic processes in the latter).

Distribution. T.glutacantha is known from central California (Farallon Is.) to Los Angeles. Previous

known records for T. glutacantha were based solely on the type series, which was from near the Farallon 1

Islands (bottom dredge on green mud; 128 to 231 m), near Catalina Island (loose rock bottom; 304 to 310 m), m

and near the Los Angeles breakwater light (large boulders; 320 to 360 m).

Key References. Delaney and Brusca, 1985. i
i

I

I

I

I

Figure 1.14. Tridentella glutacantha Delaney and Brusca, 1985. Holotype CASIZ 025948 (male).

California, Los Angeles Co., off Los Angeles breakwater light, loose rock, 320-360 m,

I 1953, coll. R/V VeIero/V, Sta. 2413-53.

I
I

I

I

" |

i I

'I'

!

!

I

!

I

!

!":'"'" '):""':"":: I

l , '"" " : "; ""' " '_' ,I

i

Figure 1.15. Tridentella glutacantha Delaney and Brusca, 1985. Allotype LACM

53-113.1 (female). California, Los Angeles Co., Santa Catalina Island,

loose 1323-41. rock bottom, 304-310 m, 18 May 1941, coll. RN Velero III, Sta. I

I

I

40 I
!

I Suborder Gnathiidea Leach, 1814

I Description. Eyes usually well-developed, some species on processes (ocular or

in short lobes, ocular

peduncles). Cephalon in males broad and flattened, often with tubercles or bosses; cephalon in females small

and narrow. Antennular peduncle 3-articulate, (rarely 2-articulate), usually with well-developed flagellum.

I Antennal 5-articulate; both antennules and antennae uniramous, without a scale. Males with peduncle

rudimentary maxillae and greatly enlarged mandibles (reminiscent of certain ant or termite castes); mandibular

palp absent. Females without mandibles or maxillae. Both sexes with only 6 free pereonites and 5 pairs of

I pereopods. Female large and rotund, with pereonites HI-V partly or largely fused. Pereonite I entirely

pereon

fused with cephalon in males, its pereopods forming a second pair of maxillipeds (the pylopods) that cover the

buccal field. Pereonite I only partly incorporated into the cephalon in females, but first pereopods still form a

I second pair of maxillipeds (pylopods). In both sexes pereonite VII is greatly reduced and lacking pereopods (in males pereonite VII is narrow and the same width as the pleonites, in females pereonite VII is generally not

discernible). Pleon abruptly narrower than pereon, with 5 free pleonites plus pleotelson. Pleopods tend to be

I simple, saclike structures, often without marginal setae, the mmi positioned side-by-side (rather than asflattened overlapping plates as in most other isopods). Pleotelson triangular or T-shaped. Uropods bimmous and attached

laterally to form a '_tailfan" in conjunction with pleotelson.

I Remarks. The reduced number of pereonites and pereopods, unusual male mandibles (which are not

used in feeding), and distinctive pleotelson, quickly distinguish gnathiids. Females incubate their embryos

internally, and when gravid nearly the entire body cavity is filled with developing embryos, the internal organs

I being hardly discernible. Gnathiids occur from the littoral zone to the deep and they are often quite

sea,

numerous in shallow soft-bottom benthic samples. Adults probably do not feed, and they are often found in

association with sponges. Adults are benthic but the juvenile stage (the "praniza") is a temporary parasite on

I marine fishes, although they are also often collected free-living in benthic samples. Praniza are good swimmers, whereas adults apparently have only limited swimming capabilities. The mouth parts of praniza are styliform,

with acute anteriorly-projecting mandibles. Data are not yet available to allow identification of females and

I juveniles to species, and the taxonomy of this suborder is based entirely on males. About 125 species, in one

family, and 10 genera, have been described. Only Gnathia is known from California waters.

Key References. Richardson, 1905; Monod, 1926; Menzies and Barnard, 1959; Menzies, 1962;

i Miller, Holdich and Harrison, Juilfs and 1987; 1987; 1988.

1975; 1980; Wiigele, W_igele, Camp,

I Genus Gnathia Leach, 1814

Description. Gnathiids with male pylopod 2- or 3-articulate, the first article being large and operculate,

with the outer (straight) margin much longer than the second article, and the third article (if present) much

I smaller than the second article.

Remarks. The MMS survey recovered 4 of the 8 species known to occur in California waters. The

following key allows for the identification of all known California species.

I Key References. See above.

!

!

!

|
!

Key to the California Species of Gnathia (adult males) I

I

1A. Pleotelson triangular or subtriangular in outline .............................................................................. 2 I

==

1B. Pleotelson T-shaped ......................................................................................................................... 5

2A. Mandible with large, distinct tooth on outer margin; no epimeres visible on pleonites in dorsal aspect i

................................................................................................... ...................................................... 3

2B. Mandible without an outer tooth, or with a minute weakly-developed outer tooth; pleonites with weak I

or distinct epimeres, either small, truncate, and ventrally directed, or subacute and laterally directed I

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

3A. Frontal margin of cephalon with pronounced medial lobe, larger than other frontal lobes; mandible

g

with large outer tooth, and with small scooplike inner region beating a crenulate margin; dorsum of

cephalon not tuberculate; pereon more-or-less straight-sided (pereonites all about same width); eyes

may be on ocular lobes .................................................................... Gnathia steveni, Menzies, 1962 1

3B. Frontal margin of cephalon with medial lobe no larger than other lobes; mandible with modest outer

tooth, and with broad scooplike inner region with several large marginal cusps; dorsum of cephalon a

weakly tuberculate; pereon tapering posteriorly (pereonites narrowing posteriorly); eyes never on II

lobes or stalks ........................................................................................................... Gnathia tridens

4A. Without eyes; frontal margin of cephalon trilobed; pleonal epimeres small, truncate, and ventrally- I

directed .................................................................................. Gnathia coronadoensis, Schultz, 1966

4B. With eyes; frontal margin of cephalon not lobed, but minutely crenulate; pleonal epimeres subacute, !

laterally directed ............................................................................................. Gnathia crenulatifrons

5A. Eyes set on distinct ocular peduncles; frontal margin of cephalon 4-lobed; pleonal epimeres in double i

pairs (a pair of venla-ally-directed and a dorsally-directed epimeres on each pleomere) .....................

................................................................................................. Gnathia clementensis, Schultz, 1966 iI_

5B. Eyes not on ocular peduncles; frontal margin of cephalon 1 or 3-lobed; pleonal epimeres in single II

pairs (double pairs may be present in Gnathia sanctaecrucis) ....................................................... 6

!

6A. Pleonal epimeres occur as doublets (2 pairs of epimeres, a dorsal and a ventral, on each pleonite);

frontal margin of cephalon produced into a single large lobe; dorsum of cephalon (and entire body)

strongly hirsute; pleotelson with a pair of large subapical setae; pleonal epimeres truncate ............. I

....................................................................................................................... Gnathia sanctaecrucis 1=

6B. Pleonal epimeres occur as a single pair on each pleonite; frontal margin of cephalon trilobed; dorsum 1

of cephalon weakly hirsute; pleotelson with or without a pair of subapical setae; pleonal epimeres II

subacute ........................................................................................................................................... 7

7A. Mandible outer margin without crenulations or setae; pleotelson without a pair of large apical setae I

....................................................................................................... Gnathia trilobata, Schultz, 1966

7B. Mandible outer margin with setose crenulations; pleotelson with a pair of large subapical setae (not 1

set side-by-side in transverse line, but off-set from one another) ................ Gnathia productatridens W

I
1

Gnathia crenulatifrons Monod, 1926

Figure 1.16

I Description of male. Body about3.3 to 3.7 times as long as broad; sides parallel. Eyes present, never

on lobes orpeduncles. Frontal margin of cephalon broad, slightly convex, minutely crenulate, not produced

I into distinct lobes. Inner margins of mandibles with 3 teeth, sometimes obliterated; tooth on outer margin I

weakly developed. Pylopods 3-articulate. Body with distinct separation between free pereonites 1I and III.

Pleonal epimeres subacute, laterally directed.

Distribution. Santa Cruz Point, Monterey Bay to Pt. Loma, San Diego County, 9 to 1300 m; coastal

shelves, slopes, and submarine canyons. This species has been collected by the City of San Diego Ocean

m Monitoring Program at Pt. Loma, thereby extending its known range. Specimens have been collected from

! gray sand, green mud, and green mud with hydrogen sulfide. MMS survey voucher material was examined

1P

from PJ-7 and PS-14.

Key References. Monod, 1926; Menzies and Barnard, 1959; Schultz, 1964, 1966; Iverson, 1974.

Gnathia productatridensFiguMenzies and Barnard, 1959 re 1.17

on lobesDorespeduncles. cription ofDorsum male. Bofodycephalon about 3tuberculate; to 3.5 times frontal as long margin as broad,produced, sides paral trilobed. lel. EyesInner presemargins nt, neverof

mandibles with 4-5 small teeth; outer margins with a series of 3-5 setose crenulations. Pylopod 3-articulate,

dtransverse istal articleline, minute. but off-set Pleon from small.onePleote another). lson with a pair of submedian subapical setae (not set side-by-side in

Distribution. Until this time, this species had been reported from Point Conception to the Southern

I Californi examineda fromBightstation in 20 R-5. to 164 m. The type material is from green silt. MMS survey voucher material was

Key References. Menzies and Barnard, 1959.

!

G_¢1_ sa_C_eerue_ Seh_tz, 1972

I Figure 1.18

Gnathia hirsuta Schultz, 1966 (not G. hirsutus of G.O. Sars, 1870).

Description of male. Body about 3 to 3.75 times as long asbroad. Cephalon wider than long, covered

with many dorsaltubercles; frontalmarginwith acutely rounded medial projectionandsome lateralcrenulations.

l Eyes present, never on ocular peduncles or lobes. Maxilliped with many plumose setae along lateral margin; endite with 2 coupling spines. Pylopod 3-articulate, apical article minute. Mandible acutely pointed with few

inner teeth and with smooth, toothless outer margin. Entire body, especially anterior pereonites, covered with

I long hairlike setae. Each pleonite with 2 pairs of lateral epimeres, a dorsal and a ventral pair on each side; pleonites with stiff, hairlike setae arising from posterior margin. Pleotelson long, with 2 pairs of large submedian

setae, one apical pair and one subapical pair. Uropodal rami both with large plumose setae; endopod slightly

longer than exopod and just reaching posterior margin of pleotelson.

Remarks. Schultz (1966) originally described this species as Gnathia hirsuta (as it also appears in

his 1969 handbook). This name, however, was preoccupied (Gnathia hirsutus G.O. Sars, 1870), and in 1972

i Schultz proposed the new replacement name, Gnathia sanctaecrucis, for this species.

I 43
!

!

/f

!

Figure 1.16. Gnathia crenulatifrons Monod, 1926. Male. California, San Diego, Co., off Pt. Loma I

ouffall, 05 October 1989, coll. Pt. Loma Biology Lab, Sta. A-5, SDNHM A.0114

I
i

:l

i

,|

i

I

I

I

I

I

i

ii

I

I

I

I

I Figure 1.17. Gnathia productatridens Menzies and Bamard, 1959. Holotype AHF 5712 (male). California, Santa Barbara Co., off Santa

Barbara Pt. Light, green silt, 89 m, 03 July 1957, coll. R/V Velero

i /V, Sta. 5173-57.

i 45
I

I

i

I

I

I

I,

l

I

l

i

i

I

I

I

I

Figure l,lS, Gnathia sanctaecrucis Schultz, 1972. Holotype AHF 5927 (male).

California, December 1959, Channel Islands, Santa Cruz Channel, green sand, 20t m, 22 I

coll. R/V Velero IV, Sta 6805-59.

!

4_ |
!

I Distribution. Santa Mafia Basin to Southern California Bight. The type material is from Santa Cruz

Canyon, from a depth of 218 m and a bottom substratum characterized as rocks and green sand.

I Key References. Schultz, 1966, 1972.

I Gnath/a it/dens Menzies and Barnard, 1959

Figures 1.19 and 1.20

I Description of male. Body about 2.5 to 3.1 times longer than broad, tapering in width posteriorly.

Eyes present, neveron lobes or peduncles. Mandible outer marginwith modest-sized tooth, without crenulations;

inner margin with 6-7 small teeth. Pylopod 3-articulate. Dorsum of cephalon weakly tuberculate; frontal

I margin produced, trilobed. Body separated or not separated between flee pereonites 11and 111.Pleotelson with

pair of submedian subapical setae, and pair of submedian apical setae.

i Distribution. Specimens of this species have been collected from Point Conception (11 to 27 m) and San Clemente (14 m). The type material was from abenthic sample containing dead kelp fragments and red

algae.

I Key References. Menzies and Barnard, 1959.

I Suborder Valvifera G.O. Sars, 1882

Description. Antennular peduncle 3-articulate, uniramous (without a scale), flagellum reduced to

I one or a few vestigial articles. Antenna uniramous, peduncle 5-articulate, flagellum multiarticulate or

uniarticulate. Frontal lamina, clypeus, and labrum well-developed; mandible with or without 3-articulate

palp. Maxillipedal palp of 3-5 articles. Coxal plates prominent. Vas deferens (and penes) of male opening on

I 1 articulation of 1 and VII than the thorax, in all other marine pleonite or on pleonite pereonite (rather on as

isopods). Pleonites variously fused, of 4 or fewer free segments (plus the pleotelson). Uropods biramous or

uniramous, attached laterally on pleotelson, but modified as ventral opercular plates covering pleopods.

I Remarks. The suborder is composed of 7 families. Valviferans are characterized by the absence of

mandibular palps (except in Holognathidae); the presence of penes on pleonite 1; the unique possession of

i uropods opercular to the pleopods; flagellum of antennute reduced to 1 or a few vestigial articles; pleon of 4

or fewer free somites (plus the pleotelson); uropods biramous or uniramous. Thirty-two species are known to

occur in California waters, representing 3 families. The MMS survey recovered 4 species from 2 families.

i Key References. Sheppard, 1957; Miller, 1975; Brusca, 1984; Poore, 1985; Poore and Lew Ton, 1990.

I Key to the California Families of Valvifera

1A Body cylindrical, often geniculate, flexed between pereonites IV and V; anterior pereopods setose for

feeding, posterior pereopods ambulatory; pereonite IV manifestly enlarged or elongated; first pleopods of males with accessory gonopod; cephalon usually fused medially to pereonite I ........... Arcturidae

lB. Body not cylindrical or geniculate; pereopods not as above; pereonite 4 not as above; first pleopods of

I accessory gonopod; cephalon not medially to pereonite ....................

males without fused I Idoteidae

!
J

i I

I

Figure 1.19. Gnatlu'a tridens Menzies and Bamard, 1959. Holotype AHF 5711 (male). California,

Santa VeleroBarba IV, Sta. ra 4822-57. Co., off Santa Barbara Point Light, 16 m, 17 January 1957, coil R/V I

I
!

!

!

!

I /

!

t Figure 1.20. Gnathia tridens Menzies and Barnard, 1959. Paratype AHF 5711 (female). California, Santa Barbara Co., off Santa Barbara Point Light, 16 m, 17 January 1957, coll. R/V

Velero IV, Sta. 5164-57.

!
!

Family Arcturidae G.O. Sars, 1897 I

Synonym. Astacillidae G.O. Sars. n

I

Description. Body cylindrical or tubular, often geniculate (bent between pereonites IV and V, except

inPleuroprion, Neoarcturus and Idarcturus). Antennal flagellum either 2- to 3-articulate (Astacilla, Neastacilla, m

Arcturella, Pleuroprion, Arcturopsis, Arcturina, Neoarcturus, Pseudarcturella and Idarcturus) or of many I

articles (most other genera). Mandible without palp. Pereonite I either distinct, or completely or incompletely

fused with cephalon. Pereonite IV generally manifestly enlarged or elongated. Pereopods I-IV directed anteriorly m

and setose for feeding; pereopods V-VII directed posteriorly and ambulatory. Male pIeopod 1 with elongated |

peduncle and accessory appendix masculina. Uropods usually biramous, with minute endopod concealed by

larger exopod. Sexual dimorphism often marked. m

Remarks. Four species, representing three genera of astacillids occur in California waters:

Microarcturus Nordenstam, 1933, NeastaciUa Tattersalt, 1921, and Idarcturus Barnard, 1914. The MMS

survey recovered one species. II

Key References. G.O. Sars, 1897a; Richardson, 1905; Nordenstam, 1933; Sheppard, 1957; Menzies II

and Bamard, 1959; Bmsca, 1984.

II

Genus Idarcturus Barnard, 1914 B

Description. Body not geniculate. Cephalon fused with pereonite I, sutures visible laterally. Only I

distalmost flagellar article of antennule with aesthetascs. Antermal flagellum 2-articulate; flagellum shorter II

than 5th peduncular article; flagellum often 4-articulate in male. Maxillipedal palp usually 5-articulate. Pereonite

IV longer than others. All pleonal segments fused into one piece. Penes fused in male. II

Remarks. The original diagnosis of the genus is superficial at best and has not been revised since it II

was created.

Key References. Bamard, 1914; Nordenstam, 1933; Menzies and Barnard, 1959. i

Idarcturus aUelomorphus Menzies and Barnard, 1959

Figures 1.21 and 1.22

Description. Eyes lateral and bulging. Antennules not reaching third antennal peduncular article, i

Cephalon indistinguishably fused with pereonite I, narrow, lateral margins nearly parallel, with two prominent m

horns located medially just slightly posterior to eyes. Maxilliped with 2 coupling spines; palp 5-articulate.

Pereonltes II-VII, each with 1 pair of spines set medially on dorsum near posterior margin of each pereonite; i

pereonites V-VII also with paired lateral spines. Pereopod V dactyl with secondary unguis, in some specimens II

often badly worn or sometimes completely absent. Pleon with 1 pair of medial spines. Lateral margins of

pleotelson with 2 posteriorly directed medium-sized, angulate spines; posterior margin produced, apex blunt. I

U

Remarks. L allelomorphus is easily distinguished fromL hedgpethiMenzies, 1951, its only congener

in California waters, by its comparatively weakly ornamented dorsum and longer cephalon. L hedgpethi is i

easily distinguished by the large triangulate anterolateral extensions on pereonites I-VI and large acute

posterolateral spines on pereonites IV-VI. ml

Distribution. Monterey to Pt. Loma (San Diego County), including Cortes and Tanner Banks; 12to

92 m. A common mud bottom species. MMS survey voucher material was examined from station R-5. This n

w

species has been collected by the City of San Diego Ocean Monitoring Program at Pt. Loma, thereby extending

its known range. Iii

Key References. Menzies and Barnard, 1959. I

m
I

I

!

\

I

I

I

I

I

I

I i...

'...', "_.

I

I

,I

I Figure 1.21. Idarc California, turus allel SantaomBarbara orphus Co., McnzoffiesGoleta, and Barnard mediu,m-coarse 1959. Hgray olotypesand,AHF17 m, 571309.

April 1957, coll. R/V VeIero IV, Sta. 4938-57.

!
i

I

I

I

I

I

I

I

' ii

I

I

I

I

I

I

I

Figure 1.22. ldarcturus Co., offBirdaltelomorphus Rock, 32°49.25_N, Menzies 117°19.60 and Barnard, awV,sandy1959. silt, 60Cali m,fornia, 19 October San Diego 1989, I

coll. Pt. Loma Biology Lab, Sta. B-5, SDNHM A.0014.

I
I Family Idoteidae Fabricius, I798

I usually Description. Body slightly to strongly depressed. Cephalon notfused medially to pereonite I.Antermules shorter than antenna, and with flagellum reduced to 1-4 minute articles. Antennal flagellum either

multiarticulate, reduced to one or a few vestigial articles, or reduced to a large clavate article (occasionally

I wi spltahyed, minuovate, te terminal sometimes articles). reduMandible ced. Pereopods withousubequal t palp. Maxillipedal in length, ambulatory; palp of 3-5 articles. pereopodsCoxal I-IIIplmore ates usually or less

anteriorly directed; pereopods IV-VII more or less directed posteriorly. Pleonites tend to fuse; pleon with at

,I most genera.3 pleonites Pleopods defined 1 and 2 with laterally, short2apical or fewer plumose articulating marginalorsetae. markedUropods dorsally;uniramous all pleonites or biramous. fused in Penes some

fused basally, or rarely free at base (only in Idotea and Lyidotea).

I tropical Rseas. emarMost ks. Idoccur oteidsinare shallow some water, of the and mostfewcommo species n isare opodsfoundof tempe at depths rate greater waters, than but they 30 m.areIdoteids rare in

usually live somewhat solitary lives. They are omnivores, many feeding primarily on the marine plants to

I whi capable ch they of undergoing cling. Several colorspecies changewhich when are transferred known toto plants occur ofonanother red, greecolor. n andThisbrown phenome marine nonplhas antsbeare en

documented for the California species ldotea resecata Stimpson, 1857 and L montereyensis Maloney, 1933.

I TSome wenty-seven of those species ranging ointo f idoteids, the Paciinfic6Northwest of the 26 know are included n generain, have Kozloff's been reported (1987) keyfromto theCalifornia idoteids.waters. Three

species in 2 genera, were recovered by the MMS survey.

I Lee, Key References. Richardson, 1905; Menzies, 1950a; Sheppard, 1957; Menzies and Barnard, 1959; 1966a, b, 1972; Lee and Gilchrist, 1972, 1975; Brusca and Wallerstein, 1979a; Wallerstein and Brusca,

1982; Brusca, 1983, 1984; Kozloff, 1987.

!

i Key to the Genera of Idoteidae Collected as part of the MMS Surveys

IA. Maxillipedal palp 4- or 5-articulate; pleon with 3discernible pleonites, with lateralsuturespresent at

I base of pleotelson ..................................................................................................................... Idotea 1B. Maxillipedal palp 3-articulate; pleon with all pleonites fused, with 1distinct pair of anteriorly placed

lateral incisions (or lateral incisions barely discernible) ..................................................... Synidotea

I Genus Idotea Fabricius, 1799

I coxal plates Descexcept ription. Antennal flagellum multi-articulate. Maxillipedal palp composed of 4 or 5 articles. All the coxal plate of pereonite I, distinctly separated from pereonites by deep dorsal groove.

Pleon three segmented, with lateral sutures present at base of terminal segment, indicating another partly

I coaslesced segment. Remarks. Menzies (1950a) synonymized Pentidotea Richardson, 1905 with ldotea (reducing the

former to a subgenus) because he felt that the single character, 4-articulate (Idotea) and 5-articulate (Pentidotea)

I maxillipedal palp did not warrant generic status. Recent unpublished studies by G.C.B. Poore suggest this synonymy may be incorrect, and both genera may once again be recognized.

Key References. Richardson, 1905; Menzies, 1950a; Brusca, 1984. !

!
I

Idotea (ldotea) rufescens Fee, 1926 I

Figure 1.23

I

Description. Anterior margin of cephalon very slightly concave, frontal process apically blunt or

notched; frontal lamina 1 semicircular and medially shorter than frontal process; frontal lamina 2 not visible i

in dorsal view. Eyes large, ovoid. Maxilliped with 1 coupling spine; palp 4-articulate. Posterior pleotelson m

margin concave in outline.

m

Remarks. ldotea rufescens can be distinguished from the closely related species L resecata by its I

very slightly concave frontal margin (distinctly concave in/. resecata), ovoid rather than pyriform eyes, J

elongate frontal process (apex blunt rather than acute) and more compressed and ovate maxillipedal palp

articles inL resecata. Also, the carpus of pereopod VII is considerably longer inL resecata than in/. rufescens, lm

with the largest propodal seta located a considerable distance from the inferior proximal angle (inl. rufescens II

the largest propodal seta occurs at the inferior proximal angle).

Distribution. British Columbia to Central California from shallow-water algal habitats, intertidal to II

82 m. Specimens have also been collected at Santa Catalina Island. MMS survey voucher material was Ii

examined from station R-5.

Key References. Fee, 1926; Menzies, 1950a; Iverson, 1974; Miller, 1975. I

Genus Synidotea Harger, 1878 •

Description. Antennal flagellum multiarticulate. Mandible with molar process. Maxillipedal palp 3- II

articulate. Pleon with all segments fused; with 1 distinct pair of anteriorly placed lateral incision lines or

lateral incisions barely discernible. Uropods uniramous, m

II Remarks. Species in this genus (approximately 40 species worldwide) occur from the littoral zone to

depths of nearly 3000 m. Ten species have been reported from California, 2 of which were collected by the

MMS survey. A good key to the eastern Pacific species can be found in Menzies and MiUer, 1972. I

I

Key References. Benedict, 1897; Richardson, 1905; Iverson, 1972; Mertzies and Miller, 1972; Miller,

1975; Brusca, 1984.

!

Key to Species of Synidotea Collected as Part of the MMS Surveys I

1A. Dorsal maxillipedal region of cephalon slightly raised; inneranterior cephalic tubercles shorter than

posterior cephalic tubercles; coxae of pereonite I entire; pleotelson with several (usually 3 or more) •

minute posterolateral serrations; pleon without clearly discernible lateral incisions; pleotelson convex, II

spatulate, evenly rounded, widest medially .......................................................... Synidotea calcarea

1B. Dorsal maxillipedal region of cephalon with 1medial tubercle; inner anterior cephalic tubercles taller i

1

than posterior cephalic tubercles; coxae of pereonite I notched; pleon with 1 distinct pair of lateral w

incisions; pleotelson with only 1or 2 minute posterolateral serrations, straight-sided or weakly convex,

widest anteriorly ....................................................................................................... Synidotea media •

l

!

!

I
i

I

I

I

I

I

I

I

I

I

I

I Figure 1.23. Idotea rufescens Fee, 1926. Female. Mexico, Baja California, Coronado Is, 08 November 1946, coll. C. C. Hubbs,H46-118, LACM 46-57.1

I
!

Synidotea calcarea Schultz, 1966 I

Figure 1.24 I

Description. Eyes on very small ocular lobes; lightly pigmented. Cephalon with anterolateral lobes

forming shallow, broad, weakly concave frontal margin; each anterolateral lobe with 1 small tubercle near its I

base; 1 pair of anteriorly-positioned submedian tubercles just behind frontal margin, narrowly rounded, tall; m

1 pair of posteriorly-positioned submedian tubercles between eyes, broad and conical. Dorsal maxillipedal

region slightly raised. Each pereonite with a tall medial conical tubercle and slightly smaller paired lateral m

tubercles. Lateral margin of pereonite I coxae entire, not notched. Entire body covered with very fine short ll

setae. Pleon without clearly discernible lateral incisions. Pleotelson convex, widest medially, spatulate, evenly

w

rounded, with several (usually 3 or more) minute posterolateral serrations. I

Remarks. S. calcarea can be distinguished from the similar appearing S. magnifica by its lightly 1

pigmented eyes with few ocelli (eyes darkly pigmented with many ocelli inS. magnifica) and the presence of

2 large, conical interocular tubercles on the cephalon (small interocular tubercles in S. magnifica). •

Distribution. Tanner and Santa Rosa Canyons at depths of 54 to 813 m. Ii

Key References. Schultz, 1966; Iverson, 1972; Menzies and Miller, 1972.

I

Synidotea media Iverson, 1972

m

Figure 1.25 I

Description. Eyes not raised on ocular peduncles or lobes; heavily pigmented. Cephalon with Im

anterolateral lobes forming shallow, broad concave frontal margin; each anterolateral lobe with 1 stout tubercle 1

near its base; 1 pair of anteriorly-positioned submedian tubercles just behind frontal margin, narrowly rounded,

tall; 1 pair of posteriorly-positioned submedian tubercles between eyes, large, broad and conical. Dorsal 1

maxillipedal region with 1 medial tubercle. Pereonites I-IV each with 1anterior and 1posterior medial tubercle; II

1pair of more laterally-positioned submedian tubercles near posterior margin of pereonite set between lateral

concentric ridges and the medial tubercles; pereonites V-VII with less pronounced lateral concentric ridges I

and 1 medial posteriorly-positioned tubercle. Lateral margin of pereonite I coxa notched. Pleon with 1 distinct l

pair of anteriorly placed lateral incisions. Pleotelson straight-sided or weakly convex, widest anteriorly;

posterolateral margin with 1-2 minute serrations.

Remarks. S. media is very similar in appearance to S. magnifica and S. calcarea; Iverson (1972) W

provides a table of distinguishing characters.

Distribution. Off Pt. Soberanes, California, 36°25'W to 36°26'N; 183 m. MMS survey voucher I

material was examined from stations R-4, R-6 and R-8.

Key References. Iverson, 1972; Menzies and Miller, 1972.

I

!

!

!
!!!!!

i:I 57
i

I

I

I

I 

I

I

I

I

I

I

I

" I

• ' ,_

' "..'.i. _ I

Figure 1.25. Synidotea media Iverson, 1972. Holotype CAS 563 (female). California, Monterey Co., off Pt. I

Soberanes, 183 m, July 1971, coll. RN Searcher.

!

i 58 I
!

I 1.3 THESUBORDERASELLOTA

by

I George D.F. Wilson 4

I Intr°ducti°n 5

I of the total. TheThe Asellota asellotes are the haveonecolonized of the most all diverse environments, group ofother isopodthanfamilies the land, comprising which seemsapproximately to be the domain 25%

of the Ortiscidea and a few Phreatoicidea. Asellota are, however, most successful and diverse in the deep-sea

I (Wilson, 1989). This work introduces an interesting assemblage of Asellota from the edge of the deep-sea in the Santa Mafia Basin, including depths from the shallow lighted realm down to nearly 1000 meters. As a

i consequence, species illustrated here have both shallow-water and deep-sea origins.

I The Asellota are perhaps the easiest isopods to define owing to their specialized copulatory apparatus. To make it clear what is meant by Asellota, I offer the following list of features as a definition:

1. Pleonites 4-5, and often always pleonite 3, fused to pleotelson, making an enlarged often wide

I terminal segment.

2. Pleonites 1-2, and rarely 3, if distinct from pleotelson, are only small tings or cuticular bars

I 3. visib Eitherle pleopods ventrally.I, II, or 117will form a distinct operculum over more posterior branchial pleopods

(AseUota never swim with their pleopods).

I 4. Male plcopods II with specialized copulatory apparatus consisting of an enlarged protopod, a

geniculate (knee-like) endopod, and typically well-muscled exopod.

5. Antenna with a distinct basal (precoxal) segment (sometimes fused in advanced groups).

i 6. Pereopodal coxae, if visible, lack coxal plates but may have coxal epimera or spines, where the

coxae will still be visible if viewed laterally.

I 7. Uropods with narrow rami and protopod generally elongate and tubular. 8. As far as is known, Asellota have a specialized sperm conduit in the female alternatively named

either the cuticular organ orthe spermathecal duct. The opening to this duct will either be adjacent

I to the oopore on pereonite 5 or onthe anterolateral or anterodorsal surface of pereonite 5 (Wilson, 1991).

I

!

4Division of Invertebrate Zoology, Australian Museum, 6 College Street, Sydney South, New South Wales 2000, Australia

I _A Disclaimer. as such. TheThdeeyscripti contain ons insufficient included in t

informatio his work naretonounambiguously t complete taxonomic identifydeeascri chptspecies ions anwithin d shoulthe d nocot nbetextuseofd

the global deep-sea asellote fauna. This work is intended to be only an identification guide for the fauna of the

i Santa Maria Basin, and will be less accurate outside the confines of this geographic area. Because deep-sea

asellote species tend to be highly provincial, workers should expect to find different species of each genus in

basins other than the Santa Maria Basin. Moreover, not all species from the Santa Maria Basin will be

covered in this work, simply because they have not been collected. Most deep-sea Asellota, are rare, and

'l therefore may have a low probability of appearing in samples.
!

This definition excludes the families Atlantasellidae (living in saltwater caves on islands) and I

Microcerberidae (living interstitially on beaches). Both families lack the distinctive male copulatory apparatus

found in the Asellota (on the male pleopod II: a geniculate endopod and a strong exopod with a copulatory •

hook). Inaddition, the pleonites 1-2 are larger and much more distinct in these two families than they are in |

most Asellota (Stenasellidae can also have large segments). The Atlantasellidae and Microcerberidae should

be classified tentatively in the suborder Microcerberidea (Brusca and Wilson, 1991). ! l

tim

General Morphology

See general isopod morphology in this chapter, (Section 1.1 and Figures 1.1 and 1.2). A glossary of I

more specific to Asellota may be found in Wilson (1989).

Collection and Preservation I

Unlike other more robust isopod groups, Asellota are extremely fragile and will be recovered only as I

fragments if handled harshly during sampling. Some groups, like the Desmosomatidae, are nearly impossible

to identify without their anterior legs or uropods. Consequently, it is essential that care be taken to prevent I

mechanical fragmentation and poor preservation during sample processing. The following suggestions assume II

that one is sampling soft sediments with a grab or box corer. Other samplers or sediment types may require

some adjustments of the procedures, although the principles are the same. I

--If possible, use a 3001Lscreen. Many Asellota are around 1 mm long. This means that they will be less m

than 0.5 mm wide in frontal profile, and will go through a standard 500ia screen. Even in shallow water, small !

body size seems to be the rule with most Asellota. A 300la screen will insure that practically all Asellota,

II

including mancas, will be retained. m

--Subject the specimens to as little washing as possible prior tofixation. Try to wash any benthic sediment I

samples gently: elutriation or gentle water flows up through screens. Do not use a strong water spray on the

screen! If you try to force the sediment through the screen, you will also force the animals through, too. A I

gentle flow either from above or below the screenis sufficient. For quantitative box corer samples, don't wash |

the top water and top 1 cm of the samples until after the material has been in fixative for a while. Putthe upper

parts of the sample directly into buffered 4% formaldehyde solution. Don't screen the upper sections of the II

sample until you transfer the material to alcohol. Most of the animals will be inthe upper few centimeters of l

the sample. For shallower water, they may occur deeper, although these sediments will also be much oozier. A

good ruleof thumb is that if the sediment is liquified, it will have the highest percentage of animals, andshould m

be fixed without washing. The bulk parts of a sample can be washed in an elutriation kettle or gently on l

m

screens, depending on the mechanical characteristics of the sample.

--Fix the specimens as soon as possible after sampling. As soon as samples are brought to the surface, •

many animals will be dead or dying, mostly due to thermal shock. Once they are dead, the animals' own |

enzymes will startto degrade their bodies; the end result is that the specimens will literally fall to pieces.

Therefore, the sooner you get the specimens infixative, the better. The formaldehyde will crosslink the proteins I

and make the specimens much tougher andable to hang together during washing and manipulation. Do not let

|

a sample sit inthe sun while deploying the next lowering. If manpower restraintsprevent you from dealing

with a sample immediately after recovery, then putthe sample in a cold room at less than 4°C. This will slow II

the degradation of the specimens. The formaldehyde fixative should be well buffered to prevent dissolution of l

the calcium carbonate in the isopod exoskeleton. Some workers use sodium borate which achieves a pH of

around 8, while others prefer sodium bicarbonate which makes a solution just above neutral. In all cases, the

solution be mixed with sea water. I
!

I --Transfer the specimens to ethanol in such a way as to avoid saltprecipitates or osmotic distortion. This

is adual problem, because (1) insoluble salt crystals precipitate on your specimens if you simply dump sea

i will waterblcowovered up like speclitime tlenballoons s into 70-8and0%popethanoffoltheir , andlegs, (2) ifheads, the specetc.imens Bothareof lef these t in effects fresh wmake ater tootheloAsellota ng, they

specimens difficult to identify. This procedure seems to work reasonably well: first wash away excess sediment

i wwash ith gentle screening in sea water. Then, briefly wash the sample with fresh water. Next, give the sample a

of 25% ethanol. Finally, wash the sample into bottles using 70-80% ethanol. This achieves a slow

transfer from sea-water to ethanol without a prolonged exposure to pure fresh water. The timings will depend

on the bulk of the sample: bulky samples will require more time, top water samples can be processed in less

I than a minute.

I Laboratory Methods

Although there are as many different ways to sort benthic samples as there are laboratories in this

I business, a few comments are in order. The fragility of the Asellota should also be taken into account when it

comes time to handle them in the laboratory. Hard forceps may damage or break the specimens, especially if

one is rushed to finish ajob. If possible, pick them up with wide mouth pipettes, Erwin loops (the meiofauna

I specialists them), or spring forceps very pressures. Pipettes are best,

know about which deform under weak

though. Do not squash the specimens with large folded labels (as were many of the voucher specimens used in

this study). Trying to identify asellote pizza is not fun [ If you cannot double vial the specimens (who has the

I time for small labels that not in contact with the and the bottom and do not have

that?), use are top sharp

edges. If you still must use large folded labels, at least put the label in the vialfirst, push it all the way to the

bottom, and then put in the specimens. Rolling the labels doesn't work because the label will expand to the

I inner circumference of the vial, and then by Murphy's Law, the specimens will between the label and vial

get

wall. More pizza! The ideal label is one that has a rounded points at either end, is narrower than the diameter

of the vial, and is shorter than its interior length.

I Despite your most careful processing efforts, some specimens will be fragmented. Save these fragments

because they may be identifiable. This is especially true for small quantitative samples, where most species

will be represented by only one or two specimens. For example, Ischnomesidae practically always lose their

I legs and antennae, my experience been that different species ischnomesids an area may have

but has of in

decidedly different cuticular textures. So match up the fragmented bits by texture and size, and voiIa, you may

be able to reassemble your fragmented specimens. If you are reasonably cautious about your assignments,

I make the identification much easier. Also if or else plans to make taxonomic you Can process you anyone

descriptions of the Asellota, these parts will be invaluable.

I Glossary and Terminology

I A detailed glossary specific for janiroidean Asellota (but largely applicable to most isopods) can be found in Wilson (1989). The terminological notes below touch on a few points that the reader might fred

different between this section and the others in this chapter.

i Setae and Spines. Setal morphology, although not a central part of the taxonomy in this Atlas, may

be more important in isopod systematics once detailed comparative studies are done. A difficulty observed in

most isopod works is the confusion of the terms "seta" and "spine." Oshel andSteele (1988) argue that the use

I of indicate confounds the and of both Poore "spine" to setae ontogenetic phylogenetic origins structures.

(1991) has a similar opinion. Often sensillate setae are found in a morphological series on the same animal

that start at hair-like structure and terminate in robust toothed spine-like setae. Often, authors will call the I
!

former "a seta" and the latter "the spine," even though one can easily fmd intermediate forms. Even worse, I

some authors will call the same structure something "a seta" at one magnification and "a spine" at a higher

magnification. To avoid this confusion, "spine" will be usedonly for simple epidermal outgrowths, and "seta" •

will refer to a complex epidermal organ that consists of a proximally articulated shaft atthe cuticular surface 1

and a variety of distal terminations, mostly pointed. Most setae have nerve extensions and are sensory in at

least one modality. Spines, on the other hand, have only mechanical functions. •

Numbering of Body Parts. This work follows previous isopod authors (Wolff, 1962; Hessler, 1970; I!

Kussakin, 1979; 1982b; 1988; Wilson, 1989) in numbering body or limb segments with Arabic Numerals and

limbs themselves with Roman numerals. The primary purpose of this convention is to avoid confusion segments n

and limbs in descriptions. The convention also has the advantage that one can abbreviate a reference to the II

carpus of the first pereopod as PerI5.

I

Key to the Families of the Asellota and the Species

Recovered from the Santa Maria Basin I

For the purposes of completeness, I here provideakey to all families of the Asellota. The classification []

follows Brusca and Wilson (1991) in excluding the Atlantasellidac andthe Microcerberidae from the Asellota I

and putting them instead inthe Microcerberidea (see diagnosis of Asellota). In addition,ituses new data from

Just and Poore (1992) for completeness. Although only 21 species are listed in this work, the asellote fauna off 1

California, including the deep sea, could exceed 100 species with representatives in all marine families. Most I

of the isopod diversity will be in the janiroidean families. With the addition of shallow water animals and fresh

water contaminants, any family below could be represented in a collection. Although my goal was to divide •

the key on the basis of external characters alone, in some cases it will be necessary to dissect off the mandible 1

(see above). For the first few couplets, it may also be necessary to carefully dissect offthe male pleopods I and

II to determine the morphology and organization of the sperm transfer apparatus. If this is to be done, first •

note how many distinct pleonites are present in ventral and dorsal view. Some families, such as the Janiridae, II

are extremely diverse and not well defined, so it was necessary to make several entries into the key for these

families. I would be grateful if anyone using this key could contact me on encountering difficulties in keying II

out families. This key should be regarded as preliminary, and will be updated as time goes on. This key also l

includes the species, either in the main part where only one species of a family has been found, or toward the

end where confamilial species are separated, n

u

1A. Pleopods 1-11may be opercular (completely covering more posterior pleopods) in males, pleopod II

fused and practically always opercular in the female ....................................................................... 5 II

lB. Pleopods IH are opercular in both sexes, pleopods 1-11small and non-opereular ............................ 2

2A. Pleopods I in male and pleopods 1Iin female completely separate .................................................. 3 I

2B. Pleopods I in male and pleopods I1 in female fused at least basally into single unit ........................ 4

I

3A. Endopod of male pleopod II fused into a single unit with a bulbous tip; pleonites 1-2 short, indistinct

.................................................................................................. Asellidae [fresh-water habitats only] II

3B. Endopod of male pleopod 11distinctly biarticulate separate with an often complex coiled tip; pleonites I

1-2 large, easily seen dorsally ............... Stenasellidae [fresh-water habitats only, generally in caves]

I

I
!

I 4A. Male pleopods I with protopods separate from rami ..........................................................................

...................................................................... Stenetriidae [cosmopolitan marine]: Stenetrium sp. A

I 4B. Male pleopods I with protopods fused to rami .......... Pseudojaniridae [southern hemisphere marine]

5A. Male pleopod II exopods with two distinct segments ....................................................................... 6

I 5B. Male pleopod II exopods fused into a single hook-like or curved segment ...................................... 7

I 6A. Male pleopods I protopods completely fused to distal rami; endopod of male pleopod II pointed, sometimes curved, with either a sperm groove or sperm tube ............................................................

.......................................................................... Protojaniridae [fresh water, cavernicolous habitats]

I 6B. Male I distinct from distal of male II either pleopods protopod rami; endopod pleopod complex or blunt,

but lacking distinct sperm tub ............. Gnathostenetroididae [interstitial and cryptic marine habitats]

I 7A. Male pleopods I with median sperm tube closed ventrally; male penes apparently well separated from

pereopod VII coxa .......................................................................... [Janiroidea sensu stricto] ......... 9

I 7B. Male pleopods I with median sperm groove open ventrally; penes clearly associated with medial process from the pereopod VII coxa or on coxa (interstitially-living asellotes with small, largely nonopercular pleopods) .......................................................................................................................... 8

!

8A. Uropedal exopod inserts subdistally, separate from endopod; pleonite 3 not visible ventrally, completely

i fused to pleotelson; pleotelson as long as or longer than posterior pereonites ...................................

..... "Microparasellidae" [partial, including only genera Angeliera, Paracharon, and Microcharon]

8B. Uropodal exopod inserts distally, adjacent to endopod; pleonite 3 present ventrally, separate from

I pleotelson; ................................. pleotelson shorter Vermeciadidae than posterior [southern pereonites hemisphere ..................................................................... on isolated subantarctic island beaches]

I 9A. Pereopod I distinctly subchelate with dactylus and propodus opposing each other; eyes, if present, often on stalks; antennulae with short flagellae, always with less than 14 articles total ................ 10

9B. Pereopod I often leg-like but ifsubchelate, with propodus and carpus opposing each other and dactylus

I short; eyes, if present, may protrude laterally are never on elongate stalks; may

but found antennulae

be long or short .............................................................................................................................. 13

I 10A. Anus always terminal, without terminal projections of pleotelson; if visible, female spermathecal

duct opening on ventral surface of pereonite 5 and closely associated with oopore (to see this special

I preparation may be required); eyes, if present, sometimes large with many ommatidia ................. 11 10B. Anus never terminal, with distinct, often pointed, terminal projections of pleotelson; female spermathecal

duct opening on dorsal surface of pereonite 5; eyes, if present, with few ommatidia, never large and

I distinct ............................................................................................................................................ 12

!

!

|
!

11A. Body compact, pleotelson often held at angle to rest of body; protopod of uropod absent or tiny; U

mandibular palp generally large .................................................................. Munnidae: Munna sp. A

........... [primarily cosmopolitan shallow water with a few deep-sea taxa; a few brackish water taxa] •

11B. Body more normal, sometimes elongate, pleotelson typically held in line with rest of body; protopod

of uropod distinct, often elongate; mandibular palp absent ...............................................................

.................................................................. Santiidae [cryptic habitats, cosmopolitan shallow water] N

I2A Protopod of uropod distinct, often elongate; antennal article 3 always short but may have a large n

anteromedial setose projection ...........................................................................................................

............................................... Pleurocopidae [cryptic habitats, tropical to temperate shallow water]

I

12B. Protopod of uropod absent or tiny; antennal article 3 enlarged, and generally much longer than other •

basal articles ...................................................................................................................................... |

Paramunnidae [cosmopolitan shallow water, with moderate number of deep-sea taxa] ................ 44

13A. Posterior part of body arranged as a distinctive natasome, consisting of partially or completely fused N

pereonites 5-7 and pleotelson in most groups; pereopods V-VII with enlarged, paddle-like carpi and

propodi that have articulated plumose setae on their margins; most dactyli with specialized paired •

terminal claws that enclose the distal sensillae ................................................................................... U

Munnopsididae [blind deep-sea cosmopolitan family with emergent shallow water taxa; contains

several subfamilies including the Eurycopinae and the Ilyarachninae] .......................................... 33 n

13B. Posterior part of body never with a distinct natasome; pereopods V-VII carpi and propodi rarely

expanded and natatory (except in some Desmosomatidae, e.g. Eugerda ) and never with articulated U

plumose setae; pereopodal dactyli various, but never with specialized paired terminal claws that

enclose the distal sensillae .............................................................................................................. 14 I

14A. Umpods with elongate protopod, often longer than pleotelson; ff uropods missing, uropodal socket on n

pleotelson large and projecting ....................................................................................................... 15

14B. Uropodal protopod not elongate; uropodal socket not obvious if uropods missing ........................ 20 !

15A. Anterior pereonites compressed into heavy fossorial unit; body never with dorsal or lateral spines;

anterior pereopods fossorial (strong, with many heavy setae), projecting laterally ........................... m

........................... Macrostylidae Hansen, 1916 [blind, deep-sea cosmopolitan; some emergent taxa]

u

15B. Anterior pereonltes distinctly separate; body may be dorsally or laterally spiny; pereopods ambulatory, •

never fossorial, generally projecting ventrally ............................................................................... 16 |

16A. Body moderately narrow to very narrow and elongate; pereonltes often distinctly set apart laterally; n

mandibular molar distally broad and truncate ............................................................................... 17

16B. Body broad, somewhat flattened; cephalon and pereonites often with laterally projecting tergai processes

or lappets; mandibular molar may be narrow and finger-like ........................................................ 19 I

17A. Pereonites 4 and 5 distinctly elongate ............................................................................................ 23 •

17B. Pereonites 4 and 5 near length of other pereonites ......................................................................... 18

!
!

I 18A. Body stiff and thick, never attenuated and spider-like; pleotelson never reflexed dorsally; uropodal

rami tiny or absent, protopod inserting laterally ................................................................................

i 18B. Body ............................................................. sometimes attenuated and spider-like; Echinothambematidae pteotelson often reflexed [blind, dorsally; rare deep-sea uropodal endemic ramigroup] large,

generally biramous, protopod inserting dorsolaterally on pleotelson .................................................

I ................... Dendrotiidae [rare shallow-water taxa with small eyes to blind deep-sea cosmopolitan]

19A. Maxillipedal palp with 4 articles; pereopod I setochelate (using seta on propodus as fixed finger) ..

I Katianiridae [blind, deep-sea cosmopolitan]

19B. Maxillipedal palp with 5 articles; pereopod I not setochelate ............................................................

• ........................................................ Acanthaspidiidae [some with eyes, shallow-water to deep-sea]

i I

20A. Pereopodal dactyli with 3 distinct claws: two large distal claws and one large subdistal accessory

1 claw ................................................................................................................................................ 21

I

20B. Pereopodal dactyli with 1 or 2 distinct claws, third accessory claw tiny or absent ....................... 22

I 21A. Uropods short with thick protopod and button-like rami; uropods inserting postemmedially, almost

touching; cephalon with distinctive rounded rostrum inserting in concave frontal margin; antennae

I compact, geniculate ............................................................................................................................ Joeropsididae [primarily shallow-water group, eyes well-developed to absent] ............................. 37

21B. Uropods various, but pmtopod generally narrow, not substantially larger than rami; uropods separated

I byantennae well-defined generallyanalstraight, region; elongate rostrum, if................................................................................................. present, elongate and not inserting in concave frontal margin;

Janiridae [not monophyletic - taxonomy in poor state; primarily shallow-water with some deep-sea

I genera; eyes well-developed to absent] .......................................................................................... 38

22A. Narrow elongate, sometimes flattened bodies ................................................................................ 23

I 22B. Broad, thick bodies ........................................................................................................................ 28

I 23A. Cephalon well set into and generally fused with pereonite 1; pereonites 4 and 5 elongate; uropods uniramous ................................ Ischnomesidae [blind, deep-sea cosmopolitan; some emergent taxa]

23B. Cephalon separate from pereonite 1; pereonites 4 and 5 approximately same length as other pereonites;

I uropods biramous 24

I 24A 24B. Mandibl andiblee with tsrunca mall, tetrimo angularlar, ........................................................................................................ setose molar ............................................................................... 257

I 25A. Anus inside pleopodal cavity; uropods terminal and adjacent ...........................................................

............ Micropamseliii_lae [sensu stricto, Genus Microparasellus; blind, shallow-water interstitial]

i 25B. Anus separate from pleopodal cavity; uropods separated by well-def'med anal area ..................... 26

!

|
!

26A. Major anterolateral seta on pereonites 2-4 on the coxa, not the tergite ................... Desmosomatidae U

[blind, deep-sea cosmopolitan; some emergent taxa] .................................................................... 42

26B. Major anterolateral seta on pereonites 2-4 on the tergite, not the coxa .............................................. •

....Nannoniscidae [blind, deep-sea cosmopolitan; some emergent taxa]:Nannonisconus latipfeonus

Body extremely long and thin; pereonal ventral surface often broadly "V" shaped ........................... I

27A.

................................................................................. Thambematidae [blind, deep-sea cosmopolitan]

mum

27B. Body more normal; ventral surface rounded .................................................................................. 28 n

u

28A. Cephalon may have eyes; penes not elongate; pleotelson flattened, pleonite 1 generally distinct ......

..................................................................................................................................................... 21B N

28B. Cephalon without eyes; penes fused; pleonite 1elongate; pleotelson dorsally and laterally ovoid ....

..................................................................................................................................................... 32B

!

29A. Tergal surfaces with no dorsal or lateral spination; anus clearly external to pleopodal chamber; bodies

often capable of enrolling or folding .................................................................................................. •

.................................................. Haploniscidae [blind, deep-sea cosmopolitan; some emergent taxa] l

29B. Tergal surfaces often tuberculate or spiny with lateral lappets or spines; anus generally covered by

pleopods; bodies not capable of enrolling or folding ...................................................................... 30 n

30A. Body without large lateral projections or lappets, spines only; body always deep, never flattened ... n

................................................................................. Haplomunnidae [blind, deep-sea cosmopolitan] l

30B. Body with large lateral projections or lappets; body broad, somewhat flattened to narrow ........... 31

ml

31A. Lateral projections or lappets on cephalon; male penes short ............................................................ U

........................................................................................ Janirellidae [blind, deep-sea cosmopolitan]

n

31B. Cephalon rounded laterally, lacking projections or lappets; male penes elongate .......................... 32 n

roll

32A. Spiny body; pleotelson with lateral projections ........... Mesosignidae [blind, deep-sea cosmopolitan] n

32B. Relatively smooth body; pleotelson without lateral projections .........................................................

............................................................................................... Mictosomatidae [blind, rare deep-sea]

N

33A. Cephalon with rostrum projecting anteriorly between antennulae ................................................. 34

33B. Cephalon without rostrum or anterior projections ......................................................................... 35 I

34A. Cephalic rostrum narrow, roundly pointed, not bilobed; natasome robust and deep; pereonite 7 larger

than pereonites 5 or 6 (except in manta stages 1 and 2); uropodal exopod greater than half length of l[

endopod, easy to see ..................................................................................... Eurycope californiensis

34B. Cephalic rostrum broad, distinctly bilobed; natasome flattened, tapering; pereonite 7 smaller than Ill

pereonites 5 or 6; uropod exopod tiny, difficult to see ............................................. Belonectes sp. A I

!
!

!

35A. Natasome not triangular, with flexible somites, pleotelson inflated; pereonite 7 near size of pereonites

I 5 and 6; uropodal protopod small, tubular ..................................................................................... 36 35B. Natasome triangular and only stiffly articulating somites; pereonite 7 distinctly narrower than pereonites

5 and 6; uropodal protopod large, flattened ........................................................ llyarachna acarina

!

36A. Small species, body length of adults less than 2ram; cephalon as broad as pereonite 1,vaulted posteriorly,

with flattened frons; dorsal mandibular muscle insertions in cephalon few and large .......................

I ............................................................................................................................ Munnopsurus sp. A

36B. Large species, body length of adults reaching 7turn; cephalon huge, distinctly broader than pereonite

I 1, dorsal mandibular muscle insertions in cephalon many and small .................. Munnopsurus sp. B

37A. Cephalon with distinct indentation on lateral margin below eyes; cephalic lateral margin not produced,

I into lateral plate, with only small denticulae; ventral midline of pereon with distinct ridges; body narrow, with few setae and devoid of pigment ...................................................... Joeropsis concava

37B. Cephalon lateral margin not indented, with large anteriorly curved spines, substantially produced

I laterally forming large plate; of pereon ridges; body

ventral midline without distinct broad, devoid of

pigment, but with numerous f'me setae ....................................................................... Joervpsis sp. A

I 38A. Anterior pereonites 2 and 3 with paired lappets of varying sizes; pereopod I propodus with

proximoventral denticles ............................................................................. [Genus JaniraIata] .. 39

I 38B. Anterior pereonites 2 and 3 with or without paired lappets of varying sizes; pereopod I propodus without proximoventral denticles .......................................................................................................

....................................... Other Janiridae not included here [likely genera include Ianiropsis, Iais].

!

39A. Cephalon with distinct rostrum; lappets on pereonites 2 and 3 large; pleotelson with distolateral

i 39B. spines Cephalon............................................................................................................................................. without distinct rostrum; lappets on pereonites 2 and 3 small or indistinct; pleotelson without 40

distolaterat spines ........................................................................................................................... 41

I 40A. Body with diffuse pigmented anastomosing chromatophores, with pigment anterior to eyes on cephalon;

obtuse rostral point of cephalon not extending anteriorly beyond anterior spines on lateral margin..

I ................................................................................................................................. Janiralata sp. A

40B. Body completely devoid of pigment; cephalon of adults with pointed rostnnn extending well beyond

i anterior limits of pointed lateral margins ................................................................. Janiralata sp. B

41A. Broad body completely devoid of pigment; lateral margins of cephalon broad, not near bulging eyes;

I pleotelson distinctly trilobed posteriorly .................................................................. Janiralata sp. C 41B. Narrow body with dense pigmented chromatophores on most dorsal parts of the body; lateral margins

of cephalon narrow, near large bulging eyes; pletotelson posteriorly rounded, without laterallobes ........

I ................................................................................................................................. Janiralata sp. D

!
!

42A. Pereonite 1 distinctly larger than pereonite 2 (synapomorphy of Eugerdellatinae), with large ventral m

median spine. Pereonites 3 and 7 with posteriorly curving ventral median spines. Pereopod I with

robust setochetate carpus ....................................................................................... ProcheIator sp. A •

42B. Pereonite 1 subequal or smaller than pereonite 2, without ventral median spines on any pereonites. l

Pereopod I carpus not robust, not setochelate ................................................................................ 43

! 43A. Pereonite 1 subequal to pereonite 2; cephalon with anterolateral spines; uropodal exopod present...

..................................................................................................................... Momedossa symmetrica m

43B. Pereonite 1 smaller than pereonite 2; cephalon lacking anterolateral spines; uropodal exopod absent m

.............................................................................................................................. Desmosoma sp. A

m

44A. Cephalon with small eyes below the antennulae; coxae rounded, only slightly visible in dorsal view m

......................................................................................................................... Austrosignum tillerae

44B. Cephalon with no trace of eyes; coxae with long spines clearly visible in dorsal view .................. 45 m

45A. Antennula article 1 lacking denficulate distal lateral lobe ...................... Pleurogonium californiense []

45B. Antennula article 1 with denticulate distal lateral lobe ...................................... Pleurogonium sp. A

Description of Asellotan Species m

l

Family Stenetriidae Hansen, 1905 m

Genus Stenetrium Haswell, 1881 m

Description (modified from Serov and Wilson, 1994). Head with large, reniform anterolateral eyes;

frontal margin with both lateral and antenna• spines; lateral spines generally extending past antenna• spines. []

Body lateral margins angular or quadrate with coxal extensions visible in dorsal view; pereonite 1longer than []

remaining pereonites. Antenna• article 1 with acutely pointed lateral spine. Male pereopod I robust with

blades or teeth on propodal palm and without denticulate setae or large terminal seta; dactylus subequal to or []

longer than propodal palm width. Male pleopod Ievenly rounded on lateralmargins. Male pleopod II protopod []

distal tipproduced, length subequal to exopod; endopod and exopod positioned on distomedial margin. Male

pleopod IIappendix masculina with one or moredistal spines. Pleotelson with prominent posterolateral spines

and2 free pleonites, m

!

I

I

!

68 m
!

I Stenetriura sp. A

Figure 1.26

|,, Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA-20, 90-130.5 m,

copulatory male (illustrated), early preparatory female.

I Description. Body elongate, unpigmented; all lateral margins with long simple setae; dorsal surface

lacking long setae. Cephalon distinctly broader than long, without projecting rostrum; frontal area with arched

supraclypeal ridge; clypeus semicircular in dorsal view; weakly convex with subtriangular anterolateral points.

I Eyes on cephalon lightly pigmented, not on margin, situated just

lateral behind insertions of antennae. Pereonite

1with fused pointed coxa in male and preadult female. Pereonite 2 with thin pointed anterior lappet. Pereonites

3-4 with both anterior and posterior small lappets separated by broad gap exposing coxae. Pereonites 4-5

I shortest along medial axis than other pereonites. Pleonites 1-2 distinct dorsally and ventrally. Pleonite 3

visible as distinct plate ventrally. Pleotelson shield-like, widest anteriorly, lateral margins slightly convex and

curving down to small posterolateral spines; distomedial margin rounded, projecting posteriorly between

I uropodal insertions. Antennula of male with around 12 articles, article 3 longer than articles 1-2. Antennal scale with elongate setae, some exceeding length of article 1of antennula. Pereopod I of male strongly subchelate

with subquadrate projection on propodus opposing dactylar teeth; many elongate simple setae on the following

I parts: proximomedial side of dactylus, distoventral margin of propodus, proximoventral margin of propodus, ventral margin of carpus, distoventral and distodorsal margins of merus; merus with elongate dorsal spine

extending well beyond distal margin of carpus; dorsal margin of ischium also projecting distally somewhat,

I with several short teeth and elongate setae. Coxae II-VI visible and distinctly bilobed in dorsal view between pereonal lappets. Male pleopod II endopod article 2 longer than medial margin of protopod, distal tip broader

than proximal part, grooved and setose, with small spine-bearing finger-like projection curving dorsolaterally

I from manydistal tip; exopod with distinct suture demarcating two articles on ventral side only. Uropodal rami with distally curved thin setae; exopod subequal to protopod length; endopod longer than protopod.

mml Biology. As usual, little is known about stenetriids. The male copulatory apparatus is rather different

_ from most Asellota, in that the exopod of pleopod II is not endowed with large fan-like muscles, suggesting

u

that the form of copulation in this group may also be different than assumed for most Asellota (Wilson, 1991).

I Remarks. This species of Stenetrium can be separated easily from all others by the distinct thick projection on pereopod I of the male and the presence of many elongate thin setae on the male pereopod I

segments 4-7. Pereonal lappets are better developed in this species than in most stenetriids, and the distal

f'mger-like projection on the appendix masculina is somewhat unusual. The body of this species, however, is

I generally similar to other stenetriids collected from deep and waters. Although typically not

cold it is illustrated

or noted by other authors, readers should be alert to the fact that this species has visible ventral sutures for

m pleonite 3, whereas other species have lost this plesiomorphic feature (e.g. seeStenetrium dagama pleotelson

I illustration in Wilson, 1987).

I

Type Locality and Type Specimens. None. This species is undescribed.

I Distribution. Santa Mafia Basin, 90-130.5 m.

!

!

!
I

I

.... /

C _-' 7 I

I

Figure 1.26. Stenetrium sp. A. Copulatory male from Phase I Sta BRA-20. A, dorsal view, scal

bar = lmm. B, right pereopod I. C, right pleopod U. I

I

_o |
!

I Family Munnidae Sars, 1897

I Genus Munna Kr0yer, 1839

Description (modified from Poore, 1984b). Munnidae with numerous dorsal setae often with setae

I onMandibular head, pleomolar n and ostrong percular andpleopods. subcylindrical, Antennul distally a distaltruncate, article minute, bearing distal accessory two articles setae; palp withextending single aesthetasc. beyond

incisor process, normally armed with second article bearing 1-2 serrate setae. Maxitlipedal epipod apex acute.

I Pereopod dimorphic;I sexually dactyli with dimorphic, two claws, enlarged anterior in malclawe, sometimes largest. Pleopod carposubehelate. I of male Pereopods with acute II-VII lateralnot lobes. substantially Pleopod

HI with broad second article distinctly longer than endopod.

!

Mun_ sp. A

I Figure 1.27

Material Examined. California, Santa Maria Basin, off Purisima Point, Sta. B RC- 14, 105-117 m, 1

i brooding female, illustrated; off Point Arguello, Sta. BRA-04, 168-237 m, 5 individuals, illustrated male pleoped;

Description (generic features modified from Poore, 1984b). Munnidae with large we[l-developed rounded eyes, projecting laterally as far as width of pereonlte 2. All dorsal surfaces covered with fine simple

setae. Frontal margin of cephalon forming smooth quadrate arc in dorsal view. Pereonites axially compressed,

medial pereon length subequal to pleotelson length. Pereonites projecting radially from body centre, so that

I pereonlte 1extends anteriorly under cephalon and pereonite 7 posteriorly pleotelson.

extends under Pereonite

1 shorter and narrower than pereonite 2 in both sexes. Pleotelson with enlarged posterior subanal projection

no wider than anus, not extending under uropods. Antennula with tiny distal article and two aesthetascs.

I Mandible with functional incisor with curved distal article

normal, palp, extending beyond process having

cleaning setae. Pereopods H-VII longer than body, lengthening posteriorly so that pereopod VII nearly 3 body

lengths. Male pleopods I-II not opercular, not covering exopod of pleopod 111;female pleopod II opercular.

Male pleopod I distally flared and truncate with pointed laterally projecting lateral lobes. Pleopod Ill endopod

broad and elongate, extending well beyond endopod, forming operculum for pleopods IV and V in the male.

Uropodal protopod absent; exopod tiny with only one large simple seta; endopod tapering distally with around

i 5 penicillate setae and several simple setae; endopodal distal tip lacking claws, spines, or large denticles.

Biology. Munnids are small, agile janiroideans that seem to favor hard substrates, especially in the

long legged forms such as described here, but can be found in many different micro habitats. Not much is

I known of their life other than behavioral information in Hessler and history, Strcmberg (1989). some Large

males are known to sequester manca 3 females in a lengthy precopula (ibid).

i

species inRethmearkges.nerThe a Munna best entry andintoUromunna the literat(aurfewe on mexcep unnids tionsis Poore could (no19t84bbe),placed). which places Of themNorth ost desceasribterned

Pacific species of Munna, this species can be distinguished fromM, stephenseni by no large lateralspine-like

i sensillate setae on the pleotelson and no spines on the uropodal endopod; fromM, chromatocephala by amore compact body and no spines on the uropodal endopod; from M. halei by a subanal shelf that does not extend

under the uropods; from M. spinifrons by no large robust sensillate setae on the cephalon frons and coxae;

from M. fernaldi by no large sensillate setae on the pleopods I and II and by much longer legs and a more

i compact body. Of these, Munna sp. A is most similar to M. fernaldi.

Type Locality and Type Specimens. None. This species is not formally described.

I Distribution. Santa Maria Basin, hard substrates, 105-237 m.

I 71
I

i

• I

I

!

c I

!

!

• ' " I

1

!

i

Figure 1.27. Munna sp. A. A-B, brooding female from Phase I Sta_ BRC-14, lateral and dorsal views, scale bar

= 1 ram. C, right uropod, lateral view, brooding female. D, pleopod I, ventral view, male from Sta _D

BRA-4.

|

!

I

I
!

i Family Paramunnidae Vanhtffen, 1914

I Munnogonium George Str6mberg,

Genus and 1968

Description. Head partially recessed into first pereonite; eyes, if present, with few ocelli, placed

I below coxae E-VII antennulae visibleoninsma dorsal ll lateral view lacking bumps.spines. Body ovate, Femaleswithout with dorsal laterallyspermathecal projecting duct tergalonspines pereoniteor plates, 5 well

separated from articular membrane between pereonites. Pleotelson lenticular, distally pointed; pleonite one

i large, Antennae length subequal to pereonite 7. Antennulae short, with six articles and with one terminal aesthetasc. short, never longer than body, strongly geniculate: articles 1-3 projecting medially, article 4

dorsolaterally and remaining articles laterally; article 3 distinctly longer than articles 1,2, or 4, without large

:! spines or projections. Mandibular molar process distally broad and quadrate, palp absent. Pereopod I prehensile

I! with major hinges between dactylus and propodus; carpus narrow, longer than broad, participating in grasping

by robust sensillate setae and spines. Pereopods II-VII ambulatory, paucisetose. Male pleopod I sagittate

i distally, smooth ventrally with tufts of small setae on lateral comers; vas deferens entering pleopodal medial sperm duct internally. Uropods tiny, biramous; protopod absent. Anus ventral, covered by opercular pleopods,

anal cavity confluent with pleopodal cavity.

I Munnogonium cf. tUlerae (Menzies and Barnard, 1959)

Figure 1.28

I Synonymy (assuming M. tiUerae =M. erratum).

Austrosignum Austrosi gnum tiIlerae erratumMenzies Schultz, and 1966:307,309-310. Barnard, 1959:8-9.

Munnogonium tillerae Bowman and Schultz, 1974:270; Wetzer et al., 1991:15-16.

I Munnogonium erratum Bowman and Schultz, 1974:270; Wetzer et al., 1991:15.

Material Examined. MMS Collections: California, Santa Maria Basin, off Point Estero, Sta. 1,

l from 158 mrock, 98 m (1); off Morro Bay, Sta. 6, 109 m (2); Sta. BRA-27, 96-126 m (1); off Point Sal, Sta. PJ-17, (2); off Purisima Point, Sta. R-4, 92 m (20, 17); off Point Arguello, Sta.BRA-6, 54-63m (22).---Other

collections: LACM cat. no. 56-26.2: R/V Velero IV station 4753 (type locality), M. tillerae brooding female

t paratype, paratype male illustrated pereopod I, (Holotype missing, see Type dam). LACM cat. no. 57-132.2: M. tillerae, from off Pt. Conception, dissected parts only. LACM cat. no. 57-228.2: M. tiIlerae, paratype

brooding female from Pt. Santa Barbara, decalcified. LACM cat. no. 57-22.2: M. tillerae, paratype from Pt.

I Arguello, station 6003, nothing M. erratum in vial,, perhaps holotypedissected male, apparently by Bowmadissected n and Schultz. by previous LACMauthors, cat. no.only 58-166.1: a few Rlimbs /V Velero in vial. IV

Description. Body broad, ovate. Cephalon somewhat recessed into first pereonite. Eyes situated on

I short bumps ventral to antennulae, small, with few ocelli. Pleotelson lenticular, distally pointed. Anus ventral, covered by opercular pleopods, anal cavity confluent with pleopodal cavity. Antennulae short, with six articles

and one terminal aesthetasc. Antennae shorter than body length; article 3 distinctly longer that articles 1,2, or

I 4; antenna strongly geniculate: articles 1-3 projecting medially, article 4 dorsotaterally and remaining articles laterally. Mandibular palp absent; molar process distally broad and quadrate. Pereopodal coxae laterally

rounded, only slightly visible in dorsal view. Pereopod I prehensile with major hinges between dactylus and

I propodus; robust subtriangular carpus alsoprojection participating on theinbasis grasping distoventml with robust comersensillate and proximal setae. Pereopod projection IIonofischium; mature male juveniles with

and females lacking such lobes on proximal articles of pereopod 11.Pereopods H-VII ambulatory, paucisetose.

Male tiny, biramous, pleopod I always protopodsagittate absent.distally, smooth ventrally with tufts of small setae on lateral comers. Uropods
%. / _'

c I

I

Figure 1.28. Munnogonium cf. tillerae. A and C, male (from Phase II, Sta. R-4). A, dorsal view, scale bar = 0.5 I

mm. B, left pereopod I, brooding female (LACM cat. no. 56-26.2). C, right pereopod 1-I.

I

I

!

_4 |
!

i Biology. Like most janiroideans, species of Munnogonium appear to be epibenthic detritivores. I

have observed M. waIdronense alive at Friday Harbor Labs in the San Juan Archipelago. When alive, this

I spec on theies dorsal appearssurface. to be a small This species ball of detritus seems towithbelegs, clumsyowingandtoslow, its habit perhaps of sticking relyingmaterial on itsfromcamouflage the bottomfor

protection from predators instead of nimbleness (likeMunna). The various populations ofM. fillerae, however,

i are malesmuch less covered with detritus in the samples, so perhaps their methods of defence are different. The of the Northern California sites have a unique enlargements on the basis and ischium of pereopod II.

These lobes may be used in precopula, of whichM, waldronense is known to practice a passive form (Wilson,

199 1).

I Remarks. Bowman and Schultz (1974), in addition to combining Munnogonium waldronense with

Austrosignum tillerae, assert that the following Austrosignum species should be assigned to Munnogonium

I grande owing to(=anglaciale absent) mandibular is the type forpalp: Austrosignum grande, erratum , and,that tillerae it probably , globifrons does, have maltinii a mandibular . The problempalp,heresinceis that the

female glaciale has the palp. Hodgson (1910) never said the palp ofgrande was absent - he only said it was

I "not observed." Moreover, in the pleopod part of his descriptions, Hodgson (1910) describes male pleopods for grande and female pleopods for glaciale (some crustacean taxonomists at the turn of the century didn't

understand the sexual dimorphism in asellote pteopods). The length of the eye stalks might make a good

i cha Hooker racter for separating Austrosignum and Munnogonium. Concerning the composition of Munnogonium, (1985) adds another species, M. wilsoni, toMunnogonium, and gives akey to the species of the genus

in which he also includes a Kensley species subtilis. Both wilsoni and maltini probably should be transferred

to Metamunna Tattersall, 1905 (which should be removed from Paramunna) because of their overall resemblance to M. typica (The type specimens, unfortunately, are probably lost (R. Lincoln, pers. comm.).

Other species that possibly should be included inMetamunna are subtilis Kensley, 1976, Paramunna simplex

j Menzies, P. kerguelensis Vanh6ffen, Munnogonium polynesiensis Miiller, 1989, possiblyAustr/munna serrata

|

Richardson, possibly P. dubia Hale, and possibly Heterosignum mutsuensis Gamo, 1976. An undescribed

species of Metamunna also was found in the La Jolla area by Eric Vetter, Scripps Institution of Oceanography.

i waldronense The.following I am not species sufficiently can beconvinced definitely that assigned M. waldronense toMunnogonium should: erratu be synonymised m, tiIlerae, globifrons into tillerae, andas

proposed by Bowman and Schultz (1974) because they did not discuss specimens from the San Juan Archipelago.

I MMunnogoniu . erratum mis not distinct from the Northern California form of M. tillerae. Much of the confusion in the species tillerae, waldronense, and erratum arises from the distinct elongation of males over

two to three adult (copulatory) molts from the more compact female form. Furthermore, if the specimens are

i exposed to fresh water during fixations, the segments are telescoped, giving a strong separation of the posterior segments. Schultz's (1966) specimen of M. erratum clearly shows everted articular membranes along the

margins of the pereortites, suggesting his holotype is damaged in this way. Therefore, the elongation and

i separation of the lateral pereonites is not a useful character for def'ming species in this genus. Bowman and Schultz (1974) indicated that this was the primary character separating M. tillerae from M. erratum.

Consequently, we must consider the two species identical for the purposes of this Atlas. Additional speciesspecific characters were not found by previous authors, probably because these species are so tiny. This

I will of material from the localities of all three problem require complete redescriptions near type species.

Unfortunately, the type material has been so badly depleted by previous descriptions that the types will not

provide sufficient information. Male types of both M.tillerae and M. erratum are lost or are thoroughly

dissected, of which are missing. The of the northern California population ofM. tillerae

many parts eyes may

have larger and darker ocelli than those of the San Diego specimens. The two populations may differ in the

size of the antennulae, as well. Because quantitative data are not yet available, I have no choice but to

I provisionally synonymizeM, erratum with M. tillerae. If sufficient evidence is found to separate the northern and southern populations atthe species level, then the northern population will be calledM, erratum, and the

southern population will be called M. tillerae.

!

| "
!

Type Locality and Type Specimens. Holotype lost (note in vial left by Bowman and Schultz; see i

also Wetzer et al., 1991:16). Brooding female paratype, 1.0 mm, LACM cat. no. 56-26.2: R/V Velero IV

station 4753, 8 December 1956:5.2 miles at 294° True from Pt. Loma Light, San Diego Co., California, i

32°41.8'N, 117°20.4'W, 101 m, sediment green mud (data from LACM label). Other paratypes listed by II

Menzies and Barnard (1959) and Bowman and Schultz (1974) should not be considered representative of the

holotype because they are not from the San Diego type locality. Unfortunately the latter authors do not an

identify the source of the paratype illustrations in their paper. i

Distribution. Waters off San Diego County to central California (provisional), 18-135 m.

i

Genus PIeurogonium Sars, 1883 g

Description. Head partially recessed into first pereonite; eyes always absent. Body ovate, sometimes B

with laterally projecting spines; coxae II-VII visible in dorsal view with lateral spines. Females with dorsal

spermathecal duct on pereonite 5 well separated from articular membrane between pereonites. Pleotelson i

lenticular, distally pointed; pleonite one large, distinct, ring-like. Antennulae short, with six articles and with g

one terminal aesthetasc. Antennae short, never longer than body, strongly geniculate: articles 1-3 projecting

medially, article 4 dorsolaterally and remaining articles laterally; article 3 distinctly longer that articles 1,2, or

4, without large spines or projections. Mandibular molar process distally narrow with 1-2 spines and 1-2 II

curved setulate setae; palp absent. Pereopod I prehensile with major hinges between dactylus and propodus;

carpus enlarged, as broad or broader than long, participating in grasping with robust sensillate setae and short i

spines. Pereopods II-VII ambulatory, paucisetose. Male pleopod I sagittate distally, smooth ventrally with II

tufts of small setae on lateral corners; vas deferens entering pleopodal medial sperm duct internally. Uropods

tiny, biramous; protopod absent. Anus ventral, covered by opercular pleopods, anal cavity confluent with Jl_

pleopodal cavity. II

Pleurogonium californiense Menzies, 1951 i

Figure 1.29

Pleurogonium californiense Menzies, 1951a:139, figs. 25-26; Menzies and Barnard, 1959:14, fig. 8; Kussakin, I

1962:99; Schultz, 1966:12; Schultz, 1969:291.

Material Examined. California, Santa Maria Basin, off Port San Luis, Sta. R-l, 91 m (15); off I

Purisima Point, Sta. 42, 100 m (7); off Purisima Point, Sta. R-4, 92 m (17); Sta R-5, 154 m (1).--Western

Santa Barbara Channel, off Point Conception, Sta. 79, 98 m (4). i

Description. Body broad, ovate. Cephalon somewhat recessed into first pereonite. Cephalon without

eyes or eye stalks. Pereon and pleotelson with rounded lateral margins lacking large teeth or serrations.

Pereonite 1of males (not intersexes) distinctly longer and more robust than that of females. Cuticle on body B

and limbs smooth to lightly scaled. Antennulae without anterior projection on distal margin. Mandible lacking I

palp; molar process thin, finger-like, with rounded distal tip having only one distal spine and one seta. Pereonal

coxae visible in dorsal view with single projecting spines; spine on coxa VII reduced or absent. Uropods •

biramous, positioned dorsolaterally on pleotelson, protopod absent.

Biology. Populations of this benthic species often may be protogynous hermaphrodites. Fully

hermaphroditic populations will exhibit small primary males, and larger secondary males that were females in i

a previous moult. The large secondary males (approximately the size of a brooding female) are broad and I

have a highly depressed lateral profile and a concave ventral surface, probably retained from the brooding

!
I

i

I A

I

1 B

I

Figure 1.29. Pleurogonium californiense Menzies, 1951. A, Dorsal view of hermaphroditic brooding female,

Phase II Sta. R-4, scale bar = 0.5 mn'L B, left antennula. C, left mandible, dorsal view.

I condition. The smaller primary males have a more normal, convex ventral surface. Preparatory and brooding

females generally have either developing or fully mature male pleopods. Currently, it is unknown whether the

i testes stationare42acti sample) ve in the seems se transitional to lack hermaphrodites, females. Surprisingl indicating y, somthate pthis opulations feature (ofe.g.their thosesexual in thebiology MMS Phase may beI

facultative.

Pleurogonium n. sp. A could also be a different morphotype of P. californiense. Such a hypothesis, however, must accommodate the presence of these differing morpbotypes at all life stages as well as the

differences in the antennules, cuticle, and mandibular molar. These types of differences have not been encountered

before in one species, so at the moment they are classified as separate species.

I
!

Both species may be relatively recent siblings judging from their overall morphological similarity. If I

this is so, their sympatry is all the more interesting. If they are indeed separate species, then the unusual

protuberance (not seen in any other Pleurogonium) on the antennute ofP. n.sp. A may provide an effective n

signal to prevent interspecific matings and concomitant gamete wastage. II

Remarks. Pleurogonium californiense co-occurs with P. n.sp. A., so the two similar species must be

carefully differentiated. P. californiense lacks large projections on the first antennular article, thereby providing •

an easy identification character for separation from P. n.sp. A. P. californiense can be further verified by W

noting its smoother cuticle, its shorter antennulae, and (if dissections are made) the lack of a second short

spine on mandibular molar process. The original description of P. californiense shows a male with a much i

longer antennular article 2 than seen in the current collection. This suggests that the populations of this !1

species North of San Francisco may be somewhat differentiated from those to the South. A comparison of the

type material with the Santa Maria Basin material seems necessary to verify this feature. •

Type Locality and Type Specimens. "Holotype (U.S.N.M. no. 87412), allotype (U.S.N.M. no. J

87413), and 1 male paratype (U.S.N.M. no. 87414), collected at type locality, 3 miles west of mouth of

Russian River, Sonoma County., Calif., July 13, 1947, by R.J. Waidzunas and Paul B. Quyte; found in fine

mud with the sea-star Luidia foliata Grube ..." (Menzies, 1951 a:143).

Distribution. Off northern and central California, 90-154 m.

i

Pleurogonium sp. A

Figure 1.30 I

Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. 42, 100 m (1); off I

Purisima Point, Sta. R-4, 92 m (13); Sta. R-5, 154 m (1).--Western Santa Barbara Channel, off Point I[

Conception, Sta. 79, 98 m (6);

,j

Description. Body broad, ovate. Cephalon somewhat recessed into first pereonite. Cephalon without l

eyes or eye stalks. Pereon and pleotelson with rounded lateral margins lacking large teeth or serrations. l

Pereonite 1 of males (not intersexes) distinctly longer and more robust than that of females. Cuticle on body

and limbs rough with apparent microscales on all surfaces. Antennulae with large curved anterolateral projection lm

on distal margin. Mandible lacking palp; molar process thin, finger-like, with rounded distal tip having two n

distal spines and one seta. PereonaI coxae visible in dorsal view each with single projecting spines; spine on

m

coxa VII large. Uropods biramous, positioned dorsolaterally on pleotelson, protopod absent. /

Biology. Populations of this benthic species may contain protogynous hermaphrodites. See remarks i

above under Pleurogonium caIiforniense.

Remarks. Pleurogonium n.sp. A. co-occurs with Pleurogonium californiense, so the two similar I

species must be carefully differentiated. P. californiense lacks large projections on the first antennular article

found in this species, thereby providing an easy identification character for separation of the two species. P.

n.sp. A. can be further verified by noting its rougher cuticle, its longer antennulae, and (if dissections are Ill

made) the presence of a second short spine on mandibular molar process. I!

Type Locality and Type Specimens. None. This species is undescribed.

Distribution. Santa Mafia Basin and western Santa Barbara Channel, 90-154 m. l_

I

!

!
| B

_//i

I

I

I

Figure 130. Pleurogonium sp. A. A-B, Dorsal view of hermaphroditic brooding female, Phase II Sta. R-4, scale

bar = 0.5 mm. C, left antennula. D, left mandible, dorsal view.

!

I

I
!

Family Janiridae Sars, 1897 I

I

Genus Janiralata Menzies, 195lb

Description (modified from Wilson and W_igele, 1994).Head with dorsal eyes; lateralmargins straight,

with anterolateral points or spines; vertex obtusely pointed to rostrate. Body broad and flattened. Lateral

margins of pereonites 2-3 divided into 2 lobes by deep cleft or broad niche, coxae visible on dorsal view; II

pereonite I with coxal lappet dorsally visible on anterolateral border. Pleonite 1 short, visible dorsally, much

narrower than pereonite 7 or pleotelson. Pleotelson near width of pereonite 7. Antennulae multiarticulate, II

with approximately 12-20 flagellar articles; antennal article 1 Ionger than wide. Antenna article 3 with U

conspicuous scale (exopod); basal article 1-4 subequal; articles 5-6 long, distinctly longer than basal articles

1-4; flagellum multiarticulate, with more than 20 articles; basal articles of flagellum wider than long but not il

conjoint. Mandibular molar process truncate; palp long, article 3 elongate, curved and setose. Maxillipedal II

endite longer than wide, width subequal to palp; palp articles 4-5 narrow, straight sided, distinctly narrower

than broad articles 1-3, palp article 3 distally broad, setose, not tapering, near width of article 2. Pereopod I IB

carposubchelate, dactylus short, with 2 claws; propodus on proximal inferior border serrated, carpus with 2 II

rows of short spine-like setae on venlxal margin. Pereopod II-VII dactylus with two subequal claws, with

subdistal accessory seta enlarged into third claw. Male pleopod I distal tips laterally expanded, with projecting Ii

subtriangular lateral lobes, and broad setose medial lobes merging smoothly into lateral lobes. Male pleopod I_

II distal tip of protopod blunt, setiferous; basal article of endopod only somewhat wider than maximum width

of styler, but not inflated; stytet elongate, curved, longer than sympod. Female pleopod II broad, ovate, with

distal median concavity. Pleopod HI endopod with 3 plumose setae having distinct gap between medial seta i_

and two lateral setae; exopod narrower and longer than endopod; exopod with 2 segments, with distal plumose

setae. Uropods biramous, rami subequal; exopod inserting apically; protopod subequal to rami. I

Jan/ra/_ sp. A

Figure 1.31 I

Material Examined. California, Santa Maria Basin, off Point Sal, Sta. BRA-17, 160.6-168 m (1 il

preparatory female). m

Description. Body broad and flattened, with large dark-pigmented diffuse chromatophores on all tit

dorsal surfaces, including cephalon anterior to eyes. Dorsal surfaces with many f'me setae. Cephalon with m

large dorsal eyes, well separated from lateral margins; lateral margins convexly curved, with triangular I

anterolateral points; anterior margin with obtuse rostral point not extending anteriorly beyond lateral points.

Lateral margins of pereonites 2-3 divided into 2 lappets with broad notches bearing coxae, anterior lappets I

approximately 2 times length of associated coxal lobes in dorsal view. Pereonite 1 bearing only posterior I

lappet shorter than associated coxa; pereonites 4-7 bearing anterior lappets, with that of pereonite 4 being

relatively narrow and that of pereonites 5-7 being broad and laterally rounded; lappets of pereonites 6-7 with B

distinct posterolateral projections. Tergite 5 distinctly shorter medially than other tergites. Pleotelson with I!

semicircular lateral margins, posterolateral medially curving spines, and posterior margin being obtusely

rounded; margin between posterolateral spines and distal tip slightly convex, not concave; tip of pleotelson •

extending beyond posterior limits of posterolateral spines. Antennula article 3 distinctly longer than wide; II

flagellum with around 12 articles, distal article tiny. Antenna with conspicuous pointed scale. Mandibular

article 3 elongate, longer than palp article 1. Articles 1-3 of maxillipedal palp expanded. Pereopod I m

carposubchelate; dactylus short, with 2 claws; propodus with 9 denticles on proximoventral border, carpus II

with 2 rows of short spine-like sensillate setae on ventral margin and fine fringe of cuticular comb-like spinules.

I

80 |
I

I

I

I

I Figure 1.31. Janiralatasp. A., preparatory female, MMSPhaseISta. BRA-17. A, dorsalbody view, scalebar=

1 mm. B, left antennula. C, left pereopod I, with enlargement of opposing margins of propodus and

i carpus.

i Co of xabothI bluntlobesy coxae pointed,II-IIIlargsubequal. e, longer than Femalepereoperculum nal lappet. broad Coxaeovate, H-V laterally distally with bilobedmedian in dorsal concavity. view, length Male

pleopods I-II unknown. Uropods (unknown in this species) probably biramous with branches of subsimilar

i length, sympod about as long as rami.
!

Biology, Little is known about the biology of Janiralata species in general, although they seem to I

favor hard substrates and are quite agile. Janiralata occidentalis is especially quick: it is difficult to capture

with forceps from a petri dish during live sorting because it easily avoids the forceps tips by scampering under Im

debris and clinging tightly. J. solasteri is found as a commensal in the ambulacral groove of the asteroid, II

Solaster species, where it typically co-habits with other commensals like polynoid scale worms. The dense gut

contents of the specimen examined here suggests that, like most janiroideans, this species is a detritivore, a_

Remarks. This species is very similar to Janiralata occidentalis (Walker, 1898). Specimens of J. II

occidentalis from the San Juan Archipelago (Menzies, 1951a), and some fresh specimens from the same

general locality collected by the author suggest thatthe two species can easily be separated by their pigmentation. II

J. occidentalis has dense, non-anastomosing chromatophores that are absent from the region anterior to the !1

eyes, whileJ, sp. A has diffuse anastomosing chromatophores that extend into the region anterior to the eyes.

J. sp. A also has a thinner, more elongate antennula. Readers should be aware, however, that the two species

are nearly identical in most other features, including the rostrum. The rostrum of J. occidentaIis is shown U

much less prominent in (Menzies, 1951a) than it actually is, possibly because Menzies illustrated the specimen

with the cephalon angling down anteriorly. Additionally J. sp. A has fewer proximoventral denticles on propodus g

of pereopod I. II

Type Locality and Type Specimens. None. This species is undescribed.

Distribution. Santa Mafia Basin, 160-168 m, I

Janiralata sp. B

II Figure 1.32

Material Examined. California, Santa MariaBasin, off Point Arguello, Sta. BRA-4, 168-237 m, 1 I

brooding female, 2 juvenile females, I manca.

lit

Description. Brooding female only. Body broad and flattened, with no pigment on any dorsal surface. Ill

Dorsal surfaces with many fine setae. Cephalon with rostrum extending welt beyond anterior limits of pointed |

lateral margins; anterior margin with lateral medially-curving flattened spines adjacent to distinct concave

indentations; eyes large, bulging but not near lateral margin except in early juveniles and mancas. Lateral m

margins of pereonites 2-3 divided into 2 lappets with broad notches bearing coxae, anterior lappets approximately |

2 times length of associated coxal lobes in dorsal view. Pereonite 1bearing only posterior lappet shorter than

associated coxa; pereonites 4-7 beating anterior lappets, with that of pereonite 4 being relatively narrow and II

that of pereonites 5-7 being broad and laterally rounded; lappets of pereonites 6-7 with distinct posterolateral l_

projections. Tergite 5 distinctly shorter medially than other tergites. Pleotelson with nearly straight lateral

margins, with posterolateral medially curving spines, and with posterior margin being obtusely rounded; m

margin between posterolateral spines and distal tip slightly convex, not concave; tip of pleotelson extending i

beyond posterior limits of posterolateral spines. Antennula article 3 distinctly longer than wide; flagellum

with around 12 articles, distal article tiny. Antenna with conspicuous pointed scale. Mandibular article 3 l

elongate, longer than palp article 1. Articles 1-3 of maxillipedal palp expanded. Pereopod I carposubchelate; l_

dactylus short, with 2 claws; propodus with 17 dentictes on proximoventral border, carpus with 2 rows of m

short spine-like sensillate setae on ventral margin and fine fringe of cuticular comb-like spinules. Coxa I

bluntly pointed, large, longer than pereonal lappet. Coxae II-V laterally bilobed in dorsal view, length of both l_

lobes of coxae II-m subequal. Female operculum broad, ovate, distally with median concavity. Male pleopods lit

I-II unknown. Uropods (known only in early juvenile) biramous with branches of subsimilar length, sympod

about as long as rami. i

li

Biology. See remarks under Janiralata sp. A.

I
"1 Figure 132. Janiralata sp. B, brooding female, MMS Phase I Sta. BRA-4. A, dorsal view, scale bar = 1 mm. B, pereopod II with enlargement of proximoventral denticles on propodus.

I

I

I
!

Remarks. Janiralata sp. B is most similar to J. sp. A, J. occidentalis and J. solasteri, but has a i

different combination of features. Unlike J. sp. A and J. occidentaIis, this species lacks pigment and has a

decidedly longer rostrum in the adult. Juveniles, however, have much shorter rostra than in adults and could l]

be confused with other species on this character alone. J. spp. A and B have nearly identical antennulae, so II

readers can consult the plate for the former species for the antennular form of the latter. J. sp B. differs from

J. solasteri in having different lengths of coxal projections on pereopods II and HI, the relative lengths of i

antennular articles 2-3, the terminal extension of the pleotelson beyond the posterior limits of the posterolateral I!

spines, and the number of proximoventral denticles on propodus I in the adult.

Type Locality and Type Specimens. None. This species is undescribed. _l

Distribution. Santa Mafia Basin, 168-237 !1

Janiralata sp. C I

Figure 1.33 l

I

Material Examined. California, Santa Maria Basin, off, Purisima Point, Sta. BRA-16, 91.5-123 m,

1 preparatory female, illustrated, and another 9 individuals, parts of 1 male illustrated.

Description. Body broad and flattened, with no pigment on any dorsal surface. Dorsal surfaces with I

many fine setae. Cephalon with no rostrum, anterior margin smoothly convex or with slight medial rounded

point, margin posterior to limits of anterolateral angles; anterolateral margin broad flattened angle, not spine- I

like; lateral margin only slightly curved; eyes large, bulging, but not near lateral margins. Lateral margins of U

pereonites 2-3 with 2 short lappets, posterior lappets indistinct, separated by broad indentations only slightly

exposing coxae in dorsal view. Pereonite 1 with no posterior tappet; pereonites 5-7 laterally broad and rounded, I

with no obvious anterior lappets. Pereonite 4 with short, broad anterior lappet. Tergite 5 distinctly shorter II

medially than other tergites. Pleotelson broad with smoothly rounded margins, with no posterolateral spines,

and with indistinctly trilobed posterior margin; margin between posterolateral lobes and rounded distal tip Ill

concave; tip of pleotelson extending only slightly beyond posterior limits of posterolateral lobes. Antennula II

article 3 distinctly longer than wide; flagellum with around 6 articles, distal article small, but easily seen.

Antenna with conspicuous pointed scale. Mandibular article 3 elongate, longer than palp article 1. Articles 1- •

3 of maxillipedai palp expanded. Pereopod I carposubchelate; dactylus short, with 2 claws; propodus with 6 1

denticles on proximoventral border, carpus with 2 rows of short spine-like sensillate setae on ventral margin

(3 on lateral side and 10 on medial side) and fine fringe of cuticular comb-like spinules. Coxa I bluntly 1

pointed, large. Coxae II-V not obviously bilobed, barely visible in dorsal view. Female operculum broad, II

ovate, distally with median concavity. Male pleopod I with elongate laterally-pointed copulatory horns and

broad truncate medial lobes. Male pleopod II with short stylet not exceeding length of protopod. Uropods I

unknown, probably biramous with branches of similar length and sympod about as long as rami. !1

Biology. See remarks under Janiralata sp. A.

Remarks. JaniraIata sp. C is most similar to J. koreaensis Jang, 1991 although this new species l

lacks pigment, has a broader pleotelson, has fewer articles in the antennula, and has much smaller lappets on

pereonites 1-4. In this last regard, J. sp C is similar to J. rajata Menzies, 1951, although the width of the body

immediately separates the two species. In Menzies' (1951a) key, this species will key out to J. erostrata, i

although again pereonal lappets and coxal projections immediately distinguish the two species. II

Type Locality and Type Specimens. None. This species is undescribed. a

Distribution. Santa Maria Basin, 91.5-123 m. i

I

84 I
!

i j...

I _.._._,___..

I

I B

I

I Figure 1.33. Janiralata sp. C. A, dorsal view, preparatory female (MMS Phase I Sta. BRA-16), pleotelson

slightly damaged, scale bar = I mm. B-D, copulatory male (MMS Phase I Sta. BRA-16). B, right

i pereopod I, with enlargement of proximoventral spines on propodus. D, pleopod I.

!

!

!

| 8_
I

Janiralata sp. D (cf. rajata) I

Figure 1.34

! Material Exmined.Cafifomia, Santa Maria Basin, off Purisima Point, Sta. BRA- 16, 91.5-123 m, 1

preparatory female body length 3.7 ram.

Description. Body narrow, with numerous dark-pigmented chromatophores on dorsalsurfaces. Dorsal !

surfaces with few fine setae. Cephalon with no rostrum, anterior margin with slight medial rounded point;

anterolateral margins rounded, not spine-like, not projecting anteriorly, with single sensiIlate seta; lateral Ill

margin narrow, only slightly curved, with single sensillate seta adjacent to eyes; eyes large, bulging, extending l[

to lateral margins. Lateral margins of pereonites 2-3 with very short lappets, posterior lappets indistinct,

Ill

separated by broad indentations only slightly exposing coxae in dorsal view. Pereonite 1 with no posterior m

lappet; pereonites 5-7 laterally broad and rounded, with no obvious anterior lappets. Pereonite 4 with short, 1

narrow anterior lappet. Tergite 5 only slightly shorter medially than other tergites. Pleotelson with smoothly

w

rounded margins, lacking posterolateral spines or lobes; posteromedial tip rounded, most posterior part of

body. Antennula article 3 distinctly longer than wide; with 12-13 articles, distal article small, but easily seen. Jl

Antenna with conspicuous pointed scale. Mandibular article 3 elongate, longer than palp article 1. Articles 1- l

3 of maxillipedal palp expanded. Pereopod I carposubchelate; dactylus short, with 2 claws; propodus with 18

denticles on proximoventral border, carpus with 2 rows of short spine-like sensillate setae on ventral margin l_

(5 on lateral side and 18 on medial side) and f'me fringe of cuticular comb-like spinules. Coxa I triangular, II

pointed. Coxae II-V weakly bilobed, barely visible in dorsal view. Female operculum broad, ovate, with

median concavity distally. Male pleopods I-IJ unknown. Uropods unknown, probably biramous with branches

of similar length and sympod about as long as rami. Ill

Biology. See remarks under Janiralata sp. A I

Remarks. This species is very similar to Janiralata rajata Menzies, 1951a, especially in body form. I

J. sp. D differs from J. rajata in the following: the eyes are smaller, the body has numerous dense

chromatophores, the antennula is longer, andthe propodus of pereopod I has more proximoventral denticles •

(18 instead of 12 or 13). Nevertheless, this new specimen and the male holotype ofJ. rajata are near the same II

size. Males from the Santa Mafia Basin population must be collected to establish more differences.

Type Locality and Type Specimens. None. This species is undescribed. _l

Distribution. Santa Maria Basin, 91.5-123m. II

!

Family Joeropsididae Nordenstam, 1933

(originally Jaeropsini) 1

Genus Joeropsis Koehler, 1885

Description (derived from data of Wilson, 1994). Body broad; pereonal tergites laterally rounded, I

coxae not visible in dorsal view. Head wider than long; frons wide with distinct vertex, rostrum projecting

anteriorly between antennulae. Pleotelson with posterolateral spines; pleonite 1fused topleotelson. Antennular I

fh'st article length greater than width, with spines on distolateral margin, flagellum with less than 5 articles. II

Antennal flagellum with more than 12 articles but proximal articles conjoint; articles 5-6 longer than 1-4,

article 5 with broad lateral flange; article 3 not especially enlarged compared to articles 1-2 and 4, scale II

absent. Mandibular molar thin, elongate, distally tapering. Maxillipedal palp slender, endite extremely broad, II

more than twice as broad as palp. Pereopod I ambulatory, not subchelate, carpus and propodus with few

setae. Pereopods H-VII dactyli with 2 subequal claws, accessory seta claw-like. Penes situated medially on I

86 I
I

I

I

U A

I Figure 1.34. Janiralata sp. D (cf. rajata), MMS Phase I Sta. BRA-16, preparatory female. A, dorsal view of

body, scale bar = 1 mm. B, right pereopod I with enlargement of proximoventral denticles on

I propodus, lateral view.

i 8_
!

pereonite 7 sternum. Pleopod Ill[ endopod with 3 plumose setae; exopod narrower and longer than endopod, I

lacking plumose setae, with 2 free articles. Uropods squat, inserting terminally on pleotelson, rami distinctly

shorter than protopod, exopod shorter than endopod; enlarged protopod with distinctive medial claw. I

Joeropsis concava (Sehultz, 1966)

Figure 1.35 I

Material Examined.California, Santa Mafia Basin, off Purisima Point, Sta. BRA-16, 91.5-123 m _l

(5).--Western Santa Barbara Channel, Sta. BRC-2, 120-123 m (3). II

Description. Body devoid of pigment, but covered with numerous fine simple setae. Body elongate

andparallel-sided; all segments width subequal, body length exceeding 4 times body width. All lateral margins _l

with small denticulae, and pleotelson with 7 large posteriorly curved spines. Cephalic lateral margin with W

distinct indentation below eyes; eyes small, with fewer than 10 ocelli. Antennula with 5 distinct articles

(Schultz, 1966, reports 6 - see remarks below) article 1 with large distolateral projection beating around 6 i

sharp, flat spines curving toward article 2; article 2 distinctly longer than wide. Male pleopod II with distinctly II

separate medial and lateral lobes; medial lobes longest, with medially converging sides. Uropodal protopod

with distinct distomedial spine; rami with short setae no longer than protopod. _!

if Biology. Living joeropsidids look like slow-moving little tanks. They are relatively well calcified

compared to other more speedy shallow-water janiroideans, such as laniropsis. Shallow-water species are

usually well pigmented although this deep-water form seems to lack any pigment. In

II

Remarks. Several species of Joeropsis are substantially similar toJ. concava in many details, especially

J. brevicornis Koehter, J. dubia Menzies, t 951, and J. paucispinis Menzies, 1951 (varieties and subspecies am

of previous authors should be considered separate species unless clear evidence for their conspecific nature is l

I

given). J. concava differs from all of these species by having a more elongate body, no pigment, and a

distinctive indentation in the cephalic margin below the eyes. /11

The specimens illustrated here from the Santa Mafia Basin conform with Schultz's (1966) description i

ofJoeropsis concava in practically all points, but their identification must be considered tentative because the

antennulae do not match. Schultz (1966) illustrates a6 articled antennula with the terminal sixth article being Jim

elongate and pointed, while these specimens clearly have 5-articled antennulae, with the presumptive sixth I1

article being fused to the 5 article. Some joeropsidids may have a free sixth article, but it is always tiny and

truncate, never elongate and pointed. Other species illustrated in Schultz (1966) are sometimes inaccurate on Ill

details of this scale, so the illustration and his description may be erroneous. Alternatively, the type specimen 1

could be simply a deviation from the common joeropsidid form. The types of this species must be examined to

settle this problem. I

The ventral surface of this species showed distinctive ridges on the ventral midline, that were not i

present inJ. sp. A (see next description). Formal descriptions ofjoeropsidid species nearly always ignore the

ventral surface of the body, so these features could not be included here. For the future, however, taxonomists IB ]

of this morphologically conservative group should be aware that important characters may be obtained from I

the lateral and ventral sides of the body.

Type Locality and Type Specimens. Holotype male, 3.1 mm long, R/V VeIero IV station 6806: I

33°56.1'N, 118°52.28'W, Santa Cruz Canyon, 221 m, 22 December 1959, rocks and coarse green sand (from !1

Schultz, 1966). j

Distribution. Central to southern California, 60-221m. Additional specimens of this species have i 

been collected by the City of San Diego Ocean Monitoring Program at Station B5 (32°49.25'N, 117°19.60%V, II

60m).

I
!

!

I A

II

II

II

II

II

I

I1

1 i: ,;/

_ t _ kk

!

F_tre 1.35. JoeropsisconcavaSchultz, 1966, male f_mPhase I Sta. BRA-16. A, bodydorsalview, scalebar=

I 1 mm. B, antennula. C, male pteopod I with enlargement of distal tip.

!

I _9
!

Joeropsis sp. A I

Figure 1.36

II

Material Examined. California, Santa Maria Basin, off Purisima Point, 90-130.5 m, Sta. BRA-20, I

90-130.5 m (1); off Point Arguello, 54-63 m, Sta. BRA-6, 4 individuals, illustrated brooding female and

copulatory male. i

Description. Body devoid of pigment, but covered with numerous fine and some longer simple setae.

Body broad and laterally curved; pereonites 3 and 4 widest, body length between 2.5 and 3 times body width; I

all pereonites with tergites extending much further than ventral insertions of pereopods. All lateral margins I

with small denticulae, pleotelson with 8 large posteriorly curved spines. Cephalic lateral margin with large

anteriorly curved spines, substantially produced laterally forming large lateral plate; eyes small, with I1

approximately 10 ocelli. Antennula with 5 distinct articles; article 1with small distolateral projection bearing 1

several laterally curving fiat spines; article 1 with small medial projection with several denticulae; article 2

distinctly longer than wide. Male pleopod II with distinctly separate medial and lateral lobes; medial lobes n

longest, with medially converging, straight sides; medial margin of lateral lobe distinctly arc-like. Uropodal !

protopod with distinct distomedial spine; rami with elongate setae longer than protopod.

I

Biology. See general remarks above, J. concava. •

Remarks. This new species is most likeJoeropsis lata Kussakin, but differs in the following respects II

(J. lata in parentheses): the antennulae have large distinct denticulae (only tiny denticulae) and article 2 is

much longer than broad (length and width approximately equal); male pleopod I medial lobe on the distal tip i

straight sided (not curved). This species of Joeropsis is also similar to J. setosa George and Strrmberg, II

although the latter is largely denticle-free on all margins, and seems to have more obvious dorsal setae. George

and Str6mberg (1968) provide a useful key to species of Joeropsis, although it is out of date now that the I

number of species in the genus has nearly doubled. II

Type Locality and Type Specimens. None. This species is undescribed.

Distribution. Santa Maria Basin, 54-130.5 m. I

Family Munnopsidae Sars, 1869 i

i

Subfamily Eurycopinae Hansen, 191 i

Genus Eurycope G.O. Sars, 1864 .!

Description (from Wilson, 1982b). Eurycopinae with deep, vaulted and rounded natasome; venter

with no enlarged or recessed area; body without dorsal or lateral spines. Tergat articulations of pereonites 5- i

7 distinct; pereonite 7 subequal to or longer than pereonites 5 and 6. Rostrum and frons distinct; clypeus II

narrow, striplike; labrum longer than clypeus. Pleotelson posterolateral margin parallel to pteotelson longitudinal

axis or angled downward in lateral view. Antennular f'u-starticle broad, with well-developed medial lobe. nit

Mandibular molar triturating surface broad, oval, with tiny denticles and small setae on posterior edge; II

ventral edge flattened into angular blade. Mandibular palp well-developed and functional; flattened distal

article strongly curled laterally. Bases of pereopods I-IV subequal to body depth. Bases of pereopods V-VII I

subequal, short, and robust. Uropods short, biramous; protopod broad or tubular, not leaflike; exopod subequal II

to or shorter than endopod.

i
I

I

I A

I

I

I B

I

I

t

I 1 \

i .....

i _ ./ ::y _::!: :

I _! • :_"../ , _" J. _, _"' ,,,-"_1I,

t

!

Figure 1.36. Joeropsis sp. A. A-B, brooding female from Phase I Sta. BRA-6. A, body dorsal view, scale bar =

I s1tat ramion.. B, antennula. C, male pleopod I with enlargement of distal tip, male specimen from same

I

| 9_
!

Eurycope cahforniensis Schultz, 1966 I

Figure 1.37 I

Eurycope californiensis: Schultz, 1966:4, 8, 28-29. I

Material Examined. California, Santa Maria Basin, off Point Sal, Sta. R-7, 565 m (1 female, I

illustrated; another female); off Purisima Point, Sta. 50, 591 m, (3, illustrated male); off Point Arguello, Sta. $

63, 930 m, (1 female).

Description. Eurycope (Eurycopinae) with deep, vaulted, and rounded natasome. Venter of natasome I

with no enlarged or recessed areas; tergal articulations dorsally distinct. Pereonlte 7 medially longer than I

pereonites 5-6; bullae on the ventrolateral surface of pereonite 7 indistinct. Pleotelson posterolateral margin

only slightly angled downward compared in lateral view to anterolateral margin. Rostrum of cephalon broadly I

acute, anteriorly rounded, only slightly overhanging frons, lacking setae, spines, or lateral keels. Body without II

dorsal or lateral spines. Antennular first article broad, medial lobe with around 4 sensillate setae, lateral lobe

with short, conical setae. Mandible with narrow molar process having numerous distal denticles surrounding I

triturating surface. Mandibular palp large, well-developed, with distal article strongly curled laterally. Maxilliped I

with normal 5 articled palp. Pereopod I-IV bases subequal and elongate, lengths longer than axial length of

cephalon and pereonites 1-4. Pereopod V-VII bases robust, short and subequal, basis VII with only one small 1

seta. Male pleopod II protopod lateral fields only slightly recurved ventrally; distal tip with short narrow I

projection. Uropods robust and bimmous; protopod with modest medial lobe and distoventral fringe of sensillate

setae, length subequal width; exopod longer than half length of endopod; both rami with distal tufts of sensillate I

setae; endopod with several penlcillate setae. l

Biology. Genera and species of the Eurycopinae are strongly swimming, epibenthic animals, usually

found on soft deep-water substrates. Hessler and Str6mberg (1989) review most of what is known about their I

behavior. Compared to other asellotes, members of the genus Eurycope are often large, with much higher O

biomass concentrations than other more abundant but smaller species. Analyses of their population biology

(Wilson, 1981, 1983a-b) suggests that some abyssal members of Eurycope may grow rapidly to an adult size I!

(as evidenced by the rarity of intermediate life stages) and reproduce multiple times as adults (shown by I

various sizes of copulatory adults as well as a strong and broad modal distribution of adults).

Remarks. Eurycope californiensis resembles members of the Eurycope compIanata complex (see I

Wilson, 1983a). However, it differs from that group in the following features: rostrum distally rounded, U

lacking a distinct anterior notch; no distinct bullae on the ventrolateral surface of pereonite 7; lateral fields of

pleopods II not strongly recurved; and uropodal protopod not broader than long. Schultz (1966) described this 1

species with a reduced palp. Actually, the palp is quite normal but is often broken off during collection and u

processing. The specimens that lack the palp can be seen to have a distinct socket where the palp was prior to

breaking off. I

Type Locality and Type Specimens. Holotype female, body length 3.5mm. R/V Velero IV station

7032. Newport Canyon, 33°31.29'N, 117°54.95'W, 478m. Collected with green mud and gray sand, some

very coarse. I

Distribution. Newport Canyon to Santa Maria Basin, upper and middle slope depths, 478 to 930m.

I

I
!1

II

I

!1

II

II

,,

!

,I

I

I Figure 1.37. Eurycope californiensis Schultz, 1966. A-C, copulatory male (Phase I Sta. 50), dorsal lateral views of body (scale bar = 1 ram), and enlargement of antennular basal article.

D, preparatory female (Phase I1 Cruise 3-4, Sta. R7), left uropod, ventral view.

!

!

!
!

Genus Belonectes Wilson and Hessler, 1981 I

Description (from Wilson and Hessler, 1981). Eurycopinae with elongate, streamlined natasome;

venter with pronounced medial bump at posterior margin of pereonite 5, becoming abruptly recessed dorsally •

posterior to bump. Pereonite 4 with distinct ventromedial bump. Pereonites 5 and 6 fused dorsally, pereonite I1

7 smaller and narrower than combined pereonites 5 and 6. Rostrum of cephalon broad, quadrate, sloping

directly into very low frons; clypeus dorsally prominent, approximately half length of labrum. Pleon I

posterolateral margin angles distinctly upward in lateral view. Antennular ira-starticle trapezoidal, lacking

medial lobe. Mandibular molar thin, obliquely truncate distally; ventral margin flattened into angular blade;

posterior margin with tiny denticles and small setae. Mandibular condyle shorter than molar process. Mandibular _l

palp distal article somewhat reduced, curled approximately 90 ° laterally. Bases of pereopods I-IV elongate U

and subequal; bases of natatory pereopods differing from one another; pereopod V basis shortest and widest,

pereopod VI longest but intermediate in width, pereopod VII basis thinnest but intermediate in length. Uropods _l

short, protopod biramous tubular; exopod tiny; endopod more than twice length of protopod. J

Belonectes sp. A I

Figure 1.38

I

Material Examined. California, Santa Maria Basin, off Point Arguello, Sta. 72, 401 m (1 preparatory l_

female, illustrated).

Description. Belonectes (Eurycopinae) with elongate, streamlined natasome, distinctly narrowing at I

pereonite 7. Rostrum of cephalon broad, sloping medially to low frons but with laterally flaring anterior lobes

n

with thin fringes. Clypeus of cephalon prominent in dorsal view. Ventromedial surface of pereonite 5 with I

distinct bump, becoming recessed posterior to bump. Pereonites 5 and 6 fused dorsally, pereonite 7 smaller 1

Nil

and narrower than pereonites 5 and 6. Pleotelson posterolateral margin angling distinctly upward in lateral

view. Pleonal preanal ridge not enlarged and prominent. Antennular first article oval, wider proximally than

distally. Bases of pereopods 1-IV elongated and subequal. Pereopod bases V-VII differing in length and width: I_

basis V shortest and widest, basis VI longest, basis VII thinnest. Uropods short, biramous; protopod parallel qll

sided; exopod very small; endopod less than twice length of protopod. am

Biology. See remarks under Eurycope californiensis. Belonectes tend to be much smaller both in !

body size and in population size than Eurycope, and therefore comprise a much smaller part of the benthic

biomass. I

Remarks. Unlike any of the known species or undescribed species of BeIonectes from the Atlantic J

(see Wilson and Hessler, 1981), this species has a distinctive broad and laterally flaring rostral area of the

cephalon. Unfortunately the only specimen examined had a damaged pleotelson, so the illustrated shape •_

should not be trusted for accuracy. Within the Santa Maria Basin, this eurycopine species is distinctive with I

broad indented and laterally flaring rostrum, waisted pereonite 7 and narrow uropods with tiny exopods.

Type Locality and Type Specimens. No type locality - this species is undescribed. I_

Distribution. Santa Maria Basin, 401 m.

!

!
!

!

!

II

I Figure 1.38. Bel baronec = 1tesram.sp.C,A,cepprhepalon, aratorylateral femal obe,liquPheasvieeIwSt. a.D,072. rightAuropod, -B, bodyventr , doralsalview. and lateral views, scale

!

!

!

!

i 95
!

Subfamily Ilyarachninae Hansen, 1916 I

Genus Ilyarachna Sars, 1870 n

Description. Head much broader than long, with flattened frons, lacking rostal projections, antennulae

medially adjacent; clypeus and labrum reduced, much narrower than head. Body with distinct natasome, Ill

natasome without stout dorsal setae. Pieonite 1distinct dorsally. Pleotelson dorsally subtriangular. Antennulae |

with multiarticulate flagellum. Mandibular incisor process robust, teeth broad rounded; molar process reduced,

distally tapering and setose; dorsal condyle elongate, curving; palp shorter than mandibular body, with 3 il

articles. Pereopods I-II length subequal, both prehensile. Pereopods Ill-IV extremely long, ambulatory, bases II

short and robust, lengths nearly as short as wide. Pereopods V-VI carpus-propodus paddle-like, fringed with

plumose setae. Pereopod VII carpus-propodus narrow, but fringed with plumose setae. Uropodal protopod

flattened and foliacious, longer than rami; exopod tiny or absent. II

i

llyarachna acarina Menzies and Barnard, 1959 U

Figure 1.39

gm

Ilyarachna acarina: Menzies and Barnard, 1959:9-10; Menzies, 1962:136; Wolff, 1962:96-97; Schultz, I

1964:310; Schultz, 1966:9; Hessler and Thistle, 1975:157; Thistle, 1979:382-384.

Material Examined. off Purisima Point, Sta. R-5, 154 m (14, illustrated 1 large female with most I

legs intact; 10 from another replicate; 3 damaged specimens from another sample).--Westem Santa Barbara

Channel, Sta. 87, 299 m (2 damaged specimens). 1

Description. Head short and broad, with decidedly projecting lateral cheeks bearing mandibular I!

articulations; cephalic dorsal vertex with 10-15 stout setae in adult. Pereonites I-4 freely articulated, with 11-

16 large pedestal setae on anterior dorsal margins; pereonite 4 with distinct anterolateral projection with l[

several setae. Pereonites 5-7 with visible sutures, narrowing posteriorly. Tergites 5-7 and pleotelson with II

scattered fine setae dorsally. Pereonal sternum lacking spines. Pleotelson subtriangular in dorsal view, with

uropods projecting posteriorly. Antennula article 1 with several setae; article 2 length subequal article 3. •

Antenna basal article with lateral projection having 3-4 setae. Mandible heavily sclerotized, incisor process II

massive, molar process reduced to small setose flap; mandibular palp present but reduced; distal article not

cuded, paucisetose. Pereopods I-II short, somewhat prehensile, pereopods III-IV extremely elongate with I

bases only slightly longer than wide. Pereopod 1 coxal plate with 2-3 setae; pereopod 2 coxal plate with 1-2 l

setae. Pereopod 2 basis fringed by long setae on anterior and posterior surfaces. Pereopods V-VII decreasingly

natatory toward the posterior end of the body, with pereopod VII nearly completely ambulatory. Female •

operculum with setose median keel. Male pleopod I elongate, tapering to narrow distal tip with narrow projecting II

lateral lobes and non-projecting medial lobes with 6-7 setae. Uropodal protopod broad, flattened, leaf-like;

exopod completely fused to protopod and reduced to two setae on protopod.

Biology. Ilyarachnines are interesting because they are known to burrow despite their munnopsid U

abilities of swimming (Hessler and Strtimberg, 1989). They walk on soft substrates with on elongate pereopods

[] and IV, probing with the anterior 2 pairs of pereopods. They can burrow into the mud posteriorly, pushing •

sediment with their "natatory" pereopods, and are often found at rest with their entire body in the mud with II

only their long antennae and pereopods []-IV protruding radially from surface. If sufficiently disturbed, they

will take to the water with short spasmodic strokes. This, however, does not last for long and they will settle II

to the bottom and either walk away or burrow into the sediment. Like Munnopsurus, their powerful jaws U

suggest that they can feed on shelly or hard food items. Males seem to be much smaller than females in this

species: the illustrated preparatory female was 3.9mm long, while a fully adult male was only 2.0ram long. I

96 I
!

!

I _A

I +

I ,

I iI ,,,,,,I

I i, ",i

.i 1 ,! r. I

I c _

!

Figure 139. Ilyarachna acarina Menzies and Barnard, 1959. A-B, body lateral anddorsal views, scale bar = 1

I female from MMS Phase II, Cruise 3-4, Sta. R-5. C-D, male pleopod II ventral

iflira, preparatory

view and distal tip, male from same station.

I

Remarks. Thistle (1979) provides sufficient information to separate llyarochna ocarina from L

i profunda two speciesSchultz, is the large 1966,pedestal both foundsetaeinonCalif theornian anteriorwaters. marginTheofbpereonites est character1-4that in Lallows acarinadis; tinguishing I. profunda,bohasth

much smaller non-pedestal setae on anterior pereonite margins. Menzies and Bamard (1959) thought their

i spec on theies anterior was similar margins, to L denticulata and has distinct Sars, 1897b, anterolateral althoughprojections I. ocarinaonhaspereonite considerab4.lyMancas larger ofpedestal L acarina setae,

however, lack pedestal setae or have few of them, so one should check the developmental stage of specimens

i before arriving Type Loatcalanityidenti andfType ication.Specimens. Holotype brooding female, 4.3mm long, Allan Hancock Foundation

collection number 578. Off coast of California, R/V VeleroSta. 4980, 34°15.8'N, 119°34.5'W, 92m.

I Distribution. Off Southern California from Santa Maria Basin to off Pt. Loma, 73-1118m.
!

Subfamily incertae sedis I

Genus Munnopsurus Richardson, 1912 I

Description. Head broad, vaulted, frons flattened with heavily cuticularized frontal arch; clypeus

angled sharply dorsally, sublriangular in anterior view, laterally extending near lateral margins of head; vertex

distinctly separating antennular insertions dorsally. Pereonites t-4 freely articulated. Body with distinct |

natasome, natasomal somites free, including pleonite 1.Pleotelson broad, rounded, often inflated. Antennular

article 1 flattened, longer than wide, lacking distinct medial lobe. Mandibular incisor molar process broad, I

heavily cuticularized, wider than long; dorsal condyle elongate, curving. Pereopod I slender, not prehensile. I

Pereopods II-IV elongate, thin, paucisetose, subequal; bases many times longer than wide. Pereopods V-VII

carpus-propodus paddle-like, fringed with plumose setae. Uropod small, biramous; protopod tubular, shorter I

than rami. |

Munnopsurus sp. A I

HI

Figure 1.40

/

Material Examined. California, Santa Maria Basin, offPoint Buchon, Sta. 15,393 m (1 preparatory I

female); offPoint Sal, Sta. R-9, 410 m (3, including 2 males, 1 with Per IV, and I female; second sample with

3 individuals: 1 male, 1mashed female, and 1 manca stage 3); Sta. R-7, 565 m (1 brooding female); off Point

Arguello, Sta. 62, 582 m (1, brooding female). II

Description (generic characters modified from Wolff, 1962). Small body size, adults with body

length around 1.8 ram. Head as broad as pereonite 1, vaulted posteriorly, with flattened frons; frontal arch 1

strongly calcified, semicircular; labrum and clypeus triangular. Dorsal surfaces of body with scattered fine II

simple setae. Pereonites 1-4 free and loosely articulated. Pereonites 5-7 lengths subequal and freely articulated.

Pteotelson rounded in dorsal view, somewhat inflated. Antennula with no medial projection, distally flattened; I1

flagellum in both sexes short with few articles, antennular article count 15 or less; article 2 with no medial U

projection, inserting around quarter length of article 1 from distal tip. Molar process of mandible broad and

simple, but distinctly projecting; mandibular palp article 3 fully functional, curved and setose. Maxillipedal It

second article with strongly oblique anterior margin. Pereopod I not prehensile, elongate and thin. Pereopods I

II-IV long and slender, more than twice body length. Pereopods V-VII short, with carpi no longer than depth

of body. Male pleopod I elongate and narrow, distinctly waisted proximally and broadening distally; lateral I

lobe on distal tip broadly recurved, projecting distally approximately half distance of medial. Male pleopod II l

with endopod and exopod reduced, placed distally in last fifth of protopod. Uropods short, biramous, somewhat

inflated; exopod distinctly longer than protopod. II

Biology. Little is known about the biology of species of Munnopsurus. Their somewhat reduced n

natapods suggests a reduction in the swimming lifestyle, while their elongate legs (typical in munnopsids) is in

line with a soft bottom, deep-water habitat. The distinctive feature of the genus, the powerful mouthparts, i

suggests an adaptation for crushing large hard objects, such as foraminifera (as suggested forAmuletta; see II

Wilson and Thistle, 1985). This species has more eurycopine-tike mouthparts and therefore is probably

somewhat more of a generalist. I1

II Remarks. Munnopsurus sp. A is most similar to M. minutus Gurjanova, 1933, although it differs by

being distinctly smaller with body lengths around 2 mm (a 7ram deep-sea asellote isopod is notminute, so one

wonders why Gurjanova gave it this name), by the lack of dorsal tubercles on pereonites 5-7, by having a

somewhat inflated and rounded pleotelson, and in having somewhat more articles in the antennula. M. minutus U

Gurjanova, M. atlanticus Beddard, andM. sp. A are some what different from the remainder of Munnopsurus

I
I Figure 1.40. Munnopsurus sp. A. A-B, body dorsal and lateral views, scale bar = 1mm, copulatory male,Phase

II, Cruise 1-2, St,xR-9. C, right antennula. D-E, male pleopod I, ventral view with enlargement of

I distal tip. F,left uropod, ventralview.

am

I species having a more plesiomorphic eurycopine-like a palp a

in mandible with functional mandibular and

strengthened but not massive mandibular molar and incisor. See remarks under Munnopsurus sp. B for

separating the two congeners within the Santa Maria Basin.

I Type Locality and Type Specimens. None. This species is not described.

Distribution. Santa Maria Basin, 393-582m.

I

i 99
!

Munnopsurus sp. B i

Figure 1.41 J

Material Examined. California, Santa Mafia Basin, off Point Arguello, Sta. 63, 930 m (2, adult 1

male, juvenile female, both specimens squashed by labels in vial).

Description (generic characters modified from Wolff, 1962). Large body size, adults with body g

length around 7 mm. Head huge, distinctly broader than pereonite 1, vaulted posteriorly, with flattened frons; ill

frontal arch heavily calcified, semicircular, distinctly projecting; labrum and clypeus triangular. Dorsal surfaces

of body smooth, with few simple setae. Pereonites 1-4 free and loosely articulated. Pereonites 5-7 lengths I

subequal and freely articulated. Pleotelson rounded in dorsal view, somewhat inflated. Antennula with no II

medial projection, distally flattened; flagellum in both sexes elongate, antennular article count much more

than 15 (around 45 in male); flagellar articles of male conjoint proximally, intermediate articles wider than I

long; article 2 with small medial projection, inserting around half length of article 1 from distal tip. Mandible I!

huge, heavily cuticularized; incisor process blunt; molar process broad and simple, not projecting; mandibular

palp article 3 thin, reduced, not curved or setose. Maxillipedal second article with strongly oblique anterior i

margin; basis with 6 coupling hooks. Pereopod I not prehensile, elongate and thin. Pereopods II-IV long and

slender. Male pleopod I elongate and narrow, more or less parallel-sided; lateral lobe on distal tip laterally

straight, not recurved, projecting distally as far as medial lobe. Male pleopod ITwith endopod and exopod i

reduced, placed distally in last fifth of protopod. Uropods thin, biramous; exopod approximately same length II

as protopod.

Biology. See remarks on the biology ofMunnopsurus sp. A. The huge jaws of this species indicates n

they can crush fairly heavy things. N

Remarks. Munnopsurus sp. B is somewhat similar to M. longipes (Tattersall, 1905), although the U

male pleopods are different. Unlike Munnopsurus giganteus (Sars, 1879) or M. ochotensis (Gurjanova, i

1933), M. sp. B has a larger and wider cephalon (Both the fh-sttwo taxa are considered by most authors to be

u

subspecies ofM. giganteus (Sars), although there is no evidence for their being conspecific. Therefore, they

should be thought of as distinct species.). The huge mouth parts ofM. sp. B makes this species a member of I[

Munnopsurus sensu stricto. Although not listed in the description, the dorsal mandibular muscle insertions UP

attachments provide an easy way to separate cephalon fragments of M. species A and B: if they are few and i

large, the fragment is sp. A, and if they are many and small, it is sp. B. This sorting character will work only i

within this basin where these two species may appear in the same series of samples. Outside of the Santa J

Mafia Basin, other species may have similar patterns of dorsal mandibular muscle insertions.

Type Locality and Type Specimens. None. This species is not formally described, i

Distribution. Santa Maria Basin, 930 m.

I

!

!

!

!
I Figure 1A1. Munnopsurus sp. B. A, copulatory male (Phase I, Sta. 63), dorsal view, pleotelson damaged, scale

bar = 1 mm. B, left antennula. C, male pleopod I, ventral side, and enlargement of distal tip. D,

i right uropod, ventral view.

!

!

!

i 101
!

Family Nannoniscidae Hansen, 191 I

m

Genus Nannonisconus Schultz, 1966

Description. (from Schultz, 1966, 1969 and Siebenaller and Hessler, 1981) Antennula extending

only slightly anterior of cephalon; 5-segmented, with bulbous distal article. Cephalon without anterolateral I

lobes. Pereonite 7 fused with pleon. Pleon of single somite and at widest point wider than cephaton or pereon. |

Lateral outline of body concave. Male pleopods I not apicalty pointed; proximal part much narrower than the

medial or distal part. Uropods biramous. I

Nannonis¢onus latipleonus Sehultz, 1966

Figure 1.42 I

Nannonisconus latipleonus: Schultz, 1966:3, 11, 35; Siebenaller and Hessler, 1981:248-249. II

Material Examined. California, Western Santa Barbara Channel, off Point Conception, Sta. 81,294 •

m (1 manca stage 3 individual).

Description. Broad-bodied Nannoniscidae with pereonite 6-7 and pleotelson fused medially. Rostrum

distinct with small lateral keels. Lateral flanges on margins of pereonites and pleotelson. Pleotelson with

distinct posterolateral spines, somewhat confluent with lateral flanges; anal region distinctly projecting from n

remainder of pleotelson. Venter of pereortites without large spines, although pereonite 6 may have smatl spine •

projecting from anterior margin. Pleon at widest point wider than cephalon or pereon. Lateral outline of body

concave in dorsal view. First antenna with 5 articles, distal article bulbous. Female pleopod large, covering n

almost entire ventral surface of pleotelson. Uropodal exopod more than two thirds length of endopod. I

Biology. Typical for all nannoniscids, Nannonisconus latipleonus is likely to be a generalized detritivore

that lives on soft bottoms or perhaps is associated with biogenic substrates (Levin et al., 1986). Sexual m

dimorphism is common in the family, and although the manca stage 3 specimen illustrated here is largely I

similar to the male specimen illustrated by Schultz (1966), a fully mature male may have broader lateral

flanges on the pereonites and pleotelson. Although detailed quantitative surveys have not been made at the m

localities described herein, the rarity of specimens (2 to date) suggests that this species is not a dominant i

component of the benthic community.

Remarks. Nannonisconus carinatus Mezhov, 1986 may be conspecific with N. latipleonus. Schultz •

(1966) does not appear to have described a fully mature male because the appendix masculina on the pleopod •

II, as illustrated in his plate 6, is short and blunt. The appendix masculina should be long and distally thin in

a fully mature male as in N. carinatus. All apparent differences between the two species, such as broader m

lateral flanges on the body, could be explained by ontogenetic changes taking place during the maturation to II

the male moult. Neither type has been examined at this point so it is not possible to demonstrate this with

certainty, although the reader should be aware of this problem when making identifications. If the two names n

are indeed conspecific, then the total distribution extends from Southern California to Alaska Bay. |

Type Locality and Type Specimens. Holotype male, 2.8 mm long, R/V Velero IV station 2793:

33°48.0'N, 118°32.0'W, Redondo Canyon, 465 m, 22 May 1954, blue-gray mud and large rocks (from Schultz, m

1966). I!

Distribution. Shallow bathyal slopes of Southern and Central California (but see remarks above),

collected at only 2 stations 190 km apart, m

!

102 I
!

!

II A B

I

!

I

I

I

I

I

I

I Figure 1.42. Nanno 1.5 mm, nisconus scale bar Iatipleonus = 0.5 mm. Schultz, A, body, 1966.dorMant sal aview. stageB,3 body, specimelateral n, Phase view.I Sta. C, 81, enlargement body lengthof

antermula and base of antenna, in situ on head.

I

I

I

I

!

i 103
!

Family Desmosomatidae Sars, 1897 i

II

Genus Momedossa Hessler, 1970 i

Description. (from Hessler, 1970) Pleotelson with posterolateral spines, in female widest anteriorly.

Uropod biramous. Cuticle of body well sclerotized. Pereonites 1 and 2 subequal in size. Pereopods long. il

Pereopod I moderately reduced; carpus and propodus somewhat attenuated; propodus without major setae; II

carpus with row of major setae only ventrally. Coxae of pereopods I-IV in female only moderately produced.

Sexual dimorphism modest; in known copulatory males (M. profunda, M. symmetrica) coxae of pereopods I- •

IV somewhat more strongly produced and pleotelson widest at posterolateral spines. I

I

Momedossa symmetrica (Schultz, 1966) In

Figure 1.43

Desmosoma symmetrica: Schultz, 1966:3, 5, 7, 26-27. 1

Momedossa symraetrica: Hessler, 1970:62, 77.

Material Examined. California, off San Francisco, Gulf of the Farallones, numerous specimens 1

collected Aug 1990, R/V Farnella, by J.A. Blake, as part of U.S. Navy 103 survey of proposed deep-water

disposal site: Sta. S-19, 37°38.41'N, 123°24.36'W, 2580m, 13 Aug 90; Sta. 1-19, 37°38.38'N, 123°27.77'W, m

2835m, 13 Aug 90; Sta. E-19, 37°38.40'N, 123°29.21'W, 2955m, 14 Aug 90; Sta. K-15, 37°39.38'N, i

123°27.23'W, 2830m, 14 Aug 90; Sta. E-07, 37°41.64'N, 123°29.16"_V,2462m, 15 Aug 90.

i

Description. Head with spines projecting lateral to antennulae and antennae; dorsal surface of head •

without longitudinal furrows. Pleotelson with posterolateral spines; pteotelson sexually dimorphic: broadest I

anteriorly in female, posterolateral spines larger and more distinct in male, and pleotelson more vaulted in

male. Cuticle of body well sclerotized. Pereonites 1 and 2 subequal in width and length. Pereonite 5 laterally i

convex in dorsal view. Coxae ofpereopods I-IV with distinct anterior spines, larger in male. Pereopod lengths l

distinctly greater than half body length. Pereopod I only moderately reduced but distinctly smaller than pereopod

II, propodus without major setae; carpus with ventral row of major sensillate setae only, most distal sensillate am

seta longer than adjacent penultimate sensillate seta (synapomorphies of the Desmosomatinae). Pereopods II- |

HI similar, propodi and carpi somewhat broadened with rows of both dorsal and ventral sensillate setae.

Ventral margins of meri and ischia I-Ill with no more than 3 sensillate setae. Pereopods V-VII carpi and i

propodi with distinctly fewer than t0 setae. Uropod biramous, exopod tiny compared to long thin endopod, i

Biology. Desmosomatids are epibenthic or shallow burrowing detritivores (Thistle and Wilson, 1987;

Hessler and Str6mberg, 1989), a pattern expected for Momedossa symmetrica. The anterior limbs may be •

used for burrowing in the sediment, because they are held laterally and have many lateral facing setae. This I

species is abundant compared to other deep water Janlroidea, and may be a dominant berLthic taxon in the

Santa Mafia Basin. Sexual dimorphism is apparent in this species, resulting in the female being larger than the 1

male, a typical pattern for many skinny desmosomatids. i

Remarks. Although this species does not appear in the voucher collection for the Santa Maria Basin,

it is included to prevent further misidentifications of the species. The male illustrated here adds to the female •

illustrated by Schultz (1966). Desmosomatids should be viewed on the lateral side, especially examining the II

head where useful characters are typically found, such as the cephalic spines. These cephalic spines are the

chief feature to identify this species from most other desmosomatines, as well as from Momodossa profunda •

Hessler, 1970. Other features separating M. symmetrica from M. profunda are the convex lateral margins of I

pereonite 5 in dorsal view, the generally fewer setae on the pereopods and the longer posterolateral spines on

I

104 i
!

!

I

I

I

i

I

I

i

\

:_ 'E

II D _. ..

I

II

I Figure 1.43. Momedossa symmetrica (Schultz, 1966), copulatory male (Navy 103 station). A-B, body, dorsal

and lateral views, scale bar = 1 nun. C-D, pereopods I and II, right, lateral view. E, uropod, right,

I ventral view.

!

!

i 105
!

the pleotelson. The uropodal exopods, however, are similar. Biologists new to identifying Desmosomatidae N

should also know that this species appears to be similar toEugerdella cornuta Hessler, 1970, primarily due to

the cephalic and pleotelsonic spines, as well as the fewer setae on the pereopods (compared to M. profunda). •

The features of the first pereopod, such as its size and its setation that separate the Desmosomatinae from the |

Eugerdellatinae apply here quite clearly: the pereopod I is smaller than pereopod II and the pereopod I carpus

has a distal ventral seta that is larger than the penultimate seta. The opposite situations apply for Eugerdella It

cornuta. Another desmosomatid found in the Santa Maria Basin (Desraosoma sp. A), is easily separated from |

this species by the lack of the uropodal exopods and the cephalic spines.

Type Locality and Type Specimens. Holotype brooding female, body length 3.2 mm (no paratypes). I

Tanner Canyon, R/V Velero IV station 2793: 32°36'N, 119"05.3'W, 469m, 29 January 1960, green mud with Ii

shale fragments (Schultz, 1966).

Distribution. California continental slope and submarine canyons, 469-2955 m. I

Genus Desmosoma G.O. Sars, 1864

Description. (from Hessler, 1970)Pieoteison without posterolateral spines, broadest anteriorly. Uropod

uniramous; protopod may or may not be abundantly setose. Pereonite 1 moderately to much smaller than

pereonite 2. Pereopod I moderately reduced; carpus and propodus moderately attenuated; propodus devoid of •

major setae; carpus without major dorsal setae, but usually with ventral row of slender setae. Carpus of

pereopod II broad, abundantly setose. Sexual dimorphism moderate to slight; in copulatory males pereonites

5-7 and pleotelson may be broader; coxae of pereopods I-IV may be moderately more strongly produced. In •

female, coxae of pereopods I-IV only modestly produced.

mum

Desmosoma sp. A ][

Figure 1.44

m

Material Examined. California, Santa Mafia Basin, off Point Estero, Sta. 3, 291 m (preparatory U

female, brooding female); off Point Buchon, Sta. 14, Phase I Sta. 14, 299 m (3 brooding females).

Description. All specimens in poor condition. Body long, attenuated, with thin cuticle. Frons sloping, !

mouth parts somewhat prognathous, without large frons-clypeal furrow (feature of the Desmosomatinae). U

Pereonite 1 distinctly shorter than pereonite 2. Pleotelson without posterolateral spines, at least in female.

Pereopods I-IV with triangular coxae in dorsal view, basis of pereopod I subequal in length and size to basis I

II. Bases V-VII enlarged, muscular. Uropod elongate and uniramous, with many penicillate setae on endopod I

and approximately 8-10 long coil-tip setae on lateral margin; uropod length greater than half pleotelson

length. I

Biology. Too little material of this species was available to comment here, other than to refer the

reader to the general remarks under Momedossa symmetrica above. mm

Remarks. These specimens were originally classified in the Santa Mafia voucher collection as I

Momedossa symmetrica, but are easily separated from that species by the lack in Desmosoma spp. of the

uropodal exopod and the lack of the cephalic spines seen in M. symmetrica. Although no specimens had I

pereopods, the specimens are provisionally classified inDesmosoma for the following reasons: (1) pereonite |

1 is much smaller than pereonite 2, suggesting that the associated pereopod is at least somewhat attenuated;

(2) the uropod is uniramous; and (3) the bases of pereopods V-VII are large suggesting that this species has I

enlarged natatory posterior pereopods, common among species of Desmosoma. This species, however, is •

unlike any other described species in Desmosoma, primarily because of its unusual setation of the uropod.

gll

Biologists studying this area should be aware that several undescribed species of Desmosoma sensu stricto I

106 I
I

Figure 1.44. Desmosomasp.A, Broodingfemale,dorsalview,composite illustration of severalspecimens(Phase

I I Sta. 14), scale bar = 1 ram. B, brooding female, lateral view, brood plates and mouth part detail

omitted. C, fight antennula, dorsal view. D, fight uropod, ventral view.

I are present in the California Continental Borderland. This species can be separated from another species

known to this author by a setose lateral margin of the uropod, a smaller and narrower body, and a smaller

I pereonite 1 in comparison with pereonite 2.

Type Locality and Type Spedmens. This species is not formally described.

i Distribution. Currently known only from the Santa Mafia Basin, ca. 290 m.

i 107
!

Genus Prochelator Hessler, 1970 i

Description. (from Hessler, 1970) First pereopod large, chelate; dactylus and enlarged, specialized

propodus together comprising movable finger which acts in opposition to immovable claw formed by large I

seta on distal end of ventral margin of enlarged carpus. Carpus usually not produced at base of claw. Ventral

margin of carpus with two accessory setae: a short, stout, unequally bifid seta located midway along margin

and somewhat longer, more slender seta located at base of fixed claw,just proximal to it. Pereonite 1as large m

as or larger than pereonite 2. Pleotelson with posterolateral spines, sometimes obscure. Uropods nearly always

biramous. Most dorsal tooth on lacinia mobilis reduced or absent; next tooth in line considerably enlarged.

Joints surrounding palp segment 1 of maxilliped nearly straight; segment 3 with long lateral margin. Coxae of I

pereopods 1-IV anterolaterally angular, but not produced. Head with transverse ridge on frons and with fronsclypeal furrow.

II

Prochelator sp. A D

Figure 1.45 i

I

Material Examined. California, off San Francisco, Gulf of the Farallones, numerous specimens

collected Aug 1990, R/V Farnella, by J.A. Blake, as part of survey of proposed U.S. Navy 103 deep-water •

disposal site: Sta. S-19, 37°38.41'N, 123°24.36_V, 2580m, 13 Aug 90 (10); Sta. 1-19,37°38.38'N, 123°27.77%V, |

2835m, 13 Aug 90 (3); Sta. E-19, 37°38.40'N, 123°29.21_vV,2955m, 14 Aug 90 (1); Sta. F-17, 37°38.96'N,

123°28.83'W, 2945m, 14 Aug 90 (2); Sta. K-15, 37°39.38'N, 123°27.23'W, 2830m, 14 Aug 90 (2); Sta. L- •

17, 37°38.94'N, 123°26.85'W, 2855m, 14 Aug 90 (61); Sta. E-07, 37°41.64'N, 123°29.16'W, 2462m, 15 II

Aug 90 (29); Sta. J-13, 37°40.01'N, 123°27.45%V,2770m, 15 Aug 90 (1); Sta. N-13, 37°40.01'N, 123°26.06%V,

2623m, 15 Aug 90 (10).---Santa Mafia Basin, off Point Estero, Sta. 2, 200 m (2, female and male); off Port m

San Luis, Sta. 22, (2 brooding females, illustrated 1 specimen); offPurisima Point, Sta. R-5,154 m (74; plus l

1 male from another sample, illustrated).

Description. Head with pointed spines lateral to antennae; frons clypeal ridge and clypeal furrow •

small but present. Pereonite 1 distinctly larger than pereonite 2 (feature of Eugerdellatinae), with large ventral i

spine. Pereonites 3 and 7 with posteriorly curving spines. Pereonite 5 of male with anterolaterat posteriorly

curving spine. Pereonite 5 of female distinctly convex, lacking a recurved spine. Pleotelson posteriorly rounded, •

with large subdistal ventrolateral spines. Pereopod I with robust setochelate carpus not produced at base of 1

distal seta (feature of Prochelator), with penultimate accessory seta adjacent to elongate strong sensillate seta

on distoventral comer of carpus (synapomorphy of Eugerdellatinae); shorter sensillate seta in middle of carpus •

ventral margin. Pereopod II robust with 4 elongate distally fringed sensillate setae on ventral margin, and 2 II

simple setae on dorsal margin of carpus. Male pleopod II with distinct laterally recurved spine on distal tip.

Uropod biramous, exopod approximately third length of endopod with seta longer than uropod on exopod tip. •

Biology. Although members of this species are tiny, they are among the most abundant of the Asellota m

studied from the Santa Maria Basin. Their rather more powerful anterior pereopods and anteriorly blunt faces

suggest they are more strongly burrowing forms. See also general remarks underMomedossa symmetrica. 1

Remarks. This undescribed species of Prochelator is most like P uncatus Hessler, 1970, but has

several distinctive features that separate the two species. Most important, sexual dimorphism is evident in the

species, while P. uncatus is nearly monomorphic. Other features of this species not seen inP. uncatus include:

ventral midline spines on pereonites 1, 3, and 7; recurved lateral spines on the distal tip of male pleopod I; and

a convex lateral margin of pereonite 5 in dorsal view of the female. The recurved lateral spine on pereonite 5

includes a large amount of variability, both sexual andontogenetic. The spine is distinct in males, and largely

absent in females. Juvenile males may show a gradation from the absent (female) condition to distinct and

I

108 i
!

!

i I1| A _ _" l

| :,

II

a I

| a

! E

! ,

!

_ 1.45. Prochelator sp. A. A, brooding female (Phase I Sta. 22), body dorsal view, scale bar = 1 ram. B-e,

i copulatory mate, body lateral and dorsal view. D-E, pereopods I-II, right, copulatory male (Phase II, Cruise 3-4 Sta. R-5). F, uropod, tip of pleopod I and right pleotelson posterolateral spine,

copulatory male.

I

large. Biologists sorting samples from this area should be aware that another rare and undescribed species of

Prochelator was found at the Navyl03 site. This second species is easy to distinguish because it lacks the

I recurred spine on pereonite 5, has pointed coxal spines on pereopods 1-3, and has a uniramous uropod.

Type l_aw.ality and Type Speehnens. None. This species is undescribed.

I Distribution. Santa Maria Basin to off San Francisco, 99-2955 m.

I 109
!

Literature Cited I

m

Barnard, K.H. 1914. Contributions to the Crustacean fauna of South Africa. Annals of the South African •

Museum 10(11):325-446.

n

Benedict, J.E. 1897. A revision of the genus Synidotea. Proceedings of the Academy of Natural Sciences of •

Philadelphia 49:389 -404.

Benedict, J.E. 1898a. Two new isopods of the genus Idotea from the coast of California. Proceedings of the I

Biological Society of Washington 22:53-55. I

Benedict, J.E. 1898b. The Arcturidae in the U. S. National Museum. Proceedings of the Biological Society of

Washington 22:41-51. I

U

Birstein, I.A. 1973. Deep water isopods (Crustacea, Isopoda) of the north-western Pacific Ocean. Izdatel'stvo

Akademii Nauk SSSR, Moskva. [Translated from the Russian for the Smithsonian Institution and

N.S.F. by the Indian National Scientific Documentation Center, New Delhi. 316 pp.] I

Bonnier, J. 1900. Contribution h l'ttude des l_picarides: Les Bopyridae. Travaux de la Station Zoologique de

Wimereux, VIII. 396 pp. 1

m

Boone, P.L. 1918. Descriptions of ten new isopods. Proceedings of the U.S. National Museum 54:591-604.

Boone, P.L. 1923. New marine tanaid and isopod Crustacea from California. Proceedings of the Biological l

Society of Washington 36:147-156.

Botosaneanu L., N.L. Bruce, and J. Notenboom. 1986. Isopoda: Cirolanidae. In, L. Botosaneanu (Ed.), m

Stygofauna Mundi: A faunistic, distributional, and ecological synthesis of the world fauna inhabiting I[

subterranean waters (including the marine interstitial), pp. 412-421.

I

Bourdon, R. 1980. Les esp_ces du genreBopyrella J. Bonnier (Crustacea, Isopoda, Bopyridae). Bulletin de la •

Muste nationale d'Histoire naturelle, Paris, 4 str., 2, section A, 1:185-236.

Bowman, T.E. and G.A. Schultz. 1974. The isopod crustacean genus Munnogonium George and Stremberg, am

1968 (Munnidae, Asellota). Proceedings of the Biological Society of Washington 87(25):265-272. 1

Bran&, A. 1988. Antarctic Serolidae andCirolanidae (Crustacea: Isopoda). Theses Zoologicae. Koeltz Scientific

Books, Koenigstein. Vol. 10. 143 pp. 1

I1

Bruce, N.L. 1981. Cirolanidae (Crustacea: Isopoda) of Australia: Diagnoses of Cirolana Leach, Metacirolana

Nierstrasz, Neocirolana Hale, Anopsilana Paulian and Debouteville, and three new genera--

Natatolana, Politolana and Cartetolana. Australian Journal of Marine and Freshwater Research •

32:945-966. 11

Bruce, N.L., R.C. Brusca, and P.M. Delaney. 1982. The status of the isopod families Corallanidae Hansen, •

1890 and Excorallanidae Stebbing, 1904 (Flabellifera). Journal Crustacean Biology 2(3):464-468. l

Brace, N.L. 1984. A new family for the isopod crustacean genus Tridentella Richardson, 1905, with description 1

of a new species from Fiji. Zoological Journal of the Linnean Society 80:447-455. 1Illll

Bruce, N.L. 1986. Cirolanidae (Crustacea: Isopoda) of Australia. Records of the Australian Museum, Suppl.

6, 239 pp. E

Bruce, N.L. 1988.Aega leptonica, a new species of aegid isopod crustacean from the tropical western Atlantic,

with notes on Rocinela oculata Harger andRocinela kapala, new species. Proceedings of the Biological

Society of Washington 101(1):95-101. I

110 I
!

I Brusca, R.C. 1977. Range extensions and new host records of cymothoid isopods (Isopoda: Cymothoidae) in

the east Pacific. Bulletin of the California Academy of Sciences 76(2): 128-131.

I Brusca, R.C. 1978a. Studies on the cymothoid fish symbionts of the eastern Pacific (Isopoda: Cymothoidae). I. Biology of Nerocila californica. Crustaceana 34(2): 141-154.

i Brusca, R.C. Cymothoidae). 1978b. Studies on the cymothoid fish symbionts of the eastern Pacific (Crustacea: Isopoda: II. Biology and systematics ofLironeca vuIgaris Stimpson, 1857. Occasional Papers

of the Allan Hancock Foundation (n. ser.) 2:1-19.

I Brusca, R.C. Idoteidae. and B.R. American Wallerstein. Museum1977. Novitates The marine 2643:1-17. isopod Crustacea of the Gulf of California. I. Family

i Brusca, University R.C. 1980. Common Intertidal Invertebrates of the Gulf of California. Second, revised edition. of Arizona Press, Tucson, Arizona. 513 pp.

Brusca, R.C. 1981. A monograph on the Isopoda Cymothoidae (Crustacea) of the eastern Pacific. Zoological

I Journal of the Linnean Society 73(2): 117-199.

Brusca, R.C. 1983a. A monograph of the isopod family Aegidae in the tropical eastern Pacific. I. The genus

i Aega. Allan Hancock Monographs in Marine Biology 12:1-39. Brusca, R.C. 1983b. Two new idoteid isopods from Baja California and the Gulf of California (Mexico) and

an analysis of the evolutionary history of the genus Colidotea (Crustacea: Isopoda: Idoteidae).

I Transactions of the San Diego Society of Natural History 29(4):69-79.

Brusca, R.C. 1984. Phylogeny, evolution and biogeography of the marine isopod Subfamily Idoteinae

(Cmstacea: Isopoda: Idoteidae). Transactions of the San Diego Society of Natural History 20(7):99-

I 134.

Brusca, R.C. 1987.Biogeographic relationships of Galapagos marine isopod crustaceans. Bulletin of Marine

I Science 4(2):268-281.

Brusca, R.C. and S.C. France. 1992. The genus Rocinela (Crustacea: Isopoda: Aegidae) in the tropical

i eastern Pacific. Zoological Journal of the Linnean Society (London) 106:231-275. Brusca, R.C. and M.R. Gilligan. 1983. Tongue replacement in a marine fish (Lutjanus guttatus) by a parasitic

isopod (Cmstacea: Isopoda). Copeia 3:813-816.

I Bmsca, R.C. and E.W. Iverson. 1985. A guide to the marine isopod Crustacea of Pacific Costa Rica. Revisa

de Biologia Tropical 33 (Supplement I): 1-77.

I Brusca, R.C. and George, and M. with Ninos.a key1978. The status of Cirolana californiensis Schultz, and C. deminuta Menzies to the California species ofCirolana (Isopoda: Cirolanidae). Proceedings of

the Biological Society of Washington 91(2):379-385.

I Brusca, R.C. and B.R. Wallerstein. 1979a. The marine isopod crustaceans of the Gulf of California II.

Idoteidae: new genus and species, range extensions, and comments on evolution and taxonomy within

the family. Proceedings of the Biological Society of Washington 92(2):253-271.

I Bmsca, R.C. and B.R. Wallerstein. 1979b. Zoogeographic patterns of idoteid isopods in the northeast Pacific,

with a review of shallow water zoogeography of the area. Bulletin of the Biological Society of

I Washington 3:67-105.

Brusca, R.C., R. Wetzer and S.C. France. 1995. Cirolanidae (Cmstacea: Isopoda: Flabellifera) of the tropical

i eastern pacific. Proceedings of the San Diego Natural History Museum, 30:1-96.

i 111
!

Brusca, R.C. and J.R. Weinberg. 1987. A new isopod crustacean from Pacific Panama,Excirolana chamensis m

new species (Isopoda: Flabellifera: Cirolanidae). Contributions in Science, Natural History Museum

of Los Angeles County 392:11-17. •

Brusca, R.C. and G.D.E Wilson. 1991. A phylogenetic analysis of the Isopoda with some classifactory

recommendations. Memoirs of the Queensland Museum 31:143-204.

m

Cadien, D. and R.C. Brusca. 1993. Anthuridean isopods of California and the temperate northeast Pacific. m

SCAMIT Newsletter 12 (6): 1-26.

Caiman, W.T. 1909. Crustacea. In, R. Lankester, Ed. A Treatise on Zoology. London, Adam and Charles m

Black. 346 pp.

Camp, D.K. 1988. Bythognathia yucatanensis, new genus, new species, from abyssal depths in the Caribbean •

Sea, with a list of the gnathiid species described since 1926 (Isopoda: Gnathiidea). Journal of Crustacean |

Biology 8(4):668-678.

Dana, J.D. 1853. United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under m

the command of Charles Wilkes, U.S.N. Vol. XIII. Crustacea. Part 11:692-1618.

Dana, J.D. 1854. Catalogue and descriptions of Crustacea collected in California by Dr. John L. LeConte. m

Proceedings of the Academy of Natural Sciences of Philadelphia 7:175-177. l

Delaney, P.M. and R.C. Brusca. 1985.Two new species ofTridentella Richardson, 1905 (Isopoda: Flabellifera:

Tridentellidae) from California, with a rediagnosis and comments on the family, and a key to the m

genera of Tridentellidae and Corallanidae. Journal of Crustacean Biology 5(4):728-742.

Delaney, P.M. 1990. Tridentella williamsi, a new species of isopod crustacean from the British Virgin Islands, m

Western Atlantic (Flabellifera: Tridentellidae). Proceedings of the Biological Society of Washington |

103(3):643-648.

Fee, A.R. 1926. The tsopoda of Departure Bay and vicinity, with descriptions of new species, variations, and m

colour notes. Contributions to Canadian Biology and Fisheries (n. ser.) 3(2): 15-46. m

George, R.Y., and J.-O. Strrmberg. 1968. Some new species and new records of marine isopods from San m

Juan Archipelago, Washington, U.S.A. Crustaceana 14(3):225-254. m

Gurjanova, E. 1933. Contributions to the Isopoda-fauna of the Pacific. Issledovaniia Moriya SSSR (Exploration

of the Sea of the USSR). Leningrad 19:79- 91. m

m

Hansen, H.J. 1897. Reports on the dredging operations off the 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 m

by the U.S. Fish Commission Steamer Albatross during 1891, Lieut. Commander Z. L. Tanner, m

u.S.N., commanding. XII. The Isopoda. Bulletin of the Museum of Comparative Zoology, Harvard

m

31(5):95-129.

Harrison, K. and G.C.B. Poore. 1984. Serolis (Crustacea, Isopoda, Serolidae) from Australia, with a new m

species from Victoria. Memoirs of the Museum of Victoria 45:13-31.

Hessler, R.R. 1970. The Desmosomatidae (Isopoda, Asellota) of the Gay Head-Bermuda Transect. Bulletin m

m

of the Scripps Institution of Oceanography 15:1-185.

m

Hessler, R.R. and J.-O. Strrmberg. 1989. Behavior ofjaniroidean isopods (Asetlota), with special reference •

to deep-sea genera. Sarsia 74:145-59.

Hodgson, T.V. 1910. Crustacea. IX. Isopoda. National Antarctic Expedition, Natural History 5:1-77. I

112 m
!

I Holdich, D.M. and K. Harrison. 1980. The crustacean isopod genus Gnathia Leach from Queensland waters

with descriptions of nine new species. Australian Journal of Marine andFreshwater Research 31:215-

I 240. Hooker, A. 1985. New species of Isopoda fromthe Florida Middlegrounds (Crustacea: Peracarida). Proceedings

of the Biological Society of Washington 98:255-280.

I Iverson, E.W. 1972. A new subtidal Synidotea from Central California (Crustacea: Isopoda). Proceedings of

the Biological Society of Washington 85(47):541-548.

I Iverson, E.W. 1974. Range extensions for some California marine isopod crustaceans. Bulletin of the Southern California Academy of Sciences 73(3): 164-169.

i Jay, C.V.Crangonfrancisorum 1989. Prevalence, .sizeAmeric and fecuanndMidland ity of theNaturalist parasitic 121:68-77. isopod Argeia pugettensis on its host shrimp

Juilfs, H.B. and J.-W. W_igele. 1987. Symbiotic bacteria in the gut of the blood-sucking Antarctic fish parasite

I Gnathia calva (Crustacea: Isopoda). Marine Biology 95:493-499.

Just, J. 1990. Abyssianiridae, a synonym of Paramunnidae (Crustacea: Isopoda: Asellota), with two new

species of Abyssianira from South-Eastern Australia. Memoirs of the Museum Victoria 50(2), 403-

I 415.

Just, J., and G.C.B. Poore. 1992. Vermectiadidae, a new primitive asellote isopod family with important

I phylogenetic implications. Journal of Crustacean Biology 12(1): 125-145.

Kensley, B. 1979. New species of anthurideans from the Cook and Fiji Islands (Crustacea: Isopoda: Anthuridea).

i Proceedings of the Biological Society of Washington 92(4):814-836. Kensley, B. 1982. Revision of the southern African Anthuridea (Crustacea, Isopoda). Annals of the South

African Museum 90(3):95-200.

I Kensley, B. and M. Schotte. 1989. Guide to the Marine Isopod Crustaceans of the Caribbean. Smithsonian

Institution Press, Washington, D. C. 308 pp.

I Kozloff, E.N. 1987. Marine Invertebrates of the Pacific Northwest. University of Washington Press, Washington. 511 pp.

i Kussakin,Zoologischeskogo O.G. 1962. On the fauna of Janiridae 0sopoda, Asellota) from the seas of the USSR. Trudy

Instituta Akademiya Nauk SSSR, 30:17-65 (in Russian).

Kussakin, O.G. 1979. Marine and brackish-water lsopoda of cold and temperate (boreal) waters of the Northern

I Hemisphere. Akad. nauk SSSR Part 1.(Flabellffera, Valvifera, and Tyloidea). [in Russian]. Opredeliteli po faune SSSR, [Acad. Sci. USSR, Zool., Leningrad], No. 122, 472 pp.

i Kussakin, O.G. 1982a. Marine and brackish-water Isopoda of cold and temperate (boreal) waters of the Northern Hemisphere. Part 2. (Anthuridea, Microcerberidea, Valvifera, and Tyloidea). [in Russian].

Opredeliteli po faune SSSR, Akad. nauk SSSR [Acad. Sci. USSR, Zool., Leningrad], No. 131,461

i Ku pp. ssakin, O.G. 1982b. Supplement to the isopod crustacean fauna from the shelf zones of the Antarctic

(From the material of the Soviet Antarctic Expedition 1965-1968). In, Fauna and Distribution of

I CDal'nevostochnyi rustaceans fromNauchyi the Southern Tsentr and Antarctic Waters. [in Russian] Akademiya Nauk CCCP, (Far East Science Center), Vladivostok. pp. 73-105.

I

I 113
!

Kussakin, O.G. 1988. Marine and brackish-water Isopoda of cold and temperate (boreal) waters of the Northem m

Hemisphere. Part 3. (Asellota. 1.). [in Russian]. Opredeliteli po faune SSSR, Akad. nauk SSSR

[Acad. Sci. USSR, Zool., Leningrad], No. 152, 500 pp. m

Lee, W.L. 1966a. Pigmentation of the marine isopod Idotea montereyensis. Comparative Biochemistry and

Physiology 18:17-36.

mm

Lee, W.L. 1966b. Color change and the ecology of the marine isopod Idotea (Pentidotea) montereyensis m

Maloney, 1933. Ecology 47(6):930-941.

Lee, W.L. 1972. Chromatophores and their role in color change in the marine isopod Idotea montereyensis m

Maloney. Journal of Experimental Marine Biology and Ecology 8(3):201-215. m

Lee, W.L. and B.M. Gilchrist. 1972. Pigmentation, color change and the ecology of the marine isopod Idotea •

resecata Stimpson. Journal of Experimental Marine Biology and Ecology 10:1-27. m

Lee, W.L. and B.M. Gilchrist. 1975. Monohydroxy carotenoids inldotea (Crustacea: Isopoda). Comparative

Biochemistry and Physiology 51B:247-253. mm

Levin, L.A., D.J. DeMaster, L.D. McCann, and C.L. Thomas. 1986. Effect of giant protozoans (class:

Xenophyophorea) on deep-seamount benthos. Marine Ecology Progress Series 29:99-104. m

Lockington, W.N. 1876. Remarks on the Crustacea of the Pacific Coast, with descriptions of some new

m

species. Proceedings of the California Academy of Sciences 7:28-36.

m

Lockington, W.N. 1877. Description of seventeen new species of Crustacea. Proceedings of the California m

Academy of Sciences 7:41-48.

Markham, J.C. 1974. A new species ofPleurocrypta (Isopoda, Bopyridae), the fh-st known from the western m

Atlantic. Crustaceana 26(3):267-272. m

Markham, J.C. 1975. A review of the bopyrid isopod genus Munidion Hansen, 1897, parasitic on galatheid •

crabs in the Atlantic and Pacific Oceans. Bulletin of Marine Science 25(3):422-441. m

Markham, J.C. 1988. Descriptions and revisions of some species of Isopoda Bopyridae of the northwestern

Atlantic coast. Zoologische Verhandelingen 246:1-63. mm

Menzies, R.J. 1950a. The taxonomy, ecology and distribution of northern California isopods of the genus

Idotea with the description of a new species. The Wasmann Journal of Biology 8(2): 155-195. m

Menzies, R.J. 1950b. A remarkable new species of marine isopod, ErichsoneUa crenulata n. sp., from Newport m

Bay, California. Bulletin of the Southern California Academy of Sciences 49(1):29-35. m

Menzies, R.J. 1951a. New marine isopods, chiefly from northern California, with notes on related forms, m

Proceedings of the United States National Museum 101(3273): 105-156.

Menzies,

88.

R.J. 195lb. A new species of Limnoria (Crustacea: Isopoda) from Southern California, 50(2):86- m

m

Menzies, R.J. 1951c. A new subspecies of marine isopod from Texas. Proceedings of the United States •

National Museum 101(3289):575-579. m

Menzies, R.J. 1951d. A new genus and new species of asellote isopod, Caecijaera horvathi, from Los AngelesLong Beach Harbor. American Museum Novitates 1542:1-7. m(IN

Menzies, R.J. 1951e. Notes on the structure of the marine asellote isopod laniropsis breviremis G.O. Sars.

Meddeleser fra det Zoologiske Museum, Oslo 57:153-155. m

114 m
!

I Menzies, R.J. and J.L. Barnard. 1951. The isopodan genuslais (Crustacea). Bulletin of the Southern California

Academy of Sciences 50(3): 136-151.

I Menzies, R.J. 1952a. The occurrence of a terrestrial isopod in plankton. Ecology 33(2):303.

Menzies, R.J. 1952b. Some marine asellote isopods from Northern California, with descriptions of nine new

i species. Proceedings of the United States National Museum 102(3293):117-159. Menzies, R.J. 1954a. A review of the systematics and ecology of the genus Exosphaeroma, with the description

of a new genus, a new species, and a new subspecies (Crustacea, Isopoda, Sphaeromatidae). American

I Museum Novitates 1683:1-24.

Menzies, R.J. 1954b. The comparative biology of reproduction in the wood-boring isopod crustacean L/mnor/a.

i Menzies,Bulletin of the Museum of Comparative Zoology 112(5):364-388. R.J. 1958. The distribution of wood-boring L/mnor/a in California. Proceedings of the California

Academy of Sciences 29(7):267-272.

I Menzies, R.J. 1961. Suggestion of night-time migration by the wood-borer Limnoria. Oikos 12(1): 170-172.

Menzies, R.J. 1962. The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico. Pacific

I Naturalist 3(11):337-348.

Menzies, R.J. and J.L. Barnard. 1959. Marine Isopoda on coastal shelf bottoms of southern California:

i Menzies,systematics and ecology. Pacific Naturalist 1(11- 12):1-44. R.J. and T.E. Bowman. 1956. Emended description and assignment to the new genus RonaIea of the

idoteid isopod Erichsonella pseudoculata Boone. Proceedings of the United States National Museum

I 106(3371):339-343.

Menzies, R.J. and D. Frankenberg. 1966. Handbook on the Common Marine Isopod Crustacea of Georgia.

i Menzies,University of Georgia Press, Athens, Georgia. 93 pp. R.J. and P.W. Glynn. 1968. The common marine isopod Crustacea of Puerto Rico. Studies on the

fauna of Curacao and other Caribbean Islands, Vol. 27. The Hague, Martinus Nijhoff. 133 pp.

I Menzies, R.J. and M.A. Miller. 1972. Systematics and zoogeography of the genus Synidotea (Crustacea:

Isopoda) with and account of California species. Smithsonian Contributions to Zoology 102:1-33.

I Menzies, R.J. and J.L. Molar. 1952. The occurence of the wood-boring crustaceanl_J'mnoria and of Nebaliacea in Morro Bay, California. The Wasmann Journal of Biology 10(1):81-86.

i Menzies, R.J. and J.L. Mohr. 1962. Benthic Tanaidacea and Isopoda from the Alaskan Arctic and the Polar Basin. Crustaceana 3(2): 192-202.

Menzies, R.J., J.L. Mohr, and C.M. Wakeman. 1963. The seasonal settlement of wood-borers in Los AngelesI Long Beach Harbors. The Wasmann Journal of Biology 21(2):97-120.

Menzies, R.J. and R.J. Waidzunas. 1948. Postembryonic growth changes in the isopod Pentidotea resecata

(Stimpson) with remarks on their taxonomic significance. Biological Bulletin 95(1): 107-113.

I Menzies, R.J. and T.M. Widrig. 1955. Aggregation by the marine wood-boring isopod, Limnoria. Oikos

6(2): 149-152.

I Mezhov, B.V. 1986. Bathyal and abyssal Nannoniscidae and Desmosomatidae (Isopoda, Asellota) from Alaska

Bay. [in Russian]. Archives of the Zoological Museum, Moscow State University 24:126-167.

!

i 115
!

Miller, M.A. 1940. The isopod Crustacea of the Hawaiian Islands (Chelifera and Valivera). Occasional Papers I

of Bernice P. Bishop Museum 15(26):295-321.

Miller, M.A. 1941. The isopod Cmstacea of the Hawaiian Islands, II. Asellota. Occasional Papers of Bernice H

P. Bishop Museum 16(13):305-320. i

Miller, M.A. 1968. Isopoda and Tanaidacea from buoys in costal waters of the continental United States, •

Hawaii, and the Bahamas (Crustacea). Proceedings of the Biological Society of Washington 125:1- i

53.

Miller, M.A. 1975. Isopoda and Tanaldacea. In, R. I. Smith and J. T.Carlton, Eds. Light's Manual: Intertidal B

Invertebrates of the Central California Coast. University of California Press, Berkeley, California. B

pp. 277-312. i

Miller, M.A. and W.L. Lee. 1970. A new idoteid isopod, Idotea (Pentidotea) kirchanskii, from central California i

(Crustacea). Proceedings of the Biological Society of Washington 82(61):789-798.

Miller, M.A. and R.J. Menzies. 1952. The isopod Crustacea of the Hawaiian Islands, HI. Superfamily i

Flabellifera, Family Anthuridae. Occasional Papers of the Bernice P. Bishop Museum 21(1): 1-15.

Monod, T. 1926. Les Gnathiidae, essai monographique (Morphologie, Biologie, Syst6matique). M_moires de •

la Soci6t6 des Sciences Naturelles du Maroc 13:1-668. i

Negoescu, I. and J.-W. W_igete. 1984. World list of the anthuridean isopods. Extrait des Travaux du MusEum

d'Histoire Naturelle Grigore Antipa, 25:99-146. HRI

Nierstrasz, H.F. and G.A. Brender fiBrandis. 1923. Die Isopoden tierSiboga-Expedition. II. Isopoda GenuinaI. Epicaridea. Siboga-Expeditie 32b:57-122. E.J. Brill, Leiden. i

Nordenstam, A. 1933. Marine Isopoda of the families Serolidae, Idoteidae, Pseudidoteidae, Arcturidae,

Parasellidae and Stenetriidae mainly from the South Atlantic. Further Zoological Results of the Swedish

Antarctic Expedition, 1901-1903, 3(1): 1-284. i

Oshel, P.E. and D.H. Steele. 1988. Comparative morphology of amphipod setae, and a proposed classification

of setal types. Crustaceana (Suppl. 13, Studies on Amphipoda):90-99. •

Poore, G.C.B. 1980. A revision of the genera of the Paranthuridae (Crustacea: Isopoda: Anthuridea) with a

catalogue of species. Zoological Journal of the Linnean Society 68(1 ):53-67.

Poore, G.C.B. 1984a. Paranthura (Crustacea, Isopoda, Paranthuddae) from south-eastern Australia. Memoirs i

of the Museum Victoria 45:33-69.

Poore, G.C.B. 1984b. Redefinition of Munna and Uromunna (Crustacea: Isopoda: Munnidae), with descriptions i

of five species from coastal Victoria. Proceedings of the Royal Society of Victoria 96(2):61-81.

Poore, G.C.B. 1985. Australian chaetiliids (Crustacea: Isopoda: Valvifera): A new genus, new species and •

remarks on the family. Memoirs of the Museum Victoria 46:153-171. |

Poore, G.C.B. 1991. Book review "Brandt, Angelika. 1990. Antarctic valviferans (Crustacea, Isopoda,

Valvifera). New genera, new species and redescriptions. E.J. Brill, Leiden. Pp. 1-176, 92 figs.".

Journal of Crustacean Biology 11(3):477-478.

Poore, G.C.B. and H.M. Lew Ton. 1985. New species of Cyathura (Crustacea: Isopoda: Anthuridae) from i

estuaries of Eastern Australia. Memoirs of the Museum Victoria 46:89-101. |

Poore, G.C.B. and H.M. Lew Ton. 1986. Mesanthura (Crustacea: Isopoda: Anthuridae) from south-eastern

Australia. Memoirs of the Museum Victoria 47(1): 87-104. i

116 I
!

I Poore, G.C.B. and H.M. Lew Ton. 1988a. Antheluridae, a new family of Crustacea (Isopoda: Anthuridea)

with new species from Australia. Journal of Natural History 22:489-506.

I Poore, G.C.B. and H.M. Lew Ton. 1988b. More Australian species of Haliophasma (Crustacea: Isopoda: Anthuridae). Memoirs of the Museum of Victoria 49(1) :85-106.

Poore, G.C.B. and H.M. Lew Ton. 1988c. A generic review of the Hyssuridae (Crustacea: Isopoda) with a

new genus and new species from Australia. Memoirs of the Museum of Victoria 49(1 ):169-193.

Poore, G.C.B. and H.M. Lew Ton. 1988d. Amakusanthura and Apanthura (Crustacea: Isopoda: Anthuridea)

I with new species from tropical Austrailia. Memoirs of the Museum of Victoria 49:170-147.

Poore, G.C.B. and H.M. Lew Ton. 1990. The Holognathidae (Crustacea: Isopoda: Valvifera) expanded and

i redefined on the basis of body-plan. Invertebrate Taxonomy 4:55-80. Richardson H. 1897. Description of a new genus and species of Sphaeromatidae from Alaskan waters.

Proceedings of the Biological Society of Washington 11:181-183.

I Richardson, H. 1898. Description of four new species of Rocinela, with a synopsis of the genus. Proceedings

of the American Philosophical Society 37:8-17.

I Richardson H. 1899a. Key to the isopods of the Pacific coast of North America, with descriptions of twenty- two new species. Proceedings of the United States National Museum 21:815-869.

i Richardson H. 1899b. Key to the isopods of thePacific coast of North America, with descriptions of twenty- two new species. Annals and Magazine of Natural History 7(4):157-187; 260-277; 321-338.

Richardson H. 1900. Synopses of North-American invertebrates. VIII. The Isopoda. Part 1. Chelifera,

I Flabellifera, Valvifera. The American Scientist 34:207-230.

Richardson H. 1904a. Contributions to the natural history of the Isopoda. Parts I-IV. Proceedings of the

United States National Museum 27:1-89. Parts V-VI. Proceedings of the United States National

I Museum 27:657-681.

Richardson, H. 1904b. Isopod crustaceans of the northwest coast of North America. Harriman Alaska

I Expeditions 10:213-230.

Richardson H. 1905a. A monograph on the isopods of North America. Bulletin of the United States National

Museum 54:1-727.

I Richardson H. 1905b. Further changes in crustacean nomenclature. Proceedings of the Biological Society of

Washington 18:9-10.

I Richardson, H. 1906. Descriptions of new isopod crustaceans of the family Sphaeromatidae. Proceedings of the United States National Museum 31:1-22.

I Richardson, of aH. new1908. genusOnand some isopods of the family Dajidae from the northwest Pacific Ocean, with descriptions two new species. Proceedings of the United States National Museum 33:689-696.

Richardson H. 1909. Isopods collected in the northwest Pacific by the U. S. Bureau of Fisheries Steamer

I Albatross in 1906. Proceedings of the United States National Museum 37:75-129.

Sars, G.O. 1870. Nye Dybuandscrustaceer fra Lofoten Forhandlinger I Videnskabs-Selskabet I Kristiania

I 1869:145-286

Sars, G.O. 1879. Crustacea et Pycnogonida nova in itinere 2do et 3tio expeditionis norvegicae an_no1877 and

78 collecta. Arch. Math. Naturv. 4:427-476.

!

i I17
!

Sars, G.O. 1883. Oversigt af Norges Crustaceer med foreltibige Bemaerkninger over de nye eller mindre I

bekjendte Arter. I. (Podophthalmata-Cumacea-Isopoda-Amphipoda). Forhandlinger I VidenskabsSelskabet I Kristiania 1882(18): 1-124. •

Sars, G.O. 1897a.Isopoda. Part V,VI. Idotheidae, Arcturidae, Asellidae, Ianiridae, Munnidae. Bergen, Norway:

Bergen Museum, (An Account of the Crustacea of Norway with Short Descriptions and Figures of

All the Species; II) (Plates XXXIII-XLIX). I

Ill

Sars, G.O. 1897b. Isopoda. Part VII, VIII. Desmosomidae, Munnopsidae. Bergen, Norway: Bergen Museum,

(An Account of the Crustacea of Norway with Short Descriptions and Figures of All the Species; II) •

(Plates L-LXIII). |

Sars, G.O. 1898. Isopoda. Part IX, X. Munnopsidae (concluded), Ligiidae, Trichoniscidae, Oniscidae (part).

(An Account of the Crustacea of Norway with Short Descriptions and Figures of All the Species, II.)

Bergen Museum, Bergen, Norway. (Plates LXIV-LXXXIV).

Sars, G.O. 1899. Isopoda. (An Account of the Crustacea of Norway with short descriptions and figures of all •

the species, II.) Bergen Museum, Bergen, Norway. i-x pages. (Preface Only). |

Sars, G.O. 1899. Isopoda. Part XIII, XIV. Cryptoniscidae, Appendix. (An Account of the Crustacea of

Norway with Short Descriptions and Figures of All the Species, II.) Bergen Museum, Bergen, Norway. •

233-270 pages. (Plates XCVII-C, Suppl. P1. I-IV).

Sassaman, C., G.A. Schultz and R. Garthwaite. 1984. Host, synonymy, and parasitic incidence of Bopyrella •

calmani (Richardson) from central California (Isopoda: Epicaridea: Bopyridae). Proceedings of the |

Biological Society of Washington 97(3): 645-654.

Schultz, G.A. 1964. Some marine isopod crustaceans from off the southern California coast. Pacific Science E

28(3):307-314.

Schultz, G.A. 1966. Submarine canyons of southern California. Part IV.Systematics: Isopoda. Allan Hancock •

Pacific Expeditions 27(4): 1-56. |

Schultz, G.A. 1969. The Marine Isopod Crustaceans. Wm. C. Brown Company, Dubuque, Iowa. 359 pp.

mira

Schultz, G.A. 1972. Gnathia sanctaecrucis nom. nov., substituted for the preoccupied name Gnathia hirsuta I

Schultz, 1966, from southern California (Isopoda, Gnathiidae). Crustaceana 23(Part I): 112.

Schultz, G.A. 1973. Ancinus H. Milne Edwards in the New World (Isopoda, Flabellifera). Crustaceana II

25(3):267-275.

Schultz, G.A. 1977. Anthurids from the west coast of North America, including a new species and three new n

genera (Crustacea, Isopoda). Proceedings of the Biological Society of Washington 90(4):839-848. |

Serov, P., and G.D.E Wilson. 1995. A review of the Stenetriidae (Crustacea: Isopoda: Asellota). Records of

the Australian Museum 47:39-82. I

Sheppard, E.M. 1933. Isopod Crustacea. Part I. The family Serolidae. Discovery Report 7:253-362.

Sheppard, E.M. 1957. Isopod Crustacea. Part II. The suborder Valvifera. Families: Idoteidae, Pseudidotheidae n

and Xenarcturidae fam. n. with a supplement to isopod Crustacea, Part I. The family Serolidae.

Discovery Report 29:141-198.

Strrmberg, J.-O. !971. Contribution to the embryology of bopyrid isopods with special reference toBopyroides, !

Hemiarthrus, and Pseudione (Isopoda, Epicaridea). Sarsia 47:1-46.

I

118 I
!

I Tattersall, W.M. 1905. The marine fauna of the coast of Ireland. Part V. Isopoda. Great Britain, Reports of

the Department of Agriculture and Technical Instruction for Ireland, Scientific Investigations of the

i Fisheries Branch, 1904 11(1905): 1-90. Thistle, D. 1979. A redescription of two species of Ilyarachna (Asellota: Janiroidea) from off California

(Crustacea: Isopoda). Zoological Journal of the Linnean Society 67:381-386.

I Thistle, D. 1980. A revision ofllyarachna (Crustacea, Isopoda) in the Atlantic with four new species. Journal

of Natural History, 14:111-143.

I Thistle, D. 34:73-87. and G. Wilson. 1987. A hydrodynamically modified, abyssal isopod fauna. Deep-Sea Research

i Thun, M.A. occidentaIis and R.C.Boone, Brusca. and1its98synonymy 0. On the with statusB, of patagonica the easternSchioedte Pacific andcymot Meinert hoid (Cmsta fish pacrasite ea: Isopoda: Braga

Cymothoidae). Bulletin of the Southern California Academy of Sciences 79(3): I30-132.

I Upton, N.P.D. Paragnathiaformica 1987a. Asynchronous male and female life cycles in the sexually dimorphic, harem-forming (Cmstacea: Isopoda). Journal of Zoology, London 212:677-690.

i Upton, N.P.D. 1987b. Gregarious larval settlement within a restricted intertidal zone and sex differences in subsequent mortality in the polygynous saltmarsh isopodParagnathiaformica (Crustacea: Isopoda).

Journal of the Marine Biological Association of the United Kingdom 67(3):663-678.

I Walker, A.O. 1898. Cmstacea collected by W.A. Herdman F.R.S. in Puget Sound, Pacific Coast of North America, September 1897. Liverpool Biological Society, Proceedings and Transactions 12:268-287,

1896-1899.

I Wallerstein, B.R. and R.C. Brusca. 1982. Fish predation: a preliminary study of its role in the zoogeography

and evolution of shallow water idoteid isopods (Crustacea: Isopoda: Idoteidae). Journal of Biogeography

9:135-150.

I W/igele, J.-W. 1979. Die Homologie der Mundwerkzeuge von Cyathura carinata (KrCyer, 1847) (Crustacean,

Isopoda, Anthuridea). Zoologische Anzeige, Jena 203(5/6):334-341.

I W/igele J.-W. 1981 a. Zur Phylogenie der Anthuridea (Crustacea, Isopoda). Mit Beitrfigen zur Lebensweise,

Morphologie, Anatomic and Taxonomic. Zoologica (Stuttgart) 45(2)132:1-127.

W_igele,J.-W. 198 lb. Study of the Hyssuridae (Cmstacea: Isopoda: Anthuridea) from the Mediterranean and the Red Sea. Israel Journal of Zoology 30:47-87.

W_igele, J.-W. 1987. Description of the postembryonal stages of the Antarctic fish parasite Gnathia calva

Vanh6ffen (Crustacea: Isopoda) and synonymy with Heterognathia Amar & Roman. Polar Biology 7:77-92.

I W_igele,biogeography J.-W. 1994. Notes and aondescription Antarctic ofandnewSouth genera. American Zoologisches Serolidae, Jahrbuch with remarks der Systematik on the phylogenetic 121:3-69.

Wetzer, R., H.G. Kuck, P. Ba6z R., R.C. Brusca and L.M. Jurkevics. 1991. Catalog of the isopod Cmstacea

type collection of the Natural History Museum Los Angeles County. Report

of Technical Number

3:1-59.

Wetzer, R., fromP.M. the Gulf Delaney, of Mexico, and R.C. andBrusca. a review1987. of thePolit "'Conilera olana genus-group" wickstenae newof Bruce speies(1986). of cirolanid Contributions isopod

in Science, Natural History Museum of Los Angeles County 392:1-10.

!

i 119
!

Wilson, G. 1976. The systematics and evolution of Haplomunna and its relatives (Isopods, Haplomunnidae, i

new family). Journal of Natural History 10:569-580.

Wilson, G. 1981. Taxonomy and postmarsupial development of a dominant deep-sea eurycopid isopod i

(Crustacea). Proceedings of the Biological Society of Washington 94(1 ):276-294. I

Wilson, G. 1982a. Two new natatory asellote isopods (Crustacea) from the San Juan Archipelago, Baeonectes II

improvisus n. gen., n. sp. and Acanthamunnopsis milled n. sp., with a revised description of A. !

hystrix Schultz. Canadian Journal of Zoology 60(12):3332-3343.

Wilson, G. 1982b. Systematics of a species complex in the deep-sea genus Eurycope, with a revision of six i

previously described species (Crustacea, Isopoda, Eurycopidae). Bulletin of the Scripps Institution

of Oceanography, University of California Press. Berkeley, Los Angeles, London. 64 pages.

Wilson, G. 1983a. An unusual species complex in the genus Eurycope (Crustacea: Isopoda: Asellota) from i

the deep North Atlantic Ocean. Proceedings of the Biological Society of Washington 96(3):452-467.

Wilson, G. 1983b. Systematics of a species complex in the deep-sea genus Eurycope, with a revision of six i

previously described species (Crustacea, Isopoda, Eurycopidae). Bulletin of the Scripps Institution

of Oceanography 25:1-63.

Wilson, G. 1983c. Variation in the deep-sea isopod, Eurycope iphthima (Asellota, Eurycopidae). Depth related i

clines in rostral morphology and in population structure. Journal of Crustacean Biology 3:127-140.

Wilson, G. 1987. The road to the Janiroidea: the comparative morphology and evolution of the asellote 1

isopods. Zeitschrift fiir zoologische Systematik und Evolutionsforschung 25:257-280.

Wilson, G. 1989. A systematic revision of the deep-sea subfamily Lipomerinae of the isopod crustacean •

family Munnopsidae. Bulletin of the Scripps Institution of Oceanography 27:1-138 I

Wilson, G. I991. Functional morphology and evolution of isopod genitalia. In, R. Bauer and J. Martin (Eds.).

Crustacean Sexual Biology. University of Columbia Press. pp. 228-245. i

I

Wilson, G.D.E 1994. A phylogenetic analysis of the isopod family Janiridae (Asellota). Invertebrate Taxonomy

8:749-766.

Wilson, G.D.E and J.-W. W_igele. 1994. A systematic review of the family Janiridae (Isopoda, Asellota). !

Invertebrate Taxonomy 8:683-747.Wilson, G. and R.R. Hessler. 1981. A revision of the genusEurycope

(Isopoda, Asellota) with descriptions of three new genera. Journal of Crustacean Biology 1(3):401- •

423.

W'dson, G., O. Kussakin and G. Vasina. 1989. A revision of the genusMicroprotus Richardson with descriptions i

of two new species,M, acutispinatces andM. lobispinatus (Asellota, Isopoda, Crustacea). Proceedings i

of the Biological Society of Washington 102(2):339-361.

Wilson, G. and D. Thistle. 1985.Amuletta, a new genus forllyarachna abyssorum Richardson 1911 (Isopoda, i

Asellota, Eurycopidae). Journal of Crustacean Biology 5(2):350-360. m

Wolff, T. 1962. The systematics and biology of bathyal and abyssal Isopoda AseUota. Galathea Report 6: 1- am

320. I

!

!

120 I
!

i 2. CUMACEA

I by

Les Watling 1and Linda D. Mceann 2

l Introduction

I Cumaceans are small crustaceans, generally ranging in size from 1 mm to I cm; however, a few

species, such as Diastylis goodsiri, may be 3 cm or more in length. The Order Cumacea currently contains

i more than 1200 species worldwide. This is a sizable increase over the 770 species listed by Jones (1969). Of these, and including those described herein, 49 species are known from the Pacific coast of the United

States. Many of the habitats where cumaceans are likely to be found, such as estuaries, shallow embayments,

I beaches, contrast, the tidalfarfiats, less and diverse the cumacean inner continental fauna ofshelf, the northeastern have not yetUnited had their Statescumacean is completely fauna known documented. (WatlingIn

1979).

I Cumaceans are malacostracans, considered by many to be an order in the Superorder Peracarida, but considered by Wafting (1983) and Schram (1986) to be a member of the Superorder Brachycarida. They

are distinguished by the following combination of features: the carapace covers the first three or four, or

I rarely six, thoracic somites; the anterior margin of the carapace is extended in front of the head as pseudorostral lobes; the telson may be present, reduced, or incorporated into the last abdominal somite (pleonite); the eyes

are confluent dorsally in all but a very few genera; the second antennae lack an exopod; and pleopods are

I absent in females (with the exception of one deep-sea species) and often reduced in number or absent in males (Fig 2.1).

I Cumacean Morphology

I The cumacean body is externally divided into carapace, thorax, and abdominal regions (Fig. 2.2).

The carapace covers three, four, or rarely six, thoracic somites. Externally the body can then be seen to be

divided into three regions, the carapace, the pereon (reduced thorax), and the abdomen or pleon. The pereon

I usually consists of five body somites, butfewer may be visible depending on the extent of the carapace. The

abdomen always contains six body somites, of which pleonite 5 is usually the longest. An articulated telson

may or may not be present terminally.

I The carapace is expanded ventrally and laterally to form a branchial chamber. Each side of the

carapace is produced anteriorly in the form of pseudorostral lobes which meet, but are not fused, in front of

I the are head, the tipsforming of thea

bpseranchial udorostepruipod m (Fig. s of2.2). the fiReach rst maxi ing lliped, to the endwhich of, ot

r

oprgethe ojectringformbeytheond,branthechpi

sealud

siophroostn,rumor

exhalant canal, for the respiratory current. The pseudorostrum may be directed anteriorly at various angles,

i o

isrincised be comptoletely form reaflnotch exed for suchthethafirst t theantenna. branchialBelow opening this inotc s dohrsal. the The carapace lowerbears marginan ofanteroventral the pseudorostrum corner,

which may be broadly rounded or produced into one or more teeth. Laterally and dorsally, the carapace may

!

_Depa_nent of Oceanography and Darling Marine Center, University of Maine, Waltmle, Maine 04573.

I 2Smithsonian Environmental Research Center, P.O. Box 28, Edgewater, Maryland 21037.

i 121
i

!

,_ I

i

o I

I

I

I

i

i

I

I

I

i

Figure 2.1. Representative cumaceans from various families likely to be encountered in the Santa Mafia Basin

Region: A, Lampropidae, mature male with elongate antenna 2; B, Lampropidae, female with eggs !

in brood pouch; C, Lampropidae, mature male with grasping antenna 2; D, Diastylidae, subadult

female; E, Diastylidae, subadult male; F, Nannastacidae, subadult female; G, Nannastacidae, mature

male; H, Leuconidae, subadult female, i

122 I
I

I

t

I fl c T,4 T8 PL1 PL=6 I __

........:

I m "--.................. pa

af Im ex _,._CoO

i Mxl x

I Figure 2.2, Side view of the cumacean body with details of the seven anterior headand thoracic appendages.

A1, antenna 1;A2, antenna 2; Md, mandible; Mxl, maxilla I (maxiUule);Mx2, maxilla 2 (maxilla);

Mxpl, maxilliped 1; Mxp2, maxilliped 2; P1, pereopod 1; P5, pereopod 5; PL1, pleonite 1;PL6,

I pleonite flagellum;6;an, T4,antenna thoracic]notch; somiteb,4basis; (=perebr, onitebranchial 1);T8,apparatus; thoracic soc,mite carapace; 8 (=pereco,onite coxa;5);ex,af,exopod; accessoryfl,

frontal lobe; ip, incisor process; lm, lacinia mobilis; mr, main flagellum; rap, molar process; o,

I oostegite; pa, palp;pl, pseudorostral lobe; s, siphon; t, telson; u, uropod (from Walling, 1979).

I be ornamented with tubercles, spines, ridges, or combination of these. The side of the be

any carapace may

depressed to form a sulcus (Fig. 2.1F, G). The ventral margin may be smooth or produced into a series of

teeth or serrations. In males, the ventral margin is extended some additional distance ventrally to house the

I enlarged peduncle of antenna 2 (Fig. 2.1E).

Behind the pseudorostrum is the anterior-most part of the head, known as the frontal lobe (Fig. 2.2).

The anterior part of the frontal lobe contains the eyelobe, which may be present even when eyes are absent.

I some cases, especially deep-sea species, eyelobe may range shape extremely elongate

In in the in from to

being completely absent. The frontal lobe ranges in shape from triangular to broadly rounded. It houses the

brain and optic ganglia (when eyes are present) and provides the point of attachment for the first and second

I antennae. As in all crustaceans the head bears five pairs of appendages, viz., the first and second pairs of

antennae, mandibles, and first and second pairs of maxillae.

i The first antenna (Fig. 2.2, 2.3A,B) consists of a three-articulate peduncle, the distal-most of which bears two rami, a main flagellum of 2 to 6 articles, and an accessory flagellum (when present) of 1 to 4

articles. In several families, the accessory flagellum consists of a single article ranging in size from minute

to nearly as long as the main flagellum. Occasionally, both flagellae are very small and their exact structure

I is difficult to determine using light microscopy. The main flagellum is often festooned with long sensory

setae, especially in males of the genus Leptostylis (Fig. 2.3D).

I

i 123
!

|

Figure 2.3. Detailed views of selected cumacean head and anterior thoracic appendages. A, antenna 1 from

female Leuconidae; B, antennae 1 and 2 from mature male Diastylidae (note enlarged peduncle

article 5 and large setal brush on antenna 2); C, antenna 2 modified for grasping, from male of i

Lampropidae; D, antennae 1 and 2 from mature male of Leptostytis (note enlarged antenna 1 and •

setal brush on peduncle article 3); E, antenna 2 of female Diastylidae; F, mandible typical of most

u

families; G, mandible from Campylaspis; H, a typical maxilla 1; I, a typical maxilla 2; J, maxilliped

1 with details of gill Iobules fromEudorelfa; K, endopod of maxilliped 1;L, maxilliped 2 of Diastytis; l

M, maxilliped 3 of Diastylis; N, maxilliped 3 of Campylaspis.

124 i
!

I The second antenna (Fig. 2.2) is generally rudimentary in the female (Fig. 2.3E) but may consist of

as many as five articles. In the male, the structure of the second antenna changes with age and its final form

i peduncle is determined bearingby thea very use tolongwhich multi-articulate it is going toflagellum, be put. In the peduncle most typical articlesformfouritandconsists five bear of a five-articulate a strong brush

of sensory setae, and peduncle article five is much longer than article four (Fig. 2.3B). In the immature

i male, the setal brush and elongate flagellum are not present; instead the second antenna is shaped like an elongate club. There are various modifications to this pattern, most of which appear to be related to the

antenna functioning as a grasping device (Fig. 2.3C). In this case, the antenna is shortened, the setal brush is

i lost or reduced, and the smaller number of flagellar articles bear specialized setae on their anterior margin. Little is known about the sensory functions of the setae arming the second antenna, so the exact uses of this

appendage cannot be predicted from its structure.

i paired first Theandmouth secondfieldmaxillae. is def'medIn bycontrast the upper to other lip (labrum), small crustaceans, paired mandibles, such as thelower amphipods, lip (labium), the upper and

and lower lip are of no taxonomic significance in cumaceans.

I mandibleThebody, cumacean a columnar mandible molarisprocess, a simpleanstruincisor cture process, (Fig. 2.2).andIna irowts basic of setae formwith it codistal nsists laofcinia an elmobilis ongate

between the molar and incisor (Fig. 2. 3F). A palp is never present. The paired mandibles are connected by

I aThose transverse mandibles mandi having bular thetendon elongate to which body arethereferred mandibular to as "boat-shaped." adductor and abductor In some families musclesthearemandibular anchored.

body is reduced, with that portion dorsal to the molar being lost. In this case, the mandible is referred to as

I truncate. ingested. Cumacean The Nannastacidae mandiblesgenus, usuallyCampylaspis, have stronglyandcolumnar allies havemolars, a recurved, used for stylet-like crushingmolar the material thought toto bebe

an aid to predation on foraminff"ers (Fig. 2.3G). Very few examinations of food habits have been made for

I cumaceans; yet be made.consequently strong relationships between the structure of the molar and food sources cannot

The fLrstmaxilla (maxillule) consists of a three-articulate protopod, the first and third articles of

I which article bear whichterminally bears eithersetose one orlobes two(endites) setae. The(Fig. palp2.2, extends 2.3H).intoThere the ante is aribackwardly-directed or part of the branchialpalpchamber, of one

but its exact role is unknown.

I The second maxilla (maxillula) consists of a three-articulate protopod, of which only the third is produced into an endite (Fig. 2.2, 2.31). On the outer side of the appendage is a thin, flattened plate, considered

to be the reduced exopod.

I In the generally covers the three most anterior thoracic somites. Only the first

cumaceans, carapace

of these is thought to be fused with (or incorporated into) the head. However, the appendages of all three

somites are modified and are termed maxillipeds. As with most crustacean thoracic legs, in cumaceans the

i basic form consists of a coxa immediately adjacent to the body, the basis, and five articles making the

up

endopod. The coxa may bear an epipod of varying size and the basis an exopod of no more than two articles

(in those forms where the exopod bears long swimming setae, the second article is multi-annulate, not

I multi-articulate, as can be seen by examining the musculature). The basis is often elongated, with the exopod attached proximally and the endopod distally. The endopod is the most developed part of the limb and its

articles are named ischium, merus, carpus, propodus, and dactylus, in consecutive order distally.

I The first which in the first thoracic is modified maxilliped, is, fact, appendage, very highiy (Fig.

2.2, 2.3J,K). The epipod is elaborated into a large branchial structure. Its posterior portion is composed of

branchial lobules (gills) on a supporting lappet and its anterior part is extended to form, with its pair, the

I branchial siphon. The endopod normally comprises five articles and extends forward to aid inthe manipulation of food near the mouth.

!

i 125
I

I

I

' D I

I

!

I

I

I

!

Figure 2.4. Detailed views of pereopods, umpods, and telons. A, pereopod 1 from Diastylidae; B, pereopod 2

from Diastylidae; C, pereopod 3 from Diastylidae; D, uropod and telson from Lampropidae; E,

uropod and telson from Diastylidae; F, ttropod from Nannastacidae (telson fused to pleonite 6). I

|

The endopod of the second maxilliped is more elongate than in the previous appendage (Fig. 2.2, g

Fig. 2.3L). The epipod and exopod are both absent. In mature females a small pair of posteriorly-directed

oostegites may be present. II

The third maxilliped is even more leg-like than the second maxilliped (Fig. 2.3M,N). It consists of E

an elongate basis and five-articulate endopod, and often possesses an exopod. The shape of this appendage

ranges from very elongate to broad and operculate. Whether truly opercular or not, the third maxilliped I

usually covers the ventral aspect of the mouth field. II

!

126 I
!

I Pereopod 1 is the first true ambulatory appendage, but is, in fact the fourth thoracic appendage (Fig.

2.4A). Its structure is much the same as for the third maxilliped, but the endopod is generally more elongate.

I An 2.4B,exoC); podexopods is usuallymaypresen or mayt. The not beremaining present onpereopods pereopodsdecrease 2 in length and robustness posteriorly (Fig. to 4. An exopod is never present on pereopod 5.

The appendages of abdominal somites 1 to 5 are known as pleopods, and are present (with a single

I exception pleopods, orin atheydeep-sea may bespecies) absent only altogether. in males. Depending on the family, there may be one to five pairs of

The last pair of abdominal appendages are the uropods. They consist of a uniarticulate peduncle

I bearing two rami, the endopod and exopod (Fig. 2.4D-F). The exopod is always two-articulate, but the endopod may consist of one to three articles.

The cumacean body terminates with a telson, on the ventral side of which is located the anus and

I anal valves. In the Bodotriidae, Leuconidae, and Nannastacidae, the telson is very short and is fused to the sixth pleonite (Fig. 2.4F). Freely articulated telsons of varying length can be found in the other families

(Fig. 2.4D,E).

!

Classification

i Family Bodotriidae Family Lampropidae*

I Subfamily Bodotriinae* Family Pseudocumatidae Vaunthompsoniinae Family Diastylidae*

Subfamily Mancocuminae Family Gynodiastylidae

I Family Leuconidae* Family Ceratocumatidae

Family Nannastacidae*

I The Order Cumacea is divided into 8 families. Those families and subfamilies dealt with in this

atlas are marked with an asterisk.

I MMS Attachment I is a list of names that have changed between the original identifications as part of the studies, and the present treatment.

!

!

!

!

!

!

i I27
!

Key to the Families of Cumacea N

with Species in the Santa Maria Basin Region

i

1A. With freely articulated telson .......................................................................................................... 2 i

lB. Without freely articulated telson ..................................................................................................... 3 1

m

2A. Telson with 0 or 2 terminal setae .................................................................................... Diastylidae

2B. Telson with 3 or more terminal setae ........................................................................... Lampropidae i

3A. Uropod endopod uniarticulate .................................................................................... Nannastacidae ll

3B. Uropod endopod 2-articulate ........................................................................................................... 4

4A. Male with 0 or 2 pairs of pleopods, females with exopods on pereopods 1 to 3 ............ Leuconidae N

4B. Mate with 5 pairs of pleopods; females with exopods only on pereopod 1 or on pereopods 1-4 .....

........................................................................................................................................ Bodotriidae 1

Family Diastylidae U

Diagnosis (as emended by Day, 1980). Mandible normally boat-shaped but rarely truncate. First

maxilliped branchial filament with numerous leaflets. Exopods present on maxilliped 3 (except in n

Paradiastylis), and in females on pereopods 1 and 2, absent or rudimentary on pereopods 3 and 4, in males U

present on pereopods 1-4. Telson variable, usually large, often with long post-anal part, sometimes short

and poorly armed, usually with 1 pair of terminal setae, occasionally with none. Uropods usually long and

slender, endopod with 1-3 articles. Male antenna 2 with long flagellum, reaching atleast to posterior margin !1

of thorax. Males with 2 pairs of pleopods; no outer process on inner ramus.

Remarks. The Diastylidae are a family of predominantly cold-water favoring cumaceans. On a l

global basis, by far the greatest number of species are found at latitudes exceeding 30°N and S (Day, 1980). II

From Point Conception northwards to British Columbia (the Oregonian Province), 12 species are known to

date. Most of these were described, or redescribed, by Caiman (1912) and Zimmer (1926, 1936, 1943), with i

single additional species added by Hart (1930) and Lie (1971). In contrast, more than 40 species of diastylids il

are known from the North Atlantic, suggesting that the fauna from the boreal eastern Pacific is vastly underrepresented. The present study has produced an additional 12new species with sufficient material to provide 1

adequate diagnoses, and another 7 probable new species represented by single specimens. l

N

N

l

I

128 I
!

I Key to the Diastylidae

from the Santa Maria Basin Region

!

1A. Telson elongate, with 1 pair of lateral setae; male antenna 1 with club-shaped article 3 bearing

many sensory setae ..................................................................................... Leptostylis .................. 2

I lB. Telson short, with 2 or more pairs of lateral setae; male antenna 1 elongate, with sensory setae on

distal articles ............................................................................................... Diastylis ..................... 3

I 2A. Uropod exopod and endopod subequal in length ................................................... LeptostyIis calva

2B. Uropod exopod reaching only halfway along endopod terminal article .............. Leptostylis abditis I

3A. Carapace with spines, but no ridges ................................................................................................ 4

I 3B. Carapace with ridges, but no spines ................................................................................................ 5

i 4A. Carapace with 4 horizontal rows of spines; pereonite 1 not completely visible in lateral view .......

................................................................................................................................ Diastylis sentosa

4B. Carapace with 1row of horizontal spines; pereonite 1 completely visible in lateral view ..............

I ................................................................................... Diastylis paraspinulosa Zimmer (not in text)

I 5A. 5B. Carapace Carapace with ridgesserrate smooth, or crenellate not serrateridges ................................................................................................ ......................................................... Diastylis crenellata6

I Carapace oblique ridges ................................................................ Diastylis peIlucida

6A. with 2 lateral

6B. Carapace with more than 2 lateral ridges ........................................................................................ 7

I 7A. Carapace with 3 lateral ridges, the area in front of the dorsal-most ridge slightly depressed; telson

not reaching end of uropod peduncle ............................................................... Diastylis californica

I 7B. Carapace with 4 distinct lateral ridges and no horizontal ridges; telson reaching end of uropod peduncle ........................................................................................................ Diastylis quadripIicata

i 7C. Carapace with 4 oblique ridges of different lengths and a few horizontal ridges; telson not reaching to end of uropod peduncle .......................................................................... Diastylis santamariensis

I Genus Diastylis Say, 1818

Type Species. Cuma rathkei (KrCyer, 1841).

I Diagnosis (after Sars, 1900 and Day, 1980). Carapace large, tumid, often armed with spiniform projections, sometimes also with lateral folds or ridges, but never with dorsal crest. Male antenna I article

3 not expanded distally, with sensory setae distally. Male antenna 2 flagellum extends to end of or beyond

i pleon. Third and fourth pereonites not coalesced. Basis of pereopods not widened in male and pereopods 2 and 3 not widely separated in ovigerous female. Pereopods 3 and 4 with rudimentary exopods or exopods

absent in female. Male with 2 pairs of pleopods. Post-anal part of telson longer than pre-anal part. Uropod

I endopod 2- or 3-articulate.

i 129
!

Remarks. This is the oldest of all cumacean genera. It is represented in all cold and cold temperate I

seas of the world. Including those described below, 8 species are known from the Oregonian Province. A

complete list can be found in Attachment II. I

Diastylis sentosa Watling and McCann, new species

Figure 2.5 I

Material Examined. Santa Maria Basin, Phase I stations 66, 201 m and 70, 200 m, 10 Nov 1983, I

and the following Phase II stations: Cruise 1-1, Nov 1986, Stas. R-2 (2), R-5 (4); Cruise 1-2, Jan 1987, Stas. I

R-2 (5), R-5 (31), PJ-1 (2); Cruise 1-3, May 1987, Stas. R-2 (7), R-5 (17); Cruise 2-3, Oct 1987, Stas. R-2

(2), R-5 (2), R-6 (1); Cruise 2-4, Jan 1988, Stas. R-2 (1), R-5 (33), R-6 (1), PJ-1 (3); Cruise 2-5, May 1988, •

Stas. R-2 (1), R-5 (15), PJ-1 (1); Cruise 3-1, Sep 1988, Stas. R-5 (2), PJ-1 (1); Cruise 3-4, May 1989, Stas. II

R-2 (2), R-4 (1), R-5 (23), R-6 (I7), PJ-I (2).

Description. Adult female and immature male, 6.5 mm. Carapace 2 times length of thoracic somites, _l

posterior two-thirds broadly rounded, lobes separated dorsally by mid-dorsal furrow; ornamentation of II

thorn-like spines arranged in 4 rows running approximately horizontally and a fifth row paralleling posterior

carapace margin; amongst large spines many short spines scattered over carapace; dorsolateral spine row •

continues anteriorly onto dorsolateral margin of pseudorostral lobes; eyelobe broad, bearing 2 spines; ventral il

margin serrate throughout length. Pereonite 1 not visible ventrally; with group of spines dorsolaterally.

Pereonites 2-5 with pair of large dorsolateral spines; spines may also be present on lateral and ventral •

margins. Pleonites with lateral spines; pleonite 5 with lateral spines terminally and additional pair midway |

along segment length. Pereopod 1 elongate, extending well beyond pseudorostral lobes. Telson extending

slightly beyond uropod peduncles; post-anal part about equal in length to pre-anal part; lateral margins with •

3-6 setae. Umpod peduncle with few setae on medial margin; rami subequal in length; endopod of 2 subequal !1

articles, with strong setae equally spaced along medial margin, terminal seta partially fused to distal article;

exopod with 3 terminal setae, lateral margin with 3-5 evenly spaced setae. Adult male, 7.0 mm. Similar to []

female but carapace spines less pronounced, anterolateral corner below eyelobe with twin anteriorly directed 1

spines. Pereonite 5 posteriorly produced, with 1 or 2 spines.

Remarks. This species belongs to a small group of northern and Arctic species characterized by the ll

presence of a spiny carapace: D. paraspinulosa Zimmer, D. spinulosa Heller, D. echinata Bate, and D.

ornata Lomakina. Of these, only D. paraspinuIosa, originally described from the Bering Sea, has been

recorded from Californian waters (Zimmer, 1943). Diastylis sentosa differs from D. paraspinulosa most •

noticeably in the following features: pereonite 1 is not completely visible in lateral view, there are at least II

four horizontal and one vertical row of spines on the carapace, and the telson bears only 4-5 pairs of lateral

setae. It is quite likely that D. paraspinulosa has been mis-identified from southern California waters or has I1

been only occasionally carried into these waters by the California Current. None of the specimens collected II

during this study are D. paraspinulosa.

Type Locality and Type Specimens. California, Santa Maria Basin, off Purisima Point, Sta. R-5 I

34°42.69'N, 120°50.83'W, 154 m, holotype USNM 273595, paratypes USNM 273596 and LACM 95-84.1.

Etymology. from L. sentus, = thorny, referring to the shape and arrangement of the spines on the

carapace. I

Distribution. Known from the Santa Maria Basin region to off Pt.Loma, San Diego; depth range,

92-410 m. i

!

130 I
!

i

!

!

!

!

/- /, .

..>\ _ f_

D

/ .,..._-

I / \ ",,

;./ ._ ,,I

.,..'_L ,,' '._,,"..,..

I / ._i _' , '. /' /" ,)..

/ ._ i__ " "X,

,,,,,"

_..Y. _:._,

I . ..j_ ," •

_,",,/,'._ Y:i

| "

Figure 2.5. D/_ty_ semosa W_,tl_g and MeCarm, new species.Female: A, carapace and pereo_ites, lateral

I vCiew; = 1ram, B, body, D=0.5lateral mm.view;C, carapaceand pereonites,dorsal view;D, telsonand uropods.Scale,Ai DiastyUs crenellata Watling and McCann, new species

I Figure 2.6

Material Examined. SantaMaria Basin, Phase II, Cruise 1-1, Nov 1986, Stas. R-1 (3), R-4 (43), RI 5 (21), PJ-1 (7); Cruise 1-2, Jan 1987, St,as.R-4 (7), R-5 (39), PJ-1 (2); Cruise 1-3, May 1987, Stas. R-2 (1), R-4 (22), R-5 (29), R-8 (1), PJ-1 (21); Cruise 2-3, Oct 1987, Stas. R-1 (2), R-2 (3), R-4 (13), R-5 (36), R-9

(1), PJ-1 (10); Cruise 2-4 Jan 1988, Stas. R-I (4), R-4 (11), R-5 (47), R-8 (3), PJ-1 (26); Cruise 2-5, May

I 1988, Stas. R-1 (3), R-2 (4), R-4 (44), R-5 (14), R-8 (4), R-9 (1), PJ-1 (14); Cruise 3-1, Oct 1988, Stas. R-1 (5), R-4 (61), R-5 (48), PJ-1 (21); Cruise 3-4, May I989, Stas. R-1 (13), R-4 (51), R-5 (18), R-8 (29), PJ-1 (24).

Description. Adult female and immature male, 2.5 mm. Carapace twice length of pereonites 1-5;

I posterior margin extended strongly laterally to cover pereonites 1 and 2 ventrally; with three major serrate

ridges, anteriormost originates middorsally behind frontal lobe, crosses onto pseudorostrum, eventually

joining anterior horizontal extension of posterior ridge, latter originates dorsolaterally near anterior ridge,

I extends posteriorly, then descends laterally in wide sweeping arc toward anterior, ending near antennal

angle, third ridge runs ventrally from anterior extension of posterior ridge to ventral margin; region of

carapace behind posterior ridge covered with dirt and debris, creating roughened look; eyelobe and frontal

I lobe each with pair of small spines; ventral margin finely toothed. Pereonites 1-5 without spines; pereonite

i 131

!

I 5 with subacute posterior corners. Maxilliped 3 basis distal margin exceptionally broad, forming opercularlike cover to mouth field; distal margin with few long, very finely plumose setae; medial margin with thick

i row of short plumose setae; endopod articles small, together less than half length of basis. Pereopod 1 long and slender; basis slender, heavily armed with finely plumose setae; articles 6 and 7 subequal in length,

together equal in length to rest of limb, with covering of simple setae throughout. Telson short, about 1.5

i times Uropodlength of pleonite 6; post-anal part slightly shorter than pre-anal part; lateral margins with 3-5 setae. peduncles more than twice length of telson; inner margins lined with closely spaced setae; exopod

longer than endopod, outer margin with many short setae, armed terminally with 4 setae; endopod of 3

articles, subequal in length, with continuation of setal part from peduncle on inner margin, distal article

I armed terminally with single strong seta.

Remarks. This species is very close to D. californica Zimmer, but differs from it in the following

I fefronaturtaels:lobwhi e, lthe e theridgriesdgearpeatte notrnserisradte, eceptive and thlye poste similar, rior Dmargin . caIifornica of the carapac does noet have coversa per ridge eonitcroesssing 1 andthe2

laterally whereas in D. californica it extends over only partof pereonite 1.The juveniles of this species have

i only a single pair of lateral telson setae, located subterminally, and have been identified in early benthic surveys from this region as Leptostylis sp. E. A second pair of lateral setae does not appear on the telson

until at least the third juvenile stage.

I 34°42.69'N, Type12Lo0°cal 50.83'W, ity and154Typem, hSopec lotype imens. USNMCalifo273591, rnia, Santa paratypes MariaUSNM Basin, 2735 off Puri 92 sanimad LACM Point, 95-82.1. Sta. R-5

Etymology. From Latin, crenellus, = embrasure, battlement, referring to the notches found at the

I top of the battlement, the pattern of which is seen on the carapace ridges.

Distribution. Known from the Santa Maria Basin region to off Pt. Loma, San Diego, depth range,

47-160 m, one record at 410 m.

!

Diastylis quadriplicata Watling and McCann, new species

I Figure 2.7

Material Examined. Off Point Arguello Phase I station 071,306 m; off Point Conception, 075,

I 293 m.

Description. Adult female and immature male, 4.5 mm Carapace about half again as long as

i pereonites 1-5; pseudorostrum broadly triangular; major ornamentation consists of 4 folds or ridges, three of which are parallel and ran dorsoventrally from vicinity of paired dorsolateral ridges; fourth dorsoventral

ridge originates at junction of pseudorostral lobe with carapace; eyelobe small, without spines. Pereonite 1

i nearly completely covered by posterior carapace margin. Pereonites 1-5 dorsal surfaces free of spines; posterior corner of pereonite 5 broadly triangular. Pereopod 1 articles 5-7 subequal in length. Telson very

slightly shorter than uropod peduncles; post-anal part slightly longer than pre-anal part; lateral margins with

i 5-8 setae. Uropod peduncle about 20% longer than rami; endopod slightly longer than exopod, 3-articulate, basal article longer than distal, armed terminally with a single seta; exopod with two setae on medial margin,

outer margin with several setae, armed terminally with 3 setae. Mature male unknown.

I waters Remarks. This species belongs to a group of diastylids found predominantly in the northern cold of the Atlantic and Pacific, with some connection through the low Arctic. These species all have a

series of parallel dorsoventral ridges on the carapace. The northern Atlantic species, D. scorpioides (Lepechin),

i D. edwardsi (Krcyer) and D. lepechini Zimmer, all have four ridges running the full dorsoventral distance and have serrate ridges on the frontal lobe and pseudorostrum. Four multi-ridged species are thus far known

from the colder Pacific waters: D. bidentata Caiman, D. dalli Calman,D. alaskensis Caiman, and D. pellucida

i Hart. Diastylis alaskensis has six ridges that run nearly vertically dorsoventral rather than obliquely as in

I 133
I

I

I

I

I

I

,"_ ,],. i/

......... , f_.._(/ D Ill

,_ / / i I

., , : _, II

/ ," /, t, II

! / _, _ l

.. ,_ ,_

' !

...

., . .,_.

Figure 2.7. Diastylis quadriplicata Watling and McCann, new species. Subadult male: A,carapace and 1

pereonites,lateralview; B,body,lateralview; C, carapaceandpereonites,dorsalview; D,telson and |

uropod. Scale, A-C = 1 mm, D = 0.5 ram.

!

this species. Also, in D. alaskensis the telson is much shorter than the Uml_d peduncles. In D. bidentata and m

D. dalli, the carapace folds are oblique and the telson is more nearly equal in length to the uropod peduncle; i

however, in these two species the basal article of the uropod endopod is longer than the two distal articles

mi

combined, which is clearly not the case in D. quadriplicata.

In

Type Locality and Type Specimens. California, Santa Barbara Co., off Pt. Arguello, Sta. 71, I

34°29.04'N, 120°44.01'W, 306 m, holotype USNM 273593, paratypes USNM 273594 and LACM 95-83.1.

Etymology. From Latin, quadros = four, and plicatus = fold or ridge, referring to the four ridges

ornamenting the carapace. il

Distribution. Known only from the Santa Maria Basin Region, approximately 290-310 m.

!

I

I

134 |
!

I Diastylis santamariensis Watling and McCann, new species

Figure 2.8

I Material Examined. Santa Maria Basin, Phase H, Cruise 1-1, Nov 1986, Sta. R-5 (4), PJ-1 (1);

Cruise 1-2, Jan 1987, Sta. R-5 (3); Cruise 1-3, May 1987, Stas. R-4 (3), R-5 (3), R-8 (22); Cruise 2-3, Oct

i 1 1987, Stas. R-2 (2), R-5 (5), PJ-1 (3); Cruise 2-4, Jan 1988, Stas. R-1 (3), R-5 (5), PJ-1 (1); Cruise 2-5, May

u _ 1988, Stas. R-5 (1), R-8 (4); Cruise 3-1, Oct 1988, Stas. R-1 (2), R-4 (3), R-5 (2), PJ-1 (3); Cruise 3-4, May

1989, Stas. R-1 (39), R-4 (2), R-5 (16), R-8 (1).

I Description. Adult female and subadult male. Carapace twice length of pereonites 1-5; surface with general overall wrinkled look; with 4 dorsoventral ridges, the two posteriormost originating dorsally at

dorsolateral ridge and running to ventral margin, anteriormost extending across frontal lobe but not reaching

n ventral margin, behind that a short ridge extending from psudorostral cut toward but not reaching ventral I margin; eyelobe smooth and globular, lenses not visible in SEM view. Pereonite 1 concealed ventrally in

i subadult male but not in young female. Pereonite 5 posterior corner bluntly rounded. Telson pre-anal part

i shorter than pleonite 6, post-anal part longer than pre-anal, with 4-5 setae on lateral margins. Uropod peduncle slightly longer than telson; endopod longer than exopod, 3 articles of endopod subequal in length, distal

article with single terminal seta; exopod with several setae on outer margin, armed with 3 setae terminally.

I Remarks. On the basis of its ornamentation, this species is closely allied to D. alaskensis

carapace

Caiman and D. paralaskensis Vassilenko and Tzareva. Diastylis santamariensis has fewer dorsoventral

ridges than either of the other two. In addition, its telson is more nearly equal in length to the uropod

[ i peduncles (versus much shorter in both D. alaskensis and D. paralaskensis) and the basal uropod endopod

roll article is not much longer than either of the distal two articles (versus much longer in D. alaskensis).

i Type Locality and Type Specimens. California, Santa Maria Basin, off Point Sal, Sta.PJ-3 34°56.26'N, 120°49.58'W, 138 m, holotype USNM 273589, paratypes USNM 273590 and LACM 95-81.1.

Etymology. Name derived from sample locations in the Santa Maria Basin region.

I Washington, Distribuas tion. Known from the Santa Maria Basin region, depth range, 90-161 m., and Puget Sound, shallow as 10 m.

i Diastylis cah'fornica Zimmer, 1936

Figure 2.9

I Material Examined. Off Point Conception, Phase I, Sta. 079, 98 m.S.

Description. Adult size, female 9 mm, male 12 mm. Carapace with elevated ridges or keels, lateral

n risidedgesofterminating frontal lobe;infrompaireddorsal elevated tooth teeth, ridge 1runs pairposteriorly dorsolaterallythenabventrally ove and meeting behind frventrally-directed ontal lobe and 1 pair ridgeto

originating at lateral tooth. Antenna 1 very slender, peduncle article 2 reaching end of pseudorostrum,

i pedunc laterally,le arti andcleforming 3 equalopercular-like in length to first covering 2 articles. of mouth Maxilliped field.3Uropod basis bropeduncle adly expanmore ded,than upturned twicesolength mewhatof

telson; endopod 3-articulate, all articles subequal in length; exopod longer than endopod.

i is most Rclosely emarks.related ThisisspecDiastylis ies is verycrenellata distinct wsp.ithn.,respect described to carapace above. sculpturing. While the latter The species is a species to which of theit

Oregonian Province, D. californica is probably restricted to the shelf of the San Diego Province, reaching

i its northern limit in the immediate vicinity of Point Conception. Type Locality and Type Specimens. Between Balboa and Corona Del Mar, California, holotype

USNM no. 71440.

I Distribution. Off Southern California, San Diego to Point Conception, 19 - 127

m.

i 135

I

I

I D

I c I ,ii , /

I

I

:| ,

G _H

| '.

'| ,

,

I Figure 2.9. Diastylis californica Zimmer 1936. Female: A, carapace and pereonites, dorsal view; B, same, lateral

view; E, antenna 1; F, maxilliped 3; H, telson and uropods. Juvenile: C, carapace, dorsal view. Male:

i D, body, lateral view; G, telson and uropod. From Zimmer (1936); no scale provided.

I

I 137
!

Diastylis pellucida Hart, 1930 I

Figure 2.10

II

Material Examined. Cruise 1-1 (November, 1986), Sta. R-2 (81), R-3 (2), R-5 (1),PJ-1 (9); Cruise II

1-2 (January, 1987), Sta. R-2 (43), R-3 (3), R-5 (1), R-6 (3), R-9 (2), PJ-1 (20); Cruise 1-3 (May, 1987), Sta. 1

R-2 (61), R-3 (20), R-5 (2), R-6 (4), R-9 (t0), PJ-1 (1); Cruise 2-3 (October, 1987), R-1 (1), R-2 (14), R-6 1

(1), R-9 (11); Cruise 2-4 (January, 1988), Sta. R-2 (89), R-3 (5), R-5 (6), R-9 (10), PJ-1 (16); Cruise 2-5 m

(May, t988), Sta. R-2 (42), R-3 (2), R-5 (1), R-6 (7), R-8 (2), R-9 (27); Cruise 3-1 (October, 1988), Sta. R2 (5), R-3 (2), R-5 (1), R-6 (3), R-9 (8), PJ-1 (1); Cruise 3-4 (May, 1989), Sta. R-2 (24), R-3 (9), R-4 (1), R- l[

5 (3), R-6 (7), R-9 (66), PJ-1 (5). m

Description. Adult size, female 8 ram, male 9 mm. Carapace longer than free pereonites; surface II

smooth except for two oblique, minutely crenulate ridges running from mid dorsal ridge to ventral margin;

a third very slight circumcinct ridge occurs posterior to frontal lobe. Pereonite 1 visible ventrally; pereonite

IllP

5 posterior corners subacute. Antenna 1 peduncle article 2 extending near to end of pseudorostrum. Telson m

slightly more than half length of uropod peduncle; lateral margin with 5-8 setae. Uropod exopod longer than I

3-articulate endopod.

u

!

!

I

!

I

!

!

Figure 2.10. Diastylispellucida, Hart 1930. Male:A, carapace and pereonites,lateralview; B, body,lateral view; m

c, carapace and pereonites, dorsal view. Scale = 1mm. I

I

138 m
!

I Remarks. This species is the only diastylid in thisregion with just two lateral oblique ridges. It also

has a relatively thin integument, so the ridges in newly moulted individuals are not readily visible except in

i transmittedTypelight. Locality and Type Specimens. Deep Cove, Vancouver Island, British Columbia, 120 m.

Type in British Columbia Provincial Museum. Holotypes on slides are in very bad condition.

I Distribution. Southern California to British Columbia, Canada (Oregonian Province), 50 - 410 m.

Genus Leptostylis G. O. Sars, 1869

I Type Species. DiastyIis longimana Sars, 1865.

I may be coalesced Diagnosis. or(modified not; fifth from pereonite Day, not 1980) produced Carapaceposteriorly. usually tumescent Male antenna posteriorly. 1 with club-shaped Pereonites 3article and 4

3, with dense array of sensory setae distally. Male antenna 2 not extending beyond pereonite 5. Basis of

I pereopods 3 and 4 within rudimentary male not widened exopodsandorpereopods exopods 2absent and 3innotfemale, widelyfully separated developed in ovigerous in male.female. Male withPereopods 2 pairs

of pleopods. Post-anal part of telson shorter than pre-anal; with single pair of lateral setae sub-terminally.

i Uropod Renedopod marks. 3-articul The ate. short telson and form of the male antenna 1 characterize this genus. It is widespread

throughout the colder waters of the world's oceans but not as diverse as Diastylis. One species, Leptostylis

I least villosafrom, hasPuget previously Soundbeen south.recorded from the Oregonian Province, but it is likely this record is in error, at

I Leptostylis Watling McCann, new species

calva and

Figure 2.11

i I

l Material Examined. Santa Maria Basin, Cruise 1-1 (November, 1986), Sta. R-1 (64); Cruise 1-2

(January, 1987), Sta. R-1 (90); Cruise 2-3 (October, 1987), Sta. R-8 (49); Cruise 2-5 (May, 1988), Sta. R-1

(89), R-4 (56), R-8 (82); Cruise 3-1 (October, 1988), Sta. R-1 (39), R-4 (40), R-5 (4), R-8 (30); Cruise 3-4

I (May, 1989), Sta. R-1 (64), Ro8 (54).

Description. Adult size, 2.5 mm. Carapace slightly swollen posteriorly; triangular in dorsal view;

I vmargins ery slight extended ridge preposteriorly sent lateral partially to froncovering tal lobe; pereonite surface of1.carPereonites apace smo1oth, and 2withof normal few setae; widthposterol dorsallyateral but

narrowed ventrolaterally; pereonites 3 and 4 coalesced dorsally. Pereopods 3 and 4 with rudimentary exopods

I inlateral femalsetae. e. TeFemale lson abouropod ut half peduncle as long assetose uropodalongpeddistal unclestwo-thirds in male andof medial female;margin; with singrami le pair subequal; of subtendopod erminal

articles subequal; male uropods similar but more elongate.

I Remarks. This species is unusual in the extent to which the carapace extends posterolaterally. It is very similar to the following species, L. abditis, being most easily distinguished from the latter by its

subequal uropod rami.

I Type Locality and Type specimens. California, Santa Maria Basin, off Point San Luis, Sta.R-1

35°05.83'N, 120°49.16'W, 91 m, holotype USNM 273587, paratypes USNM 273588 and LACM 95-80.1.

Etymology. from L. caIvus, = bald, bare, indicating the lack of setae on the carapace.

I Distribution. Known so far only from the Santa Maria Basin region, primarily at depths near 90 m.

It is probably an abundant member of the shallow shelf fauna.

I

I 139
!

!

!

i

!

!

i

7 ii .

i ,i,,i i i

'' ..... i i ._ i

, , 'iI

', _"-° i j _ ,,

,,,_:._, I _'_ " II

0,, ",L" _,'_ . i

_ _,,,_. . ,,

_i__; , ',_ ',',_

__" )_ _"_ if

/

i,,:F" ."

,I'"\,'

/ I '_

I

Figure 2.11. LeptosCylg_colva Watlmg and MeC_, new species. Female: A, body, lateral view; B, carapace

andpereonites, lateral view; C, same, dorsal view; D, teslon and uropods. Male: E, telson anduropods, l

Scale, A-C = 1 mm, D,E = 0.5 mm. It

I

Leptostylis abditis Watling and McCann, new species

Figure 2.12 H

Material Examined. Santa Maria Basin, PhaselI, Cruise I-1 (November, 1986), Sta. R-5 (25);

Cruise 1-2 (January, 1987), Sta. R-5 (13); Cruise 2-3 (October, 1987), Sta. R-5 (14), PJ-1 (18); Cruise 2-4 I

(February, 1988), Sta. PJ-1 (6); Cruise 2-5 (May, 1988), Sta. R-3 (1), R-4 (2), R-5 (6), PJ-1 (1); Cruise 3-1 II

(October, 1988), Sta. PJ-1 (1); Cruise 3-4 (May, 1989), Sta. R-1 (1), R-4 (6), R-5 (18), PJ-1 (15).

Description. Adult size, 3.5 mm. Carapace swollen posteriorly into tumescent lobes; anteroventral N

margin below antennal sinus with coarse serrations; surface of carapace smooth, with few setae; posterolateral ll

margins extended posteriorly to cover pereonite 1 laterally. Pereonites 1 and 2 of normal width dorsally but

narrowed ventrolaterally; pereonites 3 and 4 weakly coalesced dorsally. Pereopods 3 and 4 without I

rudimentary exopods in female. Telson less than half length of umpod peduncle. Uropod exopod shorter i

than endopod, reaching just beyond end of endopod second article; endopod terminal article longer than

second. Male uropod similar but more elongate. B

U

Remarks. This species is superficially similar to L. caIva, especially in general aspects of the

carapace and body shape. However, the unequal rami of the uropods and the lack of rudimentary exopods on Illlll

pereopods 3 and 4 of the female serve to readily distinguish it from L. calva. 1

|
I

I

I

I

I

I

I

I

I

I

I ,D

i ,._. ./#

/

I

I

I

Figure 2.12. Leptostylis abditis Wafting and McCann, new species. Female: A, body, lateral view; B, carapace

and anterior pereonites, lateral view; D, carapace and pereonites, dorsal view; E, telson and uropods.

I Male: C, carapace and antenna 1, dorsal view. Scale, A = 0.5 mm, B-D = 0.2 mm, E = 0.5 mm.

I

I

i 141
!

Type Locality and Type Specimens. California, Santa Maria Basin, off Purisima Point, Sta. R-5 I

34°42.69_, 120°50.83'W, 154 m, holotype USNM 273585, paratypes USNM 273586 and LACM 95-79.1.

Etymology. from L., abditus = hidden, concealed, referring to the initial difficulty in separating I this species from L. calva.

Distribution. A somewhat rare species of shelf and slope, known from the Santa Mafia Basin

region, primarily at depths of 145-410 m., to off Pt. Loma, San Diego as shallow as 47m. I

Family Lampropidae I

Diagnosis (from Day, 1978). Mandibles naviculoid. Maxilla 1 palp absent or bearing 1 or 2 setae.

Exopods present on maxilliped 3 and pereopod 1inboth sexes; in female present or rudimentary on pereopods 1

2-4 or absent from all three; in male exopods present on pereopods 2-4. Pleopods absent or 1-3 pairs in II

male, with outer process on inner ramus. Telson moderate to large, with post-anal part longer than pre-anal,

with 3 to 8 terminal and subterminal setae. I

Remarks. Lampropids are most easily distinguished on the basis of possessing atleast three terminal

setae on the telson. The family presently comprises 10 genera and is well represented in the cold waters of

the world' s oceans, from shallow depths to the abyss. The taxonomic history of this family has been quite 1

clear, with the exception of Stebbing's (1912) division of the group into five families on the basis of male

w

pleopod number. Given (1964) followed Stebbing's example and created the family Mesolampropidae for Im

species where the male possessed two pairs of pleopods. This arrangement of the lampropids has not been 1

followed by any other workers and is not accepted here. u

Genus Hemilamprops Sars, 1883 I

Type Species. Vaunthompsonia rosen Norman, 1863.

i

Diagnosis (From Day, 1978). Carapace not strongly dorsoventrally flattened. Eye present or absent. I

Pseudorostrum short. Antenna 1 flagella well developed. Maxilla 1 palp with 2 terminal setae. Exopods on

pereopods 3 and 4 rudimentary in female. Male with 3 pairs of pleopods. Telson usually with 3 terminal and 1

often 2 subterminal setae. I!

Remarks. Hemilamprops Sars, Mesolamprops Given, and Lamprops Sars are very closely related

genera. The only definitive method of distinguishing these genera at the present time is on the basis of the •

number of pleopod pairs in the male, viz., Hemilamprops has three, Mesolamprops has two, and Lamprops

none.

Hemilamprops californicus Zimmer, 1936 1

Figure 2.13 1

HemiIamprops (?) californica Zimmer, 1936.

Lampropoides californicus, Harada, 1959. !

II Hemilamprops californiensis, Lie, 1969 (lapsus calami).

Material Examined. Santa Mafia Basin, Phase II, Cruise 1-1 (November, 1986), Sta. R-1 (t); I_

Cruise 1-2 (January, 1987), Sta. R-1 (3), R-4 (23), R-8 (61); Cruise 1-3 (May, 1987), Sta. R-1 (2), R-3 (43),

R-4 (4), R-8 (111); Cruise 2-3 (October, 1987), Sta. R-1 (1), R-8 (18); Cruise 2-4 (February, 1988), Sta. R4 (15), R-8 (3); Cruise 2-5 (May, 1988), Sta. R-4 (1), R-8 (4); Cruise 3-1 (October, 1988), Sta. R-1 (24), R- •

4 (8), R-8 (33); Cruise 3-4 (May, 1989), Sta. R-4 (7), R-8 (12). II

142 I
I

I 'l B

I \, /i" J

I Figure 2.13. Hemilamprops and uropods. Scale, californicus A = t ram, Zimmer B = 0.5 1936. mm. Female:A, carapaceandpereonites,lateral view;B,telson

I

I_eripfion. Adult size, female, 5.5 to 8 ram, male 6 ram. Carapace as long as first 4 pereonites;

I broadly median truncate carina onanteriorly; frontal lobe; withocular singlelobe ridgedistinct, runninglarge, from with pseudorostral 7 lenses; margin pseudorostral upwardlobes and backward extend onlyto

short distance in front of ocular lobe. All 5 pereonites visible in lateral view; pereonites 2 and 3 subequal in

I length. longer than Telsonexopod, as longproximal as uropod articlepeduncles, longer thanwithdistal3 apical 2 combined, ad 2 distal dorsalarticle subapical the shortest; setae. Uropod exopod endopod reaching

to midpoint of endopod distal article, 2 articles subequal in length, external margin with heavier setae than

I on inner margin.

I Remarks. Zimmer did not have a male in his collection when he described the species, but assigned

it questionably to Hemilamprops on the basis of general features. Given (1964) noted that the specimens he

I Harada identified(1959) as H. and californicus Gam6 (1962). had three Harada pairs(1959) of pleopods, createdasthedidnewthe genus specimens Lampropoides identified astothis accommodate species by

several new species wherein the male possessed three pairs of pleopods and a recurved fold on the carapace.

I He as not inclubeing ded Zimmer's sufficientlyHemilamp distinguishable rops (?) calif fromornicaHemilamprops in this new. geCurrently, nus. Gam6 the (1962) closely dismisse related d thegenera genus

Hemilamprops, Lamprops, and Mesolamprops are defined on the basis of the number of pleopods present

I oHowever, n the male.Gladfelter Therefore,(1975) on thedescribed basis of male a newpleopod species,number, Mesolamprops this species dillonensis belongs, ininwhich the genus theHemilamprops male possessed.

two pairs of pleopods, but the female is virtually indistinguishable from H. californicus. In the material at

I hand, the description there is alsofo Ha.male californicus with tw.o There pairs oisf pleo thuspoadsmajor accompanying problem in twneedo ovigerous of solution. females Are there that als twoo fit closely well

related species assignable to two different genera (on the basis of male pleopod number), or is it possible for

i omore ne species to have two male morphologies? Perhaps Harada's Lampropoides concept needs to be given a

thorough examination, and all species redefmed on other characters. For the present time, this species

will be left in the genus Hemilamprops, in order to avoid needless cluttering of the synonymy until the

i necessaryTystudies pe Locaarelitycoand mpleted. Type Specimens. Between Balboa and Corona Del Mar, California. Holotype

USNM 71439.

I Distribution. Puget Sound, Washington to Southern California, Japan, 13-60 m.

i 143
!

Family Leuconidae I

Diagnosis (modified from Jones, 1963). Mandible truncate, molar usually present and columnar. •

Maxilla 1 palp bears single seta. Exopods present on maxilliped 3 and pereopods 1-3 (occasionally on

pereopods 1-2) in both sexes, additional exopod onpereopod 4 on most males. Pleopods absent, rudimentary,

or 2 pairs in male, without external process on inner ramus. Telson absent. Eye generally absent; if present •

may be either fully developed or rudimentary.

Remarks. The Leuconidae is one of the oldest cumacean families, having been established by Sars

(1878). Until Watling's (1991a) revision, it had consisted of only eight genera, all but one of which were i

proposed by 1907. The latest revision expanded the family so that it now comprises 12 genera, and five

subgenera in the genus Leucon. The present material adds one new genus, a new subgenus, and 4 new

species, i

Key to the Leuconidae of the Santa Maria Basin Region •

|

1A. Efferent orifice anterior or anterodorsal .......................................................................................... 2 i

U lB. Efferent orifice distinctly dorsal, pseudorostral lappets bent posteriad and directed dorsally ....... 6

2A. Male antenna 1 short, not reaching posterior margin of carapace; male without pleopods ............. i

............................................................................................................. AlIoeoleucon santamariensis

2B. Male antenna long, extending well past carapace posterior margin; male with 2 pairs of pleopods i

.................................................................................................................... Leucon ...................... 3

!

3A. Branchial siphon longer than pseudorostral lobes; male pleopods reduced (Subgenus i'

Diaphonoleucon ) ............................................................................................... Leucon (D.) declivis

i

3B. Branchial siphon normal, barely extending beyond pseudorostrum; male pleopods normal ........... •

.................................................................................................................... (Subgenus Leucon) .... 4

l

4A. Carapace with forwardly curving ventrally directed lateral ridge ................. Leucon (L)falcicosta R

4B. Sides of carapace without ridge ...................................................................................................... 5

5A. Pseudorostral lobes short, bluntly truncate anteriorly ....................................... Leucon (L.) armatus I

5B. Pseudorostral lobes elongate, subacute anteriorly, slightly upturned ...............................................

I

Leucon (L.) magnadentatus 1

6A. Antenna 1 geniculate between peduncle articles 1 and 2; carapace with 2 lateral, strongly curving •

ridges ....................................................................................................... Eudorellopsis longirostris II

6B. Antenna 1 geniculate between peduncle articles 2 and 3; carapace without lateral ridges ..............

.................................................................................................................... Eudorella ................... 7

I

!

144 i
!

I 7A. Antenna 1 main flagellum with setae only at distal end; uropod peduncles with tong marginal setae;

pereopod 5 absent ........................................................................................ Eudorella redacticruris

I 7B. Antenna ............................... 1 main flagellum , .... , .................................................... with setae along whole margin;, ............................................................... pereopod 5 normal in male and female8

I 8A. to end of endopod basal article; ventral margin bent, posterior Uropod exopod reaching only carapace

section smooth, anterior section with serrations ................................................ Eudorella pacifica

i 8B. Uropod exopod extending beyond end of endopod basal article; carapace ventral margin continuously curved, anterior section with fine serrations ................................................... Eudorella truncatula

I Alloeoleucon, new genus

Type Species. Alloeoleucon santamariensis Wafting and McCann, new species

I Diagnosis. Pseudorostrum projecting anteriorly. Antenna 1 not geniculate. Male antenna 2 extending

only to posterior margin of carapace; peduncle articles 4 and 5 without setal brush; flagellum not modified

I for endopod grasping. 2-articulate. Exopods Male on perewithout opods pleopods. 1-3 in female and 1-4 in male. Pereopod 2 article 3 very short. Uropod

Remarks. This peculiar genus resembles Hemileucon, Austroleucon, Heteroleucon, andNippoleucon

I having in its lackneither of pleopods a strongandsetalshor brush tenedonantenna the antennal 2 of thepeduncle male. Howe nor aver, flagellum it differsmodified from most for grasping. of this group In this by

latter respect it is closest to Heteroleucon, but the latter is additionally modified in possessing exopods only

I on pereopods 1 and 2 in both sexes. Etymology. from Gr., aUoios, of another kind, different, + leucon, the stem genus, referring to the

unusual collection of features characterizing this genus.

I

Alloeoleucon santamariensis Wafting and McCann, new species

I Figure 2.14

Material Examined. Santa Maria Basin, Phase/I, Cruise 1-1 (November, 1986), Sta. R-3 (12);

I Cruise 1-2 (January, 1987), R-3 (8), R-4 (4); Cruise t-3 (May, 1987), R-3 (2), R-6 (1); Cruise 2-3 (October, 1987), R-3 (6); Cruise 2-4 (February, 1988), R-3 (4); Cruise 2-5 (May, 1988), R-3 (2); Cruise 3-1 (October,

1988), R-3 (2); Cruise 3-4 (May, 1989), R-3 (5), R-9 (1).

I Adult 2 male and female, in both than Description.

size, mill, Carapace sexes slightly longer

pereonites; median carina sparsely, coarsely, serrate; pseudorostral lobes bluntly truncate; eyelobe not

distinguishable from frontal lobe. Antenna 1 not geniculate; accessory flagellum minute. Male antenna 2,

I flagellum short, not extending beyond pereonites; peduncle articles 4 and 5 without brush of setae,

sensory

article 5 with 4 long proximal setae. Maxilliped 3 leg-like, distal articles slender, basis not strongly expanded

distally, beating only four setae. Pereopod 1 articles 5 and 6 subequal in length; basis armed with single,

I long, plumose seta distally. Pereopod 2 with short article 3. Uropod similar in male and female; endopod longer than exopod, 2-articulate, articles subequal in length; exopod with 1-2 subterminal setae on inner

margin.

I Type Locality and Type Specimens. California, Santa Maria Basin, Luis, R-3,

off Point San Sta

35°05.30'N, 121°00.90'W, 409 m, holotype USNM 273583, paratypes USNM 273584 and LACM 95-78.1.

I

i 145
,46

I
I

I Remarks. This is the smallest leuconid found in Californian waters. The species is readily

distinguishable from the other leuconids found in the Santa Maria Basin area by the truncate pseudorostral

i lobes andEtymol the very ogy.similar This species appearance is named of theformaleits and principal female.place of occurrence, the Santa Mafia Basin of

southern California.

I one recordDistartib92utionm. . Known thus far only from the Santa Mafia basin area, depth range, 409-410 m, with

I Genus Leucon KrOyer, 1846

Type Species. Cuma nasica Kr0yer, 1841.

I Diagnosis (from Wafting, 1991a). Pseudorostrum projecting anteriorly, usually slightly shorter in

male; antenna 1 not geniculate; male antenna 2 with brush of setae on anterior margin of peduncle articles

i 4withandexopods 5; male antenna on pereopods 2 flagellum1-4; pereopod extending 2well articlealong3 very pleon;short female or lacking; with exouropod pods onendopod pereopods2-articulate; 1-3; male

male with 2 pairs of pleopods.

I relative Rlength emarks. of theThisbranchial genus wassiphon, subdivided the length by Wafting of the antennule (1991a) intaccessory o 5 subgenera, flagellum, usingtheaspresence criteria theof

ventral hooks on pereonite 5, and the degree of fusion of the uropod endopod terminal seta. A new subgenus

I isofdescribed the pleopods belowin which the mate. has a new combination of the above characters, but as well, shows extreme reduction

I Diaphonoleucon, new subgenus

Type Species. Leucon (DiaphonoIeucon) declivis Watling and McCann, new species

I Diagnosis. Branchial siphon elongate, longer than pseudorostral lobes; antenna 1 accessory flagellum

minute; pereonite 5 without ventral teeth; uropod endopod terminal seta not fused with distal article; pleopods

of male reduced to short peduncle bearing nub-like rami.

I Remarks. The combination of elongate branchial siphon and minute antennule accessory flagellum,

is sufficient to distinguish this subgenus from the others recently established by Watling (1991a). The

i pre Ledoyer, sence owhich f reduwas ced pleopods is not unique to this subgenus, having also been seen in L. serrulirostris assigned to the subgenus Crymoleucon on the basis of its short branchial siphon and

long antennule accessory flagellum.

I differentEtymology form of the. fmal rome. Gr. diaphonos, dissonant + leucon, the stem genus, referring to the strongly

I Leucon (Diaphonoleucon) declivis Watling and McCann, new species

Figure 2.15

I Material Examined. Santa Mafia Basin, Phase II. Cruise 1-1 (November, 1986), Sta. R-7 (5);

Cruise 1-2 (January, 1987), R-7 (1); Cruise 1-3 (May, 1987), R-7 (8); Cruise 2-3 (October, 1987), R-7 (7);

I Cruise 2-4 (February, 1988), R-7 (2); Cruise 3-1 (October, 1988), R-7 (1); Cruise 3-4 (May, 1989), R-7 (10). Description. Size of adult female, 9.0 mm, male, 9.5 mm. Carapace pseudorostral lobes upturned,

about one-fifth total carapace length; dorsal crest serrate only along anterior half, serrations conspicuous on

I frontal lobe and on posterior extension of frontal lobe, often with middorsal gap; antennal notch narrow,

i 147

!

I toothed along ventral margin of pseudorostrai lobe; anteroventral comer acute. Branchial siphon much

longer than pseudorostral lobes. Antennule elongate, thin, extending beyond end of pseudorostrum, accessory

i flagellum Uropod minute. Pereopod 1 article 5 longer than article 6. Pereopod 2 article 3 nearly as long as wide. rami longer than peduncle; endopod longer than exopod, distal article one-fourth length of basal

article, both articles setose along both inner and outer margin; exopod strongly setose along both margins.

i Male pleopods reduced, 2 pairs, each consisting of shortened peduncles bearing nub-like ramal remnants. Antenna 2 of final male stage unknown, but sub-adult shows many compressed articles forming within old

cuticle.

I very soft,Remarks. low oxygenThismudis habitat one of the at thelargest upperleuconid reaches ofcumaceans the oxygenknown. minimumIt appears zone. None to beofconfined the specimens to the

collected were fully mature, so it is possible that the reduced pleopods of the male will eventually be found

i tocuticle developof thesmall onerami specimen since there that appears is an indication ready toofundergo a biramous its f'mal condition molt. There which iscanno beindication, seen underhowever, the old

that these pleopods will be fully formed as is the case in the genus generally.

I 34°52.90'N, Type121loc°10.30'W, ality and565Type m, holotype SpecimeUSNM ns. Californi 273581,a, paratypes Santa Mafia USNMBasin, 273582off and PoinLACM t Sal, Sta 95-77.1. R-7,

Etymology. From L. declivis = downhill, sloping, referring to the slope habitat in which this species

I lives.

Distribution. Known only from the deeper waters of the Santa Maria Basin, at least 565 m.

I Subgenus Crymoleucon Wading, 1991a

Diagnosis. Branchial siphon normal; antenna 1accessory flagellum extending at least to midlength

I of main flagellum fLrStarticle; pereonite 5 without ventral teeth; uropod terminal seta not fused to distal article.

Remarks. Most of the species assigned to this subgenus are found in the deeper or colder polar

I waters of the world ocean. Otherwise, it is a rather heterogeneous assemblage of species united by the

unusually large accessory flagellum.

I Leueon (Crymoleucon) bishopi Bacescu, 1988

i Figure 2.16

EpiIeucon pacifica Jones, 1969

I Epileuco Leucon n(Epileucon) pacifica, Bbishopi ishop, Bacescu, 1981 1988

I Sta. R-7Ma(2);teriCalruiseExa1-3min(May, ed. Santa 1987),Maria R-7 Basi (2); nCruise , Phase2-4I, Sta. (Feb6ru3ary, , (1);Ph1988), ase II, R-7Cruise (1); Cruise 1-2 (Jan3-1uary, (Oc1987), tober,

1988), R-7 (2); Cruise 3-4 (May, 1989), R-7 (2).

I Description. Size of adult female, 6 mm. Carapace pseudorostral lobes directed anteriorly, less than one-fifth total carapace length; dorsal crest without serrations in female and male; anteroventral corner

slighting projecting, with several teeth below antennal notch. Branchial siphon not projecting beyond end of

I pse slightly udorostrum. beyond Antenna mid-length1 short, of mainpedflagellum uncle articles basa2l and article. 3 subequal Pereopodinlength; 1 articleaccessory 5 slightlyflage shorter llumthanextending article

6. Pereopod 2 article 3 shorter than wide. Uropod peduncle as tong as endopod basal article; exopod basal

!

i I49
I

I ", A

/" _i. "\ ,_ C

' I

]' I ,,, ,' ....-,-,

, I /t

" J ,' .;.

," i I

,,' ... .., i -;

.., . _._ ,/' ," ._ ,,' i ,', ",, _

," .'.K.. i' .

...... _ _r..__.._ l , _'J '_ ", "

__ /g ' '-- _;s ' '_l ,',_-

._- .,<._ , '_ -.. :_,:fl '_ ' ', ' |

/ _j ",,',* r:'1 '_r'., ,.1 ', _.;,_, ' ._ ,- _:' r, '_" "'

..¢

._.:","...

.M',/i i ] '.\ '--.'" ',I. '. , '.', ,', -'_It _:', , ',",G ,

• ' "''_ "# .!,. ..._ ,. ,, , .,,_.,. -

_'" _Ii_I. _" !'_;" I

-?,,' ,, _,\ "_":,..\ t ,.-_:..__ ./..... ., ..........

i _'. _...,, _"_' .'"_

, "_-"_.,\ v_:_ _._." ,t,_iH:: _,,:_

/,,: , ._Q, \ _. _" ....... _,_:_..__.t,:_,._:_._,'"_:_,_! ' ':"!_'.i!!:_\

., . - <....,j,...,;. \ \ _-'- ....._i.'_ [_.j_"_),t'_.X _.,,,,_ ; _.}

!.....-?_:.> _\ ," .,'::: !_o \ \ '/,'k I

1 " _ \

" ", !

Figure 2.16. Leucon (Crymoteucon) bishopi Bacescu, 1988. Subadult male: A, pereopod 1; B, pereopod 2; C,

uropods. Scale, A-C = 0.5 ram. I

I

article long, about half length of distal article; exopod extends at least to end of endopod basal article; •

endopod armed medially with continuous series of narrowly spaced setae, 3 on distal article.

Remarks. This species was assigned to the genusEpileucon Jones, 1969, because it lacked serrations

on the dorsal crest in the female. Bacescu (1988) and Wading (1991a) relegated EpiIeucon to subgeneric l

status because the constituent species could not be characterized by any feature other than the presence of

ventral teeth on pereonite 5. Bishop (1981) noted that E. pacifica did not possess these ventral teeth and

subsequently Bacescu (1988) moved E. pacifica into Leucon (Epileucon). Since the species L pacifica was I!

an occupied name, Bacescu proposed L. (E.) bishopi for Jones' E. pacifica.

Type Locality and Type Specimens. Galathea station 745, 7 ° 15' N, 79 ° 25' W (Gulf of Panama),

915 m, T 5°C; holotype in British Museum. 1

I

Distribution. Previously known only from the type locality; the present specimens extend the range

of the species to the north and further up the slope. L. (C.) bishopi seems to be characteristic of the upper I

reaches of sediments in contact with low oxygen content water. •HI

Subgenus Leucon Kroyer, 1846 1

Diagnosis. Branchial siphon normal; antenna 1 accessory flagellum rudimentary; pereonite 5 without

ventral teeth; uropod endopod terminal seta not fused to terminal article. n

Remarks. This is the most wide-spread of the subgenera and embodies most of the northern Atlantic 1

and sub-Arctic species included in the monograph of Sars (1900) from Norway.

I

150 I
!

I Leucon (Leucon)falcicosta Watfing and McCann, new species

Figure 2.17

I Material Examined. Santa Mafia Basin, Phase II. Cruise I-1 (November, 1986), Sta. R-2 (18), R4 (3), R-5 (3), R-6 (1), PJ-I (77); Cruise 1-2 (January, 1987), R-1 (1), R-2 (15), R-5 (3), PJ-1 (19); Cruise 1-

I 3R-2(May, 1987), R-1 (2), R-2 (52), R-4 (6), R-5 (9), R-8 (5), PJ-1 (41); Cruise 2-3 (October, 1987), R-1 (3), (15), R-8 (4), PJ-1 (29); Cruise 2-4 (February, 1988), R-2 (28), R-4 (14), R-5 (10), R-8 (3), PJ-t (36);

Cruise 2-5 (May, 1988), R-1 (4), R-2 (25), R-4 (8), R-5 (9), R-8 (9), PJ-1 (12); Cruise 3-1 (October, 1988),

I R-1 R-8 (4), (11),R-2PJ-1(10), (52).R-5 (4), R-8 (1), PJ-1 (14); Cruise 3-4 (May, 1989), R-1 (9), R-2 (14), R-4 (10), R-5 (10),

Description. Adult size, female and male, 3 mm. Carapace pseudorostral lobes subtriangular distally,

I less than one-fifth length of carapace; dorsal crest serrate anterior two-thirds, with approximately 15 serrations; with single anteriorly-directed curved ridge extending from posterior of dorsal serrations ventrally to region

behind pseudorostral cut; antennal notch deep, anteroventral comer blunt. Antennule accessory flagellum

I Maxilliped nearly half length of basal article of main flagellum; peduncle third article 1.25 times length of article 2. basis distal margin not strongly extended laterally and bearing only 3 long plumose setae; ischium

with additional long plumose seta. Pereopod 1 article 5 longer than article 6; exopod articles as long as

I basis. Pereopod 2 article 3 present, about half as long as wide; merus and carpus subequal in length. Uropod exopod slightly longer than endopod, inner margin armed with few (4-6), long, sparsely setulose setae;

endopod distal article three-fourths length of basal article, inner margin of both articles armed with short,

I stiff setae. Remarks. This species is superficially similar to L armams Given, but differs from the latter in the

strength of the dorsolateral curved ridge, which is noticeable as a very sharp line in L falcicosta, but only as

I a low, raised feature inL. armatus. The shape of the pseudorostral lobes and the uropod armature also differ in the two species. The only other species of Leucon possessing a curved ridge on the carapace is L. (L.)

kobjakovae Lomakina, 1955. It differs from L. (L.)falcicosta by having a more blunt pseudorostrum in the

I male, 1 and2.fewer setae along the inner margin of the uropod, and simple rather than plumose setae on pereopods

Type Locality and Type Specimens. California, Santa Maria Basin, off Point San Luis, Sta. R-l,

I 35°05.83'N, 120°49.16'W, 91 holotype USNM 273579, USNM 273580 and LACM 95-76.1.

m, paratype

Etymology. From L.falcis = sickle, and costa = ridge, signifying the sickle-shaped nature of the

i dorsolaterDistribution al ridge on. theKnown carapacfrome. the Santa Maria Basin region, 90-161 m (with a single record from 410

m), and off Pt. Loma, San Diego, from one specimen taken at 116 m.

I Leucon (Leucon) armatus Given, 1961

i Figure 2.18 Material Examined. California: Santa Maria Basin, Phase II, Sta. PJ-1.

i dor Description. Adult size, female, 5.5 mm. Carapace pseudorostral lobes subquadrate distally; somedian crest serrate nearly throughout entire length; dorsolateral depression extends from dorsomedian

crest anteroventrally to just behind origin of pseudorostral lobe; slight dorsolateral depression also present

i posteriorly. Pereopod 1 article 5 longer than 6. Pereopod 2 article 3 absent. Uropod endopod stout and broadened, as long as or longer than exopod with dense array of setae along inner margin.

Remarks. This species is the only member of the genusLeucon present in California waters to have

I sub species quadrat remain e pseuslightly dorostralin dlooubt. bes. As yet, an adult male has not been described, so the exact affinities of the

i 151

!

!

!

!

!

!

I Figure 2.18. Leucon body, lateral (Leucon) view.armatus Scale= Given, 1 ram. 1961.Female: A, carapace and pereortites 1-3,lateral view; B,

!

Type Locality and Type Specimens. Velero IV sta. 4851-57, Mugu Canyon, 7.75 miles from Port

I Hueneme Light, 172 m; holotype AHF 5726, in Los Angeles County Museum. Distribution. Southern California, in sandy silt, 145-187 m.

I Leucon (Leucon) magnadentata Given, 1961

Figure 2.19

!

Material E_ed. Santa Maria Basin, Phase 1I. Cruise 1-1 (November, 1986), Sta. R-3 (87), R6 (4);Cruise 1-2 (January, 1987), R-3 (66), R-5 (1), 1t-6 (1), R-9 (12); Cruise 1-3 (May, 1989), R-3 (135),

I R-6 (3), R-9 (23); Cruise 2-3 R-3 R-6 R-9 (26); Cruise 2-4 R- (October, 1987), (2), (February, 1988),

3 (118), R-6 (3), R-9 (26); Cruise 2-5 (May, 1988), R-3 (41), R-6 (1), R-9 (44); Cruise 3-1 (October, 1988),

R-3 (137), R-6 (2), R-9 (20); Cruise 3-4 (May, 1989), R-3 (195), R-6 (1 I), R-9 (41).

I Description. Size of adult female 5.5 mm, male 6.0 mm. Carapace longer than pereonites;

pseudorostral lobes acute distally, slightly upturned; dorsomedian crest with 7-10 large serrations;

i anteroventral margin below antennal notch with large teeth. Pereopod 1 basis with large serrations; article 5

longer than article 6. Pereopod 2 article 3 absent. Uropod endopod 2-articulate, basal article the longer;

endopod shorter than exopod. In mature male, carapace pseudorostral lobes more truncate than in female,

serrations on dorsal crest reduced; pereonites 4 and 5 with exaggerated pleurites; uropod exopod and endopod

I with additional medial

setae.

Remarks. Until this study, this species was known only from four specimens, all of which were

i juveniles. carapace ofThethe species juvenilesis and distinguished immature females by the very and males. large serrations, both dorsally and ventrally, on the

Type Locality and Type Specimens. R/V VeIero IV Sta. 5762-58, 8.6 miles 310.5 °true from Point

I Loma Naturalight History (nearMuseum). San Diego, California), 575 ft (208 m). Holotype AHF 585 (now in Los Angeles County

Distribution. Southern California including San Diego and southern part of Oregonian Provinces,

I 200-410 m.

i 153
I

I

I

I

I

I

I

I

I

I

Figure 2.19, Leucon (Leucon) magnadentata Given, 1961.Female: A, carapace and anterior pereonites, lateral I

view; B, body, lateral view; C, carapace and anterior pereonites, dorsolateral view.Scale = I mm.

I

Genus EudoreIla Norman, 1867

Type Species. Eudora truncatula Bate, 1856. I

Diagnosis (from Wafting, 1991a). Pseudorostrum directed dorsally in both male and female; antenna

peduncle 1 geniculatearticles between 4 andpedunc 5; malele antenna articles 22 and 3; male antenna 2 with brush of setae on anterior margin of I flagellum extending nearly to end of pleon; female with exopods

on pereopods 1-3; male with exopods on pereopods 1-4; pereopod 2 article 3 lost; uropod endopod 2-

articulate; male with 2 pairs of pleopods. I Remarks. This is a cold-water genus represented by approximately 27 known species. The genus

has a very conservative body plan, consequently, species are often differentiated on the basis of a narrow

suite of features and correct identification of species may be difficult. If the specimen at hand does not fit I

the species described here, the key to the known species of the world (Watling, 1991a) should be consulted.

Any species taken south of the immediate area encompassed by this Atlas are likely to be new to science.

I

I

154 I
I

I

I

I

I

I

I

I

I

I

I Figure 2.211. Eudorella and anteriorpacifica pereonites, Hart,dorsal 1930.view; Female: D, A, carapace carapace, andlate pereonites, ral view;

lateral B, bodview. y,latera Scalel viA, ew;CC,= 1cara mm, pace B,

D= 1 mm.

I

EudoreUa paeifiea Hart, 1930

I Figure 2.20

EudoreUa tridentata Hart, 1930.

I Material Examined. Santa Maria Basin, Phase II. Cruise 1-1 (November, 1986), Sta. R-1 (14), R2 (33), R-3 (24), R-4 (79), R-5 (53), R-6 (1), PJ-1 (45); Cruise 1-2 (January, 1987), R-1 (14), R-2 (35), R-3

I (15), R-6 R-4 (46), R-5 (54), R-9 (12), PJ-1 (18); Cruise 1-3 (May, 1987), R-1 (21), R-2 (30), R-4 (84), R-5 (62), (2), PJ-1 (31); Cruise 2-3 (October, 1987), R-1 (28), R-2 (20), R-3 (33), R-4 (153), R-5 (117), R-8 (1),

PJ-1 (36); Cruise 2-4 (February, 1988), R-1 (34), R-2 (28), R-3 (20), R-4 (168), R-5 (159), R-8 (1), PJ-1

I (43); (23); Cruise Cruise 2-5 3-1 (May, (October, 1988), 1988), R-1R-1(30),(51), R-2 R-2 (38),(22), R-3 R-3 (10),(67), R-4 R-4 (138),(208), R-5 R-5 (120),(36), R-8R-8(8),(12), R-9 (61), PJ-1 (30); PJ-1

Cruise 3-4 (May, 1989), R-1 (61), R-2 (32), R-3 (66), R-4 (156), R-5 (64), R-6 (2), R-8 (21), PJ-1 (35).

I Description. Female 4.5 mm, male 6 mm. Antennule peduncle articles 2 and 3 subequal in length; main flagellum basal article with setae along inner margin; accessory flagellum as long as main flagellum

basal article. Pereopod 1 article 5 shorter than article 6. Pereopod 2 article 5 about 1.5 times length of article

I 4. Uropod exopod much shorter than endopod, only as long as endopod basal article.

i 155
!

Remarks. This species shows considerable variation, especially with respect to the teeth in the •

vicinity of the anteroventral comer and associated notch (Bamard and Given, 1960). Additionally, variation |

in the length of appendage articles and degree of setation has also been seen, both of which undoubtedly led

Hart to create the new species E. tridentata. In the Santa Maria Basin area E. pacifica is very common and I1

exhibits varying degrees of calcification as well as size of adults. 1

Type Locality and Type Specimens. Deep Cove, Vancouver Island, British Columbia; holotype

in British Columbia Provincial Museum. •

Distribution. Very common throughout San Diego and Oregonian Provinces, 20-410 m. I

Eudorella redacticruris Watling and MeCann, new species II

Figure 2.21

I

Material examined. Pac Baroness wreck (34°21.43'N, 120°38.29'W, 410-436 m) and control 1

(34°25.01'N, 120°41.51a_V,432 m) sites, off Point Conception, 135 specimens.

Description. Size of adult female and male, 3.5-4.0 mm. Carapace in both male and female longer I

i

than free pereonites; anteroventral angle marked with strong tooth; antennal notch broad and shallow, with I

only a single tooth marking dorsal boundary in female, no tooth in male. Pereonite 1 visible dorsally,

submerged laterally; pereonites 4 and 5 with lateral pleural extension in male. Antennule peduncle articles 1

2 and 3 subequal in length; accessory flagellum two-thirds length of main flagellum basal article, outer 1

margin of latter with two setae distally. Pereopod 1 articles 5 and 6 subequal in length. Pereopod 2 article 3 1

absent; article 4 one-half the length of article 5. Pereopod 5 absent inboth sexes; male with strongly developed 1

penial lobes. Uropod exopod nearly as long as endopod; terminal seta not fused to endopod distal article; I1

endopod basal article armed with long setae proximally and short, microsetulate setae distally.

Remarks. This species is unique in the genus in having lost the fifth pereopod; however, since the l

m

animal otherwise is indistinguishable from Eudorella, the erection of a new genus for this aberrant form

seems unwarranted. The presence of penial lobes is also quite rare in the Cumacea. Two other species with 1

penial lobes are known, Campylaspis rowei, for which the new genus, Campylaspenis, was originally erected I

by Bacescu & Muradian (1974) and Archaeocuma peruanum, used to establish the new family

Archaeocumatidae by Bacescu (1972). Ledoyer (1988) was not convinced that Campylaspenis, with the I

exception of the penial lobes, was sufficiently different from the remaining Campylaspis species to warrant •

the erection of a new genus; and Day (1988) noted that Archaeocuma possessed many of the characters of I

the Lampropidae. In general, single character differences, even though they seem significant, have not I

resulted in the erection of new genera, but this view may change as more of these aberrant forms are 1

discovered.

Type Locality and Type Specimens. California, Santa Barbara Co., offPt. Conception, 34°21.43'N, I

120°38.29'W,430m, holotype USNM 273577, and paratypes USNM 273578 and LACM 95-75.1. 11

Etymology. From L. redactus = edited, abridged, and cruris = leg, referring to the highly reduced

form of the fifth leg.

Distribution. Known only from the type locality. •

EudoreUa truncatula Bate, 1856 I

!1

Figure 2.22

Material examined. Santa Maria Basin, Phase II. Cruise 1-1 (November, 1986), Sta. R-7 (7); l

Cruise 1-2 (January, 1987), R-7 (1); Cruise 1-3 (May, 1987), R-7 (2), R-9 (40); Cruise 2-3 (October, 1987),

I

R-7 (1), R-9 (22); Cruise 2-4 (February, 1988), R-7 (3), R-9 (60); Cruise 3-1 (October, 1988), R-7 (4), R-9 I

(35); Cruise 3-4 (May, 1989), R-6 (2), R-7 (1), R-9 (58). I

156 I
I Figure 2.21. Eudorella redacticruris Wafting and McCann, new species.Female: A, carapaceand pereonites,

lateral view; B, antenna 1; C, pereopod 2; D, uropods. Male: E, pereonite 5 with penial lobes and

pleonites 1 and 2with appendages;F, ventralview of penial lobes. Scale, A = 1 ram, B-E =0.5 nun,

F=0.lmm.

I Description. Adult size, female and male, 5 mm. Anteimule peduncle articles 2 and 3 subequal in

length; main flagellum basal article with setae on outer margin; accessory flagellum three-fourths length of

I main flagellum basal article. Pereopod 1 article 5 shorter than article 6. Pereopod 2 articles 4 and 5 subequal in length. Uropod exopod longer than endopod basal article; endopod terminal seta fused to distal article.

Remarks. This is a widespread northern cold-water species which seems to exhibit a significant

I amou Mediterranean nt of variation. At least four species have been described from the eastern North Atlantic and which may be variants ofE. truncatula. This species has hitherto not been recorded from the

U.S. west coast; however, since it is known to occur in boreal waters, it is possible that it will be found at

I outer shelf depths from California to Alaska.

i 157
Figure 2.22. Eudorella rruncatulaBate, 1856. Female: A, carapace and pereonites 1-4, lateral view; B, body, I

lateral view; C, D, uropods. Scale, A, B = 1 mm,scale C = 0.1 ram, D = 0.5 mm.

I

Type Locality and Type Specimen. British Isles; holotype in Natural History Museum, London. I

I

Distribution. Boreal N.E. and N.W. Atlantic, Mediterranean, and boreal N.E. Pacific, shallow

shelf to deep bathyal. Record from New Zealand is doubtful. I

Genus Eudorellopsis Sars, 1883

Type Species. Leucon deformis KrCyer, 1846. I

Diagnosis (from Watling, 1991a).Pseudorostrum directed dorsally in both male and female; antenna

1 geniculate between peduncle articles 1 and 2; male antenna 2 with brush of setae on anterior margin of I

peduncle articles 4 and 5; male antenna 2 flagellum extending well along pleon; female with exopods on

pereopods 1-3; male with exopods on pereopods 1-4; pereopod 2 article 3 lost; uropod endopod peduncle 2-

articulate; male with 2 pairs of pleopods. I

I Remarks. Species in this genus have been found, with one exception (New Zealand), in the North

Atlantic and North Pacific.

I

158 I
!

I EudoreUopsis longirostris Given, 1961

Figure 2.23

I Material E_ummed. Santa Maria Basin, Phase II. Cruise 1-1 (November, 1986), Sta. R-4 (6);

Cruise 1-2 (January, 1987), R-4 (2); Cruise 1-3 (May, 1989), R-4 (14); Cruise 2-3 (October, 1987), R-4 (4);

Cruise 2-4 (February, 1988), R-4 (4), R-5 (1); Cruise 2-5 (May, 1988), R-4 (12), R-5 (3); Cruise 3-1 (October, 1988), R-4 (5), R-5 (2); Cruise 3-4 (May, 1989), R-4 (17), R-5 (2).

Description. Size of adult female, 4 mm. Carapace with 2 oblique, curving, lateral ridges, uppermost

I terminating anteriorly in small horn; pseudorostral lobes with covering of fine setae; anteroventral margin

finely serrate. Uropod endopod nearly as long as exopod; endopod basal article strongly armed with medial

setae; distal article about one-third length of proximal article, terminal seta slender, not approaching fusion

I with distal article.

Remarks. The only other species with anterolateral horns, E. ushakovi Lomakina, also has one or

i more dorsomedial Type Localhorns ity anond Ttheypecarapa Specicme e, nsand. Rthe/V anterove Velero/VnSta. tral margin 5828-58,of10themiles carapa231.5 ce i°strue coarsely from Ventura serrate.

Pier light, depth 186 m; holotype AHF 584 (now in Los Angeles County Natural History Museum).

I Distribution. Southern California, in sandy sediments, 43-183 m.

I

!

1

!

!

!

!

!

!

Figure 2,23. Eudorellopsislongirostris Given, 1961.Subadultmale: A, carapaceandpereonites,lateral view; B,

I body,lateralview; C, carapace, dorsal view.Subadultfemale:D, body,lateral view.Scale A,B, D =

1 ram, C = 0.1 mm.

i 159
!

Family Nannastacidae I

Diagnosis (emended from Jones, 1963). Mandibles naviculoid with columnar molar or truncate II

with styliform molar. Maxilla 1 palp with 1 or 2 setae. Exopods present on maxilliped 3 and pereopods 1-4, !1

occasionally absent from pereopods 3 and/or 4, in male, and in female on maxilliped 3 and pereopods 1and

2, occasionally absent from maxilliped 3 while occurring on pereopods 1 and 2, or absent from all appendages. •

Pleopods absent in male. Telson absent. Uropod endopod uniarticulate. II

Remarks. The Nannastacidae are often difficult to distinguish from specific members of the

Bodotriidae, especially those where the male is unknown, or where, as in some members of the Mancocuminae, I1

the male lacks pleopods. In the Santa Maria Basin region, the family is represented only by the genera

Campylaspis, Procampylaspis, and Cumella, all of which have a characteristic carapace shape, and so are

easily recognizable. I

Key to the Nannastaeidae of the Santa Maria Basin I

1A. Carapace basically smooth, may be covered with fine sediment, lateral sulcus may be present ... 2

II

1B. Carapace with tubercles, ridges, spines covering large areas of the carapace or otherwise conspicuous i

......................................................................................................................................................... 6

2A. Carapace covered with fine detritus, 3 or 4 spines occasionally visible, maxilliped 2 dactyl with I

recurved teeth .............................................................................................. Procamylaspis caenosa

2B. Carapace features visible, not covered excessively with detritus ................................................... 3 !

II

3A. Pereonites 4 and 5, pleonites 1-3 with dorsolateral spikes ................................. Campylaspis blakei II

3B. Pereonites and pleonites without dorsolateral spikes ...................................................................... 4

4A. Carapace without lateral sulcus ............................................................................. Campylaspis rufa I

4B. Carapace with lateral sulcus ............................................................................................................ 5

m

5A. Carapace lateral sulcus narrow; uropod exopod without setae on outside margin ........................... I

.................................................................................................................. Campylaspis canaliculata lm

5B. Carapace lateral sulcus wide; uropod exopod with setae on outside margin Campylaspis biplicata !

6A. Carapace with conspicuous middorsal ridge or spines ................................................................... 7 I_

6B. Carapace ornamentation primarily dorsolateral .............................................................................. 8

7A. Carapace with middorsal row of large curved spines ....................................... Cumella californica !

7B. Carapace with middorsal ridge ................................................................................ Cumella morion

! 8A. Carapace with lateral ridges, without tubercles ................................................ Campylaspis hartae

8B. Carapace with tubercles or tuberculate ridges ................................................................................ 9 I

160 I
!

I 9A. Uropod peduncle and endopod with many evenly-spaced medial setae ...........................................

.............................................................................................................. Campylaspis rubromaculata

I 9B. Uropod peduncle devoid of medial setae, endopod with 1-3 medial setae .......................................

............................................................................................................ Campylaspis maculinodulosa

I Campylaspis Sars,

Genus 1865

Type Species. Cuma rubicunda Lilljeborg, 1855

I Diagnosis (after Jones, 1974). Carapace strongly elevated posteriorly in females, often coveting

anterior pereonites; anteroventral corner little or not produced; ocular lobe single, middorsal, often

I rudimentary. Mandible molar styliform. Maxilla reduced to simple plate without movable endites. Antenna

i peduncle article 2 without process, subequal to or longer than article 3. First maxilliped reduced, with only

three articles, terminal article very small. Second maxilliped with propodus articulated at near right-angle to

carpus and ending in broad seta, dactyl short and ending in 2 or more distal diverging spines. Female

i maxilliped 3, pereopods 1 and 2 with exopods; male maxilliped 3 and pereopods 1-4 with exopods. Pereopod

1 with ischium not specially elongated. Uropod peduncle usually as tong as or longer than pleonite 6;

exopod basal article short, not submerged in peduncle; exopod longer than its terminal seta.

i Remarks. The genus Campylaspis belongs to a group of genera (including Procamylaspis,

Camylaspides, Paracampylaspis) within the Nannastacidae characterized by the carapace being elevated

i posterodorsally. appendages. ForInexample, this groupintheProcampyIaspis primary generic , characters the dactyl areof associ maxilliped ated with2 isheaclawed, d or anterior rake-like, thoraciinc

Campylaspides, it is in the form of a trident, and in Paracampylaspis, while the dactyl of maxilliped 2 is

i highly reduced, the carpus of maxilliped 3 is greatly expanded.

Campylaspis canaliculata Zimmer, 1936

I Figure 2.24

I R-4 Material Examined. Santa Mafia Basin, Phase II. Cruise 1-2 (January, 1987), Sta. R- 1(2), R-3 (1), (1), R-6 (1), R-8 (2), R-9 (1); Cruise 1-3 (May, 1987), R-1 (1); Cruise 2-3 (October, 1987), R-1 (1), R3 (1), R-4 (1), R-9 (2); Cruise 2-4 (February, 1988), R-1 (4), R-4 (5), R-8 (1), PJ-1 (2); Cruise 2-5 (May,

1988), R-1 (1), R-4 (1), R-5 (1); Cruise 3-1 (October, 1988), R-1 (4), R-4 (1), R-8 (2), R-9 (1); Cruise 3-4

I (May, 1989), R-4 (5), R-9 (2).

Description. Female, 4 ram. Carapace smoothly rounded, elevated posteriorly, with small, lateral,

I udorsal pwrdview, ly curving carapasulc ce umoderately s extending pointed from ventral anteriorly; marginpseudorostral of pseudorostral lobeslobeof tnormal o middlshort e of carapace length; eyelobe side; in

with 3 lenses. Pereonites 1 and 2 with mid-dorsal projections (lappets) which fit into posteromedian depression

I onlengthcaraofpace. endopod, Pereopoandd 2slightly article longer 7 as lonthang aslast artic2 lpleortites, e 5, not darmed istally medially tapering. with Uropod2-7 pedunc heavy lsetae; e aboendopod ut twice

with 2-7 heavy setae; exopod shorter and more slender than endopod, with single subterminal median seta.

I Male, 4 mm. Carapace more elongate and less arched than that of female; lenses of eyelobe enlarged; lateral sulcus not present. Pereonites 1 and 2 with middorsal projections; pereonite 1 mostly obscured by dorsal

margin of carapace; remaining pereonites and pleonites with paired dorsolateral ridges. Uropod peduncles

i much longer than those of female, about as tong as last 4 pleonites; medial margin of endopod and peduncle

armed continuously with finely serrate setae; exopod as in female.

Remarks. Previously known from 2 specimens, both females. This is the only smooth carapace

I form in this region with a narrow lateral sulcus.

I 161
I

I

I

I

I

C \H I

_ _-_ .

, I

,_._._._ I

/ ,! |

I

!

Figure 2.24. Campylaspis body, lateral view; canaliculata H, umpods. 7__Jmmer, Female: 1936.C,Male: body, A,lateral carapace view;andD, pereonites maxilliped 13;-2,E,lapereopod teral view;1;B, F I

pereopod 2; G, uropods. Scale A, B = 1 mm, G, H = 0.5 mm (C-F from Zimmer, 1936; no scale

provided). I

Type Locality and Type Specimens. Between Corona Del Mar and Balboa, California, holotype I

USNM 7148.

Distribution. Southern California including San Diego and southern Oregonian Provinces, 14-410 m. I

162 I
!

I Campylaspis rufa Hart, 1930

Figure 2.25

I Material Examined. Santa Maria Basin region, Phase II. Cruise 1-1 (November, 1986), Sta. R-3

(2); Cruise 1-2 (January, 1987), R-7 (4).

I Description. Female, 3.5 mm. Carapace large, smooth, comprising more than half total length of

body, extending dorsally over pereonites 1-3; antennal notch obsolescent; eyelobe broadened, without lenses.

i Pereopod 2 articles 5 and 7 subequal in length. Uropod peduncles twice length of endopod and approximately equal in length to last 2 pleonites combined; medial margin of peduncle armed with large scales giving

coarsely serrate appearance; endopod with about 7 medial setae. Color dark reddish-brown to orange, with

large dark red chromatophores scattered over body.

I Remarks. The very small adult body size and smooth, reddish brown carapace serves to distinguish

this species from all others in the Santa Maria Basin region. CampyIaspis rufa has never been found in even

i moderateType abundance; Localityitsanoccurrence d Type Speci heremextends ens. Mitlenatch, its range throughout Vancouver theIsland, Oregonian BritishProvince. Columbia, 200 m.;

type in British Columbia Provincial Museum.

I Distribution. Vancouver Island to Point Conception, 200 - 565 m.

!

!

!

!

!

!

!

!

!

Figure 2.25. Campylaspis tufa Hart, 1930. Female: A, carapace and pereonites, lateral view; B, body,lateral

I view; C, carapace, dorsal view; D, uropods. Scale A-C = lmm, D = 0.1 ram.

I 163
!

!

Campylaspis rubromaculata Lie, 1971 II

Figure 2.26

I

Campylaspis nodMosa Lie, 1969 (non Sars, 1887)

Material Examined. Santa Maria Basin, Phase II, Cruise 1-1 (November, 1986), Sta. R-4 (1), R-5 U

(14), R-7 (8), PJ-1 (15); Cruise 1-2 (January, 1987), R-1 (3), R-2 (1), R-4 (1), R-5 (4), PJ-1 (10); Cruise 1- a

3 (May, 1987), R-1 (3), R-8 (3), PJ-1 (9); Cruise 2-3 (October, 1987), R-5 (3), PJ-1 (6); Cruise 2-4 (January,

1988), R-1 (2), R-8 (2), PJ-1 (16); Cruise 2-5 (May, 1988), R-1 (1), PJ-1 (2); Cruise 3-1 (October, 1988), PJ- n

1 (1); cruise 3-4 (May, 1989), R-1 (3), R-8 (1), PJ-1 (9).

Description. Female, 4 mm. Carapace elevated posteriorly, covered with low tubercles in middorsal ii

region, laterally forming dorsal border to sulcus, with single ridge of low tubercles forming ventral border 1

to sulcus and short row of 3 tubercles extending into sulcus dorsally; pseudorostml lobes without tubercles u

dorsaIly, slightly elongate; eyelobe enlarged, without noticeable lenses. Pereonites 1 and 2 with middorsal I

projections, pereonite 1 visible in lateral view; pereonites 3-5 with strong ridge along posterior border. II

Pereopod 2 article 7 nearly 1.5 times length article 5; articles 4 and 5 armed with plumose setae. Uropod i

peduncle as long as last 2 pleonites combined and more than twice length of endopod; latter terminally

quadrate, armed with 3-5 setae medially and 1 short and 1long setae terminally; exopod shorter than endopod, 1

with single subterminal seta medially. Color in life, orange with red pigment chromatophores. I1

Remarks. This species was originally described by Lie (1969), but the proposed name, Campylaspis II

nodulosa, was occupied as Sars (1887) had described a species from the North Atlantic with that name. 1

CampyIaspis rubromaculata bears considerable resemblance to C.sagamiensis Gain6 and C. maculinodulosa

Watling and McCann, which is described below. See remarks under that species for characters which can be m

used for their separation. II

U

Type Locality and Type Specimens. Puget Sound, Seattle, Washington, Holotype female, USNM

125081, 23 m.

Distribution. Southern California to Washington, 22-565 meters. D

l

Campylaspis maculinodulosa Wafting and MeCann, new species 1i

Figure 2.27

I

Material Examined. Santa Maria Basin, Phase II, Cruise t-1 (November, 1986), Sta. R-1 (19), R- I

4 (5), PJ-I (5); Cruise 1-2 (January, 1987), R-1 (3), R-4 (4), R-8 (2), PJ-1 (2); Cruise 1-3(May, 1987), R-1

(16), R-4 (1), R-5 (1), R-8 (3), PJ-1 (2); Cruise 2-3 (October, 1987), R-1 (12), R-4 (4), R-5 (2), R-8 (4), PJ- •

1 (2); Cruise 2-4 (January, 1988), R-1 (3), R-4 (3); Cruise 2-5 (May, 1988), R-1 (20), R-4 (6), R-8 (t6);

Cruise 3-1 (October, 1988), R-1 (12), R-4 (5), R-8 (2); Cruise 3-4 (May, 1989), R-1 (24), R-8 (11).

Description. Adult size, female and male, 2.0 ram. Carapace slightly elevated posteriorly, with II

coveting of non-contiguous low tubercles dorsally; latemt sulcus bordered with lightly tuberculate ridge

dorsally and row of widely spaced tubercles ventrally; pseudorostral lobes longer than normal for the genus,

with row of 3 tubercles extending anteriorly from lateral carapace ridge; eyelobe distinct, without noticeable I_

lenses. Pereonites 1 and 2 with middorsal projections; pereonites 3 and 4 with low dorsolateral tubercles. I

Pereopod 2 article 3 present; article 7 only slightly longer than article 5. Uropod peduncle as long as last 2

pleonites combined and nearly twice length of endopod; rami subequal in length; peduncle unarmed medially i

and endopod with 1 or 2 medial setae. Color in life translucent white with spots on tubercles. Male carapace U

lower and more elongate, only 2 lateral tubercles at posterior end of sulcus; uropod longer than in female,

with about 10 evenly spaced setae on peduncle medial margin, endopod with 4-5 medial setae. I

,64 |
i

!

!

!

!

!

") D

/

/

L b4

!

Figure 2.26. Campylaspis rubromaculata Lie, 1971. Subadult female: A, body, lateral view; B, carapace and

anterior pereonites, lateral view; C, carapace, dorsal view; E, uropod. Scale, A-C, 1.0mm, D = 0.5

I mm

!

Remarks. This species is very similar in general form to C. rubromaculata, but differs from the

latter in the presence of tubercles rather than a short ridge at the posterior end of the carapace lateral sulcus,

I and in the lack of setae the medial margin of the female uropod peduncle. Campylaspis sagamiensis

on

Gam6 exhibits a very similar arrangement of tubercles on the carapace, but differs from C. maculinodulosa

in having the tubercles more widely separated such that a continuous ridge is not formed above the sulcus,

I and in its weakly setose male uropod peduncle.

Type Locality and Type Specimens. California, Santa Maria Basin, off Point San Luis, Sta.R-1

35°05.83'N, 120°49.16%V, 91 m, holotype USNM 273575, paratypes USNM 273567 and LACM 95-74.1.

I Etymology. from L. maculosus, = spotted, and L., nodulus, =knob, referring to the chromatophores

that are associated primarily with the tubercles on the carapace.

i Distribution. Known only from the Santa Maria Basin region, 90-154 m.

!

!

• ! 165

|

I Campylaspis hartae Lie, 1969

Figure 2.28

I Material Examined. Santa Maria Basin, Phase II, Cruise 1-2 (January, 1987), Sta. R-5 (3), PJ-1

(3); Cruise 1-3 (May, 1987), R-1 (7), R-5 (8); Cruise 2-3 (October, 1987), R-5 (17), R-8 (4); Cruise 2-4

I (January, 1988), R-5 (5); Cruise 2-5 (May, 1988), R-5 (8); Cruise 3-1 (October, 1988), R-4 (4), R-5 (4), R- 8 (1); Cruise 3-4 (May, 1988), R-4 (2), R-5 (3).

Description. Adult female, 3.5 mm. Carapace strongly elevated posteriorly, with several series of

I strong ridges bordering deep sulci, 3 of which transverse, with first crossing frontal lobe and joining lateral

ventral ridges anteriorly, second passing obliquely posteriorly to middorsal region, and posteriormost running

from ventrolateral ridge to middorsal ridge, 2 of which are horizontal and border lateral sulcus. Eyelobe

I enlarged but without distinct lenses. Pereonite 1only with middorsal projection. Pereopod 2 article 3

strong

distinct, article 7 short and wide but longer than article 5, armed with several lateral setae. Uropod peduncle

nearly equal in length to last 2 pleonites and 2 to 2.5 times length of endopod; peduncle unarmed medially,

I but heavily scaled, endopod with 3 medial setae. Immature male similar to female; uropod endopod

very

with more medial setae, but peduncle remains without medial setae.

!

!

!

!

!

/ \D

/// I

/ / \

I

i Figure 2.28. Campylaspis hartae I_de,1969.Female: A, carapace and pereonites, lateral view; B, body, lateral

view; C, carapace, dorsal view; D, uropods. Scale, A-C = 1 mm, D = 0.5 ram.

!

i 167
I

Remarks. This species can be distinguished from all others in this region by the very deep depressions II

between the ridges combined with the absence of tubercles on the carapace. Of the 69 specimens examined

in this study only a single individual was a male. Lie did not record a mate from the type locality. While •

mature males of campylaspids are always rare, they are usually encountered at some time during the year. |

Type Locality and Type Specimens. Puget Sound, off Seattle, 22-23 m, sediment >80% sand,

mean particle size 0.113-0.235 mm. Holotype, USNM 125077. i

Distribution. Probably distributed throughout San Diego and the Oregonian Provinces, a species of

the shallow shelf, known thus far from depths of 22-154 m.

CampyIaspis blakei Watling and McCann, new species i

Figure 2.29 I

Material Examined. Santa Maria Basin, Phase II, Cruise 1-1 (November, 1986), Sta. R-4 (1), R-5

(6); Cruise 1-2 (January, 1987), R-2 (1), PJ-1 (2); Cruise 1-3 (May, 1987), R-5 (5); Cruise 2-3 (October, i

1987), R-2 (1), PJ-1 (3); Cruise 2-4 (January, 1988), PJ-1 (I); Cruise 2-5 (May, 1988), R-2 (1), R-5 (2), PJ1 (1); Cruise 3-1 (October. 1988), R-4 (1), R-5 (1), PJ-I (7); Cruise 3-4 (May, 1989), PJ-1 (2).

Description. Female, 3 mm. Carapace strongly elevated posterodistally, broadly rounded, with i

gently sloping lateral sulcus not bounded by ridges; with paired set of 3 small spines dorsolaterally near

m

posterior margin; eyelobe small, distinct; pseudorostral lobes ordinary. Pereonites 1 and 2 with middorsal i

projections; pereonite 4 with dorsally projecting lateral spines; pereonite 5 with both dorsolateral and lateral i

dorsally projecting spines. Similar spines also on pleonites 1-4. Pereopod 2 article 3 distinct; article 7 as

u

long as article 5. Uropod peduncle as long as last 3 pleonites combined and about 2.5 times length of ,me

endopod; peduncle slightly flared terminally. Endopod and distal half of peduncle armed with 4 to 6 and 4 i

setae respectively. Color in life, white to gray, translucent. Male unknown.

!1

Remarks. This species is unique among members of this genus which have a smooth carapace in i

possessing blunt, dorsally projecting spines on the pereonites and pleonites, i

Type Locality and Type Specimens. California, Santa Maria basin, off Purisima Point, Sta. R-5,

34°42.69'N, 120°50.83'W, 154 m, holotype USNM 273548, paratypes USNM 273549 and LACM 95-73.1. •

Etymology. The species is named in honor of my friend and colleague, the lead scientist of the n

Santa Maria Basin macrobenthic sampling program, and senior editor of this Atlas project, in recognition of

his many contributions to our knowledge of the benthic fauna of the Southern California continental shelf. 11

II Distribution. Known from the Santa Maria Basin area and off Pt. Loma, San Diego, 92-161 m.

i

!

I

!

!

168 I

I

l

Campylaspis biplicata Watling and McCann, new species l

Figure 2.30a, b

Material Examined. Santa Maria Basin Region, Phase II, Cruise 1-1 (November, 1986), Sta. R-6 Ii

(17), PJ-1 (3); Cruise 1-2 (January, 1987), R-1 (1), R-3 (6), R-5 (11), R-6 (1); Cruise 1-3 (May, 1987), R-3

(9), R-5 (7), R-6 (9), R-9 (10), PJ-1 (2); Cruise 2-3 (October, 1987), R-2 (1), R-3 (18), R-5 (4), R-6 (11), R- _l

9 (4), PJ-1 (3); Cruise 2-4 (January, 1988), R-3 (11), R-5 (20), R-6 (I0), R-9 (5), PJ-1 (2); Cruise 2-5 (May, II

1988), R-3 (1), R-5 (14), R-6 (12), R-9 (2), PJ-1 (1); Cruise 3-1 (October, 1988), R-1 (2), R-3 (15), R-5 (20),

R-6 (14), R-9 (7), PJ-1 (3); Cruise 3-4 (May, 1989), R-3 (3), R-4 (1), R-5 (24), R-6 (11), R-9 (3), PJ-1 (8). I

n

Description. Adult size, female and male, 2.0 mm. Carapace extended strongly posteriorly, appearing

flattened dorsally; with lateral, nearly horizontal sulcus becoming wider from posterior to anterior, bounded

above and below by soft ridges ornamented with pigment spots; eyelobe small, pseudorostral lobes slightly 1

enlarged, branchial siphons about twice length of pseudorostrum. Pereonites 1 and 2 with middorsal

w

projections; pleonite 4 pleurae extending back along peronite 5. Pereopod 2 article 3 distinct, article 7

longer than article 5, with terminal seta as long as article in length. Uropod peduncle as tong as last 3 I!

pleonites combined and twice length of endopod, with only a few weak setae; exopod basal article half n

length of distal article; endopod with 3-5 medial setae. Male uropod endopod somewhat more robust than in

female. I

Remarks. The presence of two, three, or four subparaltel lateral ridges on the carapace occurs in

several species, many of which have been found in the deep Atlantic (e.g., C. arcuata Jones, C. plicata m

Jones, C. vaIleculata Jones, C. exarata Jones, C. bicarinata Jones, C. crispa Lomakina, C.johnstoni Hale, |

C. macrophthalma Sars, C. sinuosa Gamr, C. umbensis Gurwitchi, C. undata Sars). The appendage

morphology of C. biplicata is somewhat similar to that seen in C. crispa and C. umbensis, however, both of 1

these species have three ridges that are more vertical (relative to the ventral margin). In the form of the II

ridges C. plicata resembles most closely the Atlantic species of Jones and Sars, but differs from them in the

form of the appendages, especially the uropod. I

Type Locality and Type Specimens. California, Santa Maria Basin, off Point Sal, Sta. R-9, U

34°53.68'N, 120°59.12'W, 410 m, holotype USNM 273546, paratypes USNM 273547 and LACM 95-72.1.

Etymology. from L. bis = two, and L. plicatus = fold, referring to the two lateral ridges on the 1

carapace. U

Distribution. Known from Puget Sound to San Diego, including the Santa Maria Basin Region, 47-

410 m, with most specimens found at 145-410 m. I

Genus Procampylaspis Bonnier, 1896 n

Type Species. Procampylaspis echinata Bonnier, 1896. I_

Diagnosis. Carapace strongly elevated in female, occasionally covering part of first pereonite; •

anterolateral corner not produced in females, weakly produced in males; ocular lobe single, middorsal, II

often rudimentary. Mandible molar elongate, weakly triturative distally. Maxilla with movable endites.

Antenna 1 peduncle article 2 without process, longer than article 3. First maxilliped 6-articulate, dactyl •

minute. Maxilliped 2 dactyl with recurred processes, claw-like. Female maxilliped 3, pereopods 1 and 2 I

with exopods; male maxilliped 3 and pereopods 1-4 with exopods. Pereopod 1article 3 (ischium) subequal

to or longer than article 4. Uropod peduncle usually as long as or longer than pleonite 6; exopod basal article •

short, not submerged in peduncle; exopod longer than its terminal seta. |

Remarks. This genus is distinguished from other members of the family by the shape of the dactyl

on maxilliped 2 and the elongate ischium on pereopod 1. I

170 I

I

I

I

i

,.. I

I

Figure 2.30b. Campylaspis biplicata Wafting and McCann, new species.Female: F, maxilliped3; G, pereopod

2; H, uropocLMale: I, uropod. Scale H, I = 0.5 mm. I

|

Procampylaspis caenosa Watling and McCann, new species I

Figure 2.31 I

[]

Material Examined. Santa Maria Basin Region, Phase II, Cruise 1-1 (November, 1986), Sta. R-1

(3), R-2 (3), PJ-1 (8); Cruise 1-2 (January, 1987), R-1 (6), R-2 (4), R-4 (5), R-5 (7), PJ-1 (3); Cruise 1-3 n

May, 1987), R-1 (6), R-2 (3), R-4 (1), R-5 (8), PJ-1 (9); Cruise 2-3 (October, 1987), R-1 (6), R-2 (6), R-4 I

(11), R-8 (6), PJ-1 (6); Cruise 2-4 (January, 1988), R-1 (4), R-4 (15), R-5 (3), R-8 (2), PJ-1 (1); Cruise 2-5

(May, 1988), R-1 (5), R-4 (10), R-5 (3), R-8 (4); Cruise 3-1 (October, 1988), R-1 (6), R-4 (3), R-5 (1), R-8 am

(2), PJ-1 (6); Cruise 3-4 (May, 1989), R-1 (3), R-4 (10), R-8 (3), PJ-1 (4). 1

Description. Female, 3.0 mm. Carapace slightly elevated dorsally; covered with dense coating of

sediment through which shallow lateral sulcus visible, bordered above and below by low ridges, upper of •

which may bear 3-5 large conical spines; pseudorostral lobes enlarged, directed somewhat upward; eyelobe I

not distinct. Pereonites 1 and 2 without middorsal projections. Pleonites I-5 with paired dorsolateral spines.

N

172 i
!

I

I

I

I

I

i D '..

' '1E

! i

I "_ ".

/I_,. / \

"_ ".', )' i

_ '":"7' _,

f\ i_jIl<,','_'_,, y/

I _lk<j ,. _ . :_. ,:: : .:. " ,.x. " •,. • _, " e,. : ..

Figure 2.31. Procampylaspis caenosa Watling and McCann, new species, Female: A, Carapace and anterior pereonites, lateral view; B, body, lateral view; C, carapace, dorsal view; D, maxilliped 2; E, uropods.

Scale, A-C = 1 nun, E = 0.5 nun.

I

i Maxilliped 2, articles 6 and 7 subequal in length; terminal article with 4 large recurved claw-like teeth, and 1 smaller tooth nestled between penultimate and last claw. Uropod peduncle 1.5 times length of endopod;

medial margin of peduncle unarmed, of endopod with 2 setae; exopod with very long terminal seta, extending

to end of endopod terminal seta. Male, similar to female but carapace and uropod peduncles more elongate.

I Remarks. The combination of conical carapace spines and broad lateral sulcus, along with the

details of the maxilliped 2 dactyl, help to distinguish this sediment-covered species. Of the 26 known species

i of Procampylas Type pisLocality , onlyand5 haTveypebeeSpecim n found at shallow shelf depths. ens. California, Santa Maria Basin, off Purisima Point, Sta. R-4,

34°43.01'N, 120°47.39'W, 92 m, holotype USNM 273542, paratype USNM 273543 and LACM 95-71.2.

I Distribution. Known from the Santa Mafia Basin region and off Pt. Loma, San Diego, 47-161 m.

Etymology. from L. caenosus, = muddy, dirty, refen'ing to the covering of sediment particles on

the carapace surface. !

I

i 173
!

1

Genus Cumella Sars, 1865 n

Type Species. Cumella pygmaea Sars, 1865 1 

Diagnosis (from Watling, 1991b). Carapace anteroventral comer acute to slightly rounded, not

strongly projecting; ocular lobe single, middorsal; siphons united medially, occasionally slightly separated;

pseudorostral lobes of varying length, usually slightly to strongly upturned, meeting in front of eyelobe, l

Mandible molar columnar. Maxilla with movable endites. Antenna 1 peduncle article 2 with or without

u

process, subequal to or longer than article 3. Maxilliped 1 of normal length. Female maxilliped 3, pereopods 1

1 and 2 with exopods; male maxilliped 3 and pereopods 1-4 with exopods. Uropod peduncle usually as long 1

as or longer than pleonite 6; exopod basal article short, not submerged in peduncle; exopod much longer w

than its terminal seta.

1

Remarks. Following the lead of Bacescu (1971) who erected the f'trst subgenus in this very large 1

genus, Watling (1991b) added three additional subgenera in an attempt to define some pattern among the

species. For diagnoses of the subgenera, see Watling (1991b). 1

CumeUa (Cumella) morion Watling and McCann, new species

Figure 2.32 I

Material Examined. Santa Maria Basin Region, Phase IT,Cruise 1-1 (November, 1986), Sta. R-4 •

(17); Cruise 1-2 (January, 1987), R-1 (8), R-4 (5), R-8 (11); Cruise 1-3 (May, 1987), R-1 (13), R-4 (39), R- l

8 (4); Cruise 2-3 (October, 1987), R-1 (1), R-4 (24), R-8 (14); Cruise 2-4 (January, 1988), R-1 (4), R-4 (21),

R-5 (1), R-8 (8), PJ-1 (I); Cruise 2-5 (May, 1988), R-1 (2), R-4 (107), R-8 (5); Cruise 3-1 (October, 1988), n

R-1 (4), R-4 (31), R-8 (16); Cruise 3-4 (May, 1989), R-1 (1), R-4 (110), R-8 (4). 1

Description. Female, 1.5 ram. Carapace with flared ridge along posterodorsal margin, dorsally

joining large middorsal crest which originates on frontal lobe; eyelobe without obvious lenses; pseudorostral •

lobe with short, curved horizontal ridge extending onto carapace frontal lobe. Pereonites 1 and 2 with I

thickened posterior margin; pereonite 3 with slight middorsal process. Body compressed laterally. Pleonites

1 -4 with dorsally-directed middorsal processes. Uropod peduncle and endopod subequal in length, about as •

long as last 2 pleonites combined; endopod with 2 medial setae and 2 terminal setae. 1

Remarks. This species is quite similar in carapace shape and appendage morphology to C. (C.)

rigida Gam6 and C. (C.) sadoensis Gamr, but differs from both in not possessing a dorsolateral vertical •

ridge midway along the carapace and in bearing vertical processes on pleonites 1 - 3. I

Type Locality and Type Specimens. California, Santa Maria Basin, off Purisima Point, Sta. R-4,

34°43.01'N, 120°47.39'W, 92 m, hototype USNM 273544, paratype USNM 273545 and LACM 95-71.1. n

Etymology. A morion is a visor-less helmet with a high crest worn by some 16th century soldiers,

for example, the Spanish conquistadors who frequented the Californian shores, and to which the carapace of 1

this species bears some resemblance, i

u

Distribution. Known only from the Santa Maria Basin region, most abundant at the 90 m stations,

but occasionally found to 154 m. u

1

!

1

174 I

i

I
I

I

I

I

I

I ,,\ J

I i I

I , I i,

I ' I

:',"",, _ I ,'",

,,,,,,,,,,,__! ,',,

i { ,,,,,

Figure 2.32. CumeUa (CumelIa) morion Watling and McCann, new species. Female: A, carapace, pereonites,

I and anterior pleonites, lateral view; B, body, lateral view; C, anterodorsal view; D, uropods.

carapace,

Scale, A,C, D = 0.2 ram, B = 1 mm.

I

CumeUa (CumeUa) californica Wafting and McCann, new species

i Figure 2.33

Material Examined. Santa Maria Basin Region, Phase II, Cruise 1-1 (November, 1986), Sta. R-4

i (2), PJ-1 (1); Cruise 1-2 (January, 1987), R-4 (4), R-5 (1), R-8 (2), PJ-1 (1); Cruise 1-3 (May, 1989), R-4 (1),

R-5 (5), R-8 (1), PJ-t (6); Cruise 2-3 (October, 1987), R-4 (24), R-5 (3), R-8 (4), PJ-1 (1); Cruise 2-4

(January, 1988), R-4 (7), R-5 (1), R-8 (6), PJ-1 (4); Cruise 2-5 (May, 1988), R-4 (2), R-5 (1), R-8 (13), PJI 1 (2); Cruise 3-1 (October, 1988), R-4 (9), R-5 (2), R-8 (20), PJ-1 (3); Cruise 3-4 (May, 1989), R-4 (23), R- 8 (3), PJ-1 (15).

Description. Female, 2 ram. Carapace with swollen ridge parallel to posterior margin, giving

I appearance elevated submarginal region dorsally; 4-5 spines in middorsal row beginning on frontal lobe;

of

eyelobe small; pseudorostral lobes directed forward, subacute. Pereonites with flared lateral margins,

becoming nearly horizontal. Uropod peduncle as long as last pleonite; endopod as long as peduncle, with

I elongate, partially seta; exopod as long as

fused terminal distal article terminal seta.

I

i

i I 175
I

i

!

!

!

i/ ""' i

i

'_--_ /-4 ,.,,b-!I,

r , ,_,_ ,,, I

/ I ; i '1

I " '1 i "..'.,

L il' ", _k _'l'_ •

I

I

Figure 2.33. Cumelk_ (CumeUa) eelifornica Watl_ and MeCm, new species. Female: A, carapace and I

pereonites, dorsolateral view; B, pereonites and pereopods 1-3;C, uropods. Scale,A = 1ram, B,C = •

0.2 ram. U

Remarks. In addition to the new species described here, three CumeIla (CumelIa) species are known I

to have coarse spines along the middorsal crest of the carapace, viz., C. (C.) arguta Gam6, C. (C.) meridionalis

Jones, and C. (C.) quadrispinosa Gam6. None have the broadly flaring pleurites nor the short uropod peduncles

seen in C. (C.) californica.

Type Locality and Type Specimens. California, Santa Maria Basin, off Purisima Point, Sta. R-4,

34°43.01'N, 120°47.39'W, 92 m, holotype USNM 273540, paratype USNM 273541 and LACM 95-70.1. I

m

Distribution. Known from the Santa Maria Basin Region and off Pt. Loma, San Diego, 45-154 m.

Acknowledgements I

We would like to express our appreciation to several people who helped with the production of this •

paper. Most of the line drawings in the species descriptions are the wonderful handiwork of Patrice Rossi, |

with a few being contributed by B. McKenzie. The original introductory figures are the work of C. Chisholm.

Very able assistance with the SEM was always kindly given by K. Edwards. Additional distribution records •

were made available by Tim Stebbins of the City of San Diego Marine Biology Lab, and the participants of l

a cumacean workshop held at Friday Harbor Laboratories in February, 1995. This project was supported by

the U.S. Department of the Interior, Minerals Management Service, under Contract No. 14-35-0001-30484. !

176 I
|

I Literature Cited

I Bacescu-Mester, L. 1967. Contribution to the knowledge of the genus Leptostylis Sars (Cumacea): three

new species collected by the Vema Expedition. Crnstaceana 13: 265-274.

I Bate, S. 1856. On the British Diastylidae. The Annals and Magazine of Natural History 102: 449-465. Caiman, W.T. 1912. The Crustacea of the Order Cumacea in the collection of the United States National

Museum. Proceedings of the United States National Museum 41:604-674.

I Day, J. 1978. Southern African Cumacea. Part 3. Families Lampropidae and Ceratocumatidae. Annals of

the South African Museum 76: 137-189.

I Day, J. 1980. South Southern African Museum African Cumacea. 82: 187-292. Part 4. Families Gynodiastylidae and Diastylidae. Annals of the

i Gamr, S.Marine I962. Biological On the cumacean LaboratoryCrustacea from Tanabe Bay, Kii Peninsula. Publications of the Seto 10:154-210.

Given, R.R. 1961. The cumacean fauna of the Southern California continental shelf. No. 1,Family Leuconidae.

I Bulletin of the Southern California Academy of Sciences 60: 129-146.

Given, R.R. 1964. The cumacean fauna of the southern California continental shelf. No. 2. The new family

i Mesolampropidae. Crustaceana 7: 284-292. Gladfelter, W.B. 1975. Quantitative distribution of shallow-water Cumacea from the vicinity of Dillon

Beach, California, with description of five new species. Crustaceana 29:241-251.

I Harada, I. 1959. Cumacean fauna of Japan. I. Family Lampropidae. Japanese Journal of Zoology 12: 229-

246.

I Hart, J.F.L. 1930. Some Cumacea of the Vancouver Island Region. Contributions to Canadian Biology and Fisheries, n.s., 6: 25-40.

i Jones, N.S. Oceanographic 1963. The marine fauna of New Zealand: Crustaceans of the Order Cumacea. New Zealand Institute Memoir 23: 1-81.

Jones, N.S. 1969. The systematics and distribution of Cumacea from depths exceeding 200 meters. Galathea

I Report. 180. Scientific results of the Danish Deep-Sea Expedition round the World 1950-52 10: 100-

i Kr0yer, KrCyer, H. H. 1841. Fire nye arter af Slaegten Cuma. Naturhistorisk Tidsskrift 3: 503-534. 1846. Carcinologiske bidrag. Naturhistorisk Tidsskfift, Ser. 2, 2:123-211.

i Lie, U. 1969. Cumacea from Puget Sound and off the northwestern coast of Washington, with descriptions of two new species. Crustaceana 17: 19-30.

Lie, U. 1971. Additional Cumacea from Washington, U.S.A., with description of a new species. Cmstaceana

I 21: 33-36.

Norman, A.M. 1867. On the Crnstacea, Echinodermata, Polypora, Actinozoa and Hydrozoa. Report of the

i committee appointed for the purpose of exploring the coasts of the Hebrides by means of the dredge. Report of the British Association for the Advancement of Science 36: 193-206.

Sars, G.O. 1869. Unders/igelser over Christianaf-yord dybvandsfauna. Nyt Magazin for Naturvidenskabeme

I 11: 1-58.

i 177
!

Sars, G.O. 1878, 1879 Nye Bidrag til Kundskaben om Middelhavets Invertebraffauna. II. Middelhavets •

Cumaceer. Separataftryk af Archiv for Mathematik ig Naturvidenskab 3:46I-512; 4: 1-144.

Sars, G.O. 1883. Oversigt af Norges Crustaceer med forel_bige bemaerkninger over de nye eller mindre n

bekj ende Arter. 1. F/Srhandlingar VidenskabsSelskabet i Christiana 1882:1-124. J

Sars, G.O. 1887. Report on the Cumacea collected by H.M.S. Challenger during the years 1873-1876. •

Report on the Scientific Results of the Voyage of the H.M.S. Challenger During the Years 1873- II

1876 Under the Command of George S. Nares and Frank Tourle Thomson 19(55): 1-78.

Sars, G.O. 1900. Cumacea. An Account of the Crustacea of Norway 3:1-115. I

Say. T. 1818. An account of the Crustacea of the United States (continued). Journal of the Academy of

Natural Sciences of Philadelphia 1: 313-319. •

Schram, F.R. 1986. Crustacea. New York: Oxford University Press, 606 pp.

Stebbing, T.R.R. 1912. The Sympoda (Part VI of S. A. Crustacea, for the marine investigations in South •

Africa). Annals of the South Africa Museum 10: 129-176. II

Wading, L. 1979. Crustacea: Cumacea. Marine Flora and Fauna of the Northeastern United States. NOAA

Technical Report NMFS Circular 423: 1-22. n

Watling, L. 1983. Peracaridan disunity and its bearing on eumalacostracan phylogeny with a redefinition of

eumalacostracan superorders. Crustacean Issues 1: 213-228. •

Watling, L. 1991a. Revision of the cumacean Family Leuconidae. Journal of Crustacean Biology 11: 569-

582.

n

Wading, L. 1991b. Rediagnosis and revision of some Nannastacidae (Crustacea: Cumacea). Proceedings of I

the Biological Society of Washington 104: 751-757.

Zimmer, C. 1926. Northern and Arctic invertebrates in the collection of the Swedish State Museum I

(Riksmuseum). Kungliga Svenska Vetenskaps-Akademiens Handlingar 3: 1-88. Ill

Zimmer, C. 1936. California Crustacea of the Order Cumacea. Proceedings of the United States National •

Museum 83: 423-439. |

Zimmer, C. 1943. Cumaceen des Stillen Ozeans. Archiv fiir Naturgeschichte 12:130-174.

!

!

!

!

178 I
!

I Attachment I

Phase I and II manuscript names with formal equivalents

I Family Diastylidae Family Nannastacidae

i Diastylis paraspinulosa = D. sentosa new Campylaspis sp. N = manca, probably of C.

species rubromaculata

Diastylis sp. 1 = Diastylis peIlucida Hart, 1930 Campylaspspis sp 6 = CampyIaspis rufa Hart,

I Diastylis sp. 2 =Diastylis sentosa new species

1930

Diastylis sp. 5 =Diastylis santamariensis new Campylaspis sp. 1 = Campylaspis

I species maculinodulosa, new species

DiastyIis sp. A = Diastylis crenellata new Campylaspis sp. 10, Campylaspis sp. 12 =

species* Campylaspis hartae Lie, 1969

I DiastyIis sp. E = Diastylis quadriplicata new Campylaspis sp. E = Campylaspis blakei, new

species species

I Campylaspis sp. B, sp. 7, sp. 8, sp. 9, and

Leptostylis longimana Leptostylis calva new

Campylaspis nr. crispa = Campylaspis

species biplicata, new species

I Leptostylis villosa =Leptostylis ? (specimens too damaged, but are def'mitelynotL. villosa). Campylaspis sp. 4 = Procampylaspis caenosa,

new species

Leptostylis sp. 1 = Leptostylis abditis new

I Cumella sp. i = CumeUa (Cumella) morion,

species

new species

Leptostylis sp. A =Leptostylis calva new species

• Cumellasp. 2 =CumeUa (Cumella)cah'fornica,

I Leptostylis sp. E = Diastylis crenellata new new species

species*

I Family Leucortidae

Hemileucon sp. 1 = Alloeoleucon

santamariensis new species

I Leucon sp. H = Leucon (Diaphonoleucon)

declivis, new species

I Leucon sp. A = Leucon (Leucon)falcicosta, new

species

I Leucon(Crymoleucon (? Epileu)conbishopi ) sp.Bacescu, B = L1988 eucon

Eudorella sp. 2 = Eudorella redacticruris new

I species

Eudorella sp. 1 = Eudorella truncatuIa Bate,

I 1856

*NotethatDiastytiscreneltatais correctlylistedtwiceonthechecklist.

i 179
I

Attachment H i

List of cumaceans previously found in the Oregonian Province

I

Family Diastylidae Family Leuconidae

(Caiman, 1912) as EudoreIla pacifica Hart, 1930 (includes 1

i

Anchicolurus occidentalis

Colurostylis (?) Eudorella tn'dentata Hart, 1930)

Diastylis abbotti Gladfelter, 1975 Eudorellopsis biplicata Caiman, 1912 l

Diastylis californica Zimmer 1936 Eudorellopsis integra Smith, 1879

Diastylis paraspinulosa Zimmer, 1926 EudoreIlopsis longirostris Given, 1961 1

Diastylis pelIucida Hart, 1930 Leucon armatus Given, 1961

Diastylis umatillensis Lie, 1969 Leucon magnadentata Given, 1961 n

Diastylopsis dawsoni Smith, 1880 Leucon subnasica Given, 1961

Leptostylis villosa Sars, 1869 (doubtful record Family Nannastacidae 1

of Hart, 1930)

Campylaspis canaliculata Zimmer, 1936

Family Lampropidae Campylaspis hartae Lie, 1969 i

Hemilamprops californica Zimmer, 1936

CampyIaspis rubromaculata Lie, 1969

Hemilamprops graciIis Hart, 1930 Campylaspis rufa Hart, 1930 I

Lamprops carinata Hart, 1930

Cumella vulgaris Hart, 1930

Lamprops quadriplicata Smith, 1879 •

Family Bodotriidae l

Lamprops serrata Hart, 1930

Cyclaspis nubila Zimmer, 1936

Lamprops tomalesi Gladfelter, 1975 n

Lamprops triserrata Gtadfelter, 1975

Lamprops krasheninnikovi Derzhavin, 1926 1

Lamprops obfuscata (Gladfelter, 1975)

Mesolamprops bispinosa Given, 1964 1

Mesolamprops dillonensis Gladfelter, 1975

!

!

I

I

180 I
I

I 3. TANAIDACEA

I by

I Masahiro Dojiri I and Jiirgen Sieg 2

I Introduction

! ,

The Tanaidacea is an order of free-living and exclusively benthic malacostracans belonging to the

Peracarida, which also includes the orders Mysidacea, Amphipoda, Cumacea, Isopoda, Spelaeogriphacea,

and Mictacea. Presently, the tanaids include three recent snborders (i.e., Apseudomorpha, Neotanaidomorpha,

I and Tanaidomorpha) and one extinct suborder (i.e., Anthracocaridomorpha), 21 families, and about 700

extant species. Most species are exclusively marine, but a few are known to occur in fresh or brackish water.

Species that have been reported from fresh water are typically marine, euryhaline species. There are no

I strictly fresh water Tanaidacea. This order is most closely related to the Isopoda (see Siewing, 1953; Fryer, 1964; Sieg, 1983a), although Watling (1981) considered it a sister taxon to the Cumacea.

I Taxonomic History

I The first tanaidacean described is "Gammarus heteroclitus" recorded Viviani

ever probably by

(1805) from Genoa, Italy. The published illustrations are reminiscent of members of the genus Leptochelia,

but the species cannot be identified with any certainty. For this reason, it was listed as incertae sedis under

I the family Paratanaidae (now Leptocheliidae) by Sieg (1983b). Prior to the discovery of Viviani's work,

"Cancer Gammarus Talpa," now attributed to Apseudes talpa and described by Montagu (1808), was

considered the furst known tanaidacean. In both cases, the incorrect identifications indicate that the authors

I thought that these organisms belonged to the Garnmaridea (Amphipoda). Although Latreille (1831) had

transferred these species to the isopodan group "Heteropa" (= Heteropoda), their supposed amphipod

affiliation could still be found in Gerstaecker (1883).

I These tanaid species then were taxonomically grouped with the Isopoda for quite some time.

Milne-Edwards (1840) thought that the known species should be included in the "Idoteides", while White

( 1847, 1850) placed them in the "Asellita." Dana (1852) introduced the name "Anisopoda" which indicated

I group species represented a very group among isopods, prompted

that this of miscellaneous unusual the This

Bate and Westwood (1868) to place them in the "Tribus Vagantia" of their "Isopoda Aberrantia" which

contained anthuridand gnathiidisopods among others. G. O. Sars (1882) named them "Isopoda chelifera."

I Finally, Claus (1888) elevated the Anisopoda to the ordinal level, a taxonomic position equalto that

of the Isopoda. Hansen (1895) introducedthe name Tanaidacea, which is now the commonly accepted name

i for the order.

I

Environmental Monitoring Division, 12000 Vista det Mar, Playa del Rey, California, 90293, USA

i 2 Universit_t Osnabriick, Standort Vechta, Driverstraf, e 22, 2848 Vechta, Germany

i 181
1

The systematics of some tanaidacean groups is still in a state of flux. Many new species remain to I

be described, and the diversity within this order is only now being recognized. So far about 700 species have

been described, but the estimated number of tanaidacean species calculated from deep-sea samples could •

raise this number well over 3,000 species. I

Tanaidaceans have a relatively well-known fossil record that dates back to the Lower Carboniferous

(Schram, 1974; Sieg, 1983a, 1984a) and shows the first radiation in the Jurassic (Schram et al., 1986). Even •

some Eocene specimens are known (Sieg, unpublished data).

Since G. O. Sars (1882), only two families, Tanaidae and Apseudidae, were recognized until Lang

(1949) published his f'n'st paper on the Tanaidacea; this paper marks the starting point for a more complex, •

but better foundation for the systematics of this order. In the following years, Lang (1956a, 1956b, 1970,

1971, 1973) published a series of papers not only describing a few new families, but also dividing the order

into two suborders Monokonophora and Dikonophora. Subsequently, the classification was further revised l

by Sieg (1973, 1976, 1980a, 1984a) who included the fossil record in his analysis.

Four suborders are presently recognized. The f'u'st suborder, Anthracocaridomorpha, contains

exclusively Carboniferous and Permian species. The second suborder, Apseudomorpha, consists of fossil I

species from the Jurassic (Schram et al., 1986) as well as all the extant taxa formerly united in the U

Monokonophom. The third suborder, Neotanaldomorpha, is a group of exclusively deep-sea species (Gardiner,

1975) which have no known fossil record. The final suborder, Tanaidomorpha, includes one fossil (Jurassic) 1

species and all remaining taxa formerly grouped under the suborder Dikonophora.

External Morphology I

Because the tanaidacean fauna collected for this study belongs predominantly to the suborder •

Tanaidomorpha, much of the general discussion on morphology presented here focuses on this suborder.

Additionally, however, morphological features of the Apseudomorpha, specifically the Apseudidae, that

differ from those of the Tanaidomorpha are discussed. Much of the information provided here is taken from •

the compilation published by Sieg (1984a). II

Body Shape

The general habitus of the tanaidomorphans is characterized by its cylindrical body shape, while 1

B

that of the apseudomorphans is dorsoventrally depressed.

The body (Figure 3.1), ranging in total length from 1 to 37 mm, always consists of three tagmata of •

which the first one, the eephalothorax, is formed by the fusion of the eephalon and the first two l

thoracomeres. A small carapace encloses the branchial chamber. Terminology for the descriptions of

the surfaces of body somites and appendages follow that of Lang (1968): tergal andsternal are synonymous n

with dorsal and ventral, respectively; rostral and caudal are synonyms of anterior and posterior, respectively I

(Figure 3.1).

Eight appendages, seven of which are paired, are found attached to this region: antenna 1(antennule), •

antenna 2 (antenna), labrum (unpaired), mandibles, maxilla 1 (maxillules), maxilla 2 (maxilla), maxilliped Ii

(thoracopod 1), and eheliped (thoracopod 2). The remaining six free thoracomeres are called pereonites (or

pereomeres), each bearing one pair of nniramous walking legs, the pereopods, and collectively forming •

the second tagma referred to as the pereon. Only the first pair of pereopods bear exopodites. The last |

tagma, the abdomen or pleon, is predominantly six-segmented and composed of five pleonites and a

pleotelson. Sometimes the pleonites are reduced in number or totally fused with the pleoteIson. Typically, •

each pleonite bears one pair of blramous pleopods. Pleopods may be present or absent. Usually only the 1

last abdominal somite is fused with the telson, forming the pleotelson. The appendages of the pleotelson,

the uropods, are fitiform or styliform, and uni- or biramous, but may be reduced in some tanaidomorphan •

families.

l

182 I
I

I

I

I

I

I rostral

I

I

I

I

I

I

!

I

I

P.1

i

Figure 3.1. Generalized habitus of a tanaidomorphan"top, dorsal view; bottom, lateral view.

I

I 183
!

Genital openings are found on the eoxae of the fourth pair of pereopods (thoracopod 6) in the i

female. In the male, these openings are centered on a cone located on the sternite of the last pereonite

(thoracomere 8). Some species of tanaidaceans have one slender, elongate "genital cone" having one common II

outlet for both testes ("Monokonophora" sensu Lang), but others have two cones, each with an outlet for !

one testis ("Dikonophora" sensu Lang). Various intermediate stages exist between these two extremes: a

single genital cone can be relatively broad and bear two openings, or the cone may be deeply incised •

resulting in a dicondyl (bilobed) type of genital cone. !

Hooks, referred to as hylmsphaenia, are located on the stemites of some of the pereonites in apseudids

and occasionally in tanaidomorphans. Ridges may occur on the sternal area of the pleonites (e.g., Scoloura). i

Females of some tanaidacean species lack pleopods, while females of other species have a reduced 111

number of pleonal somites.

The reduction in the number of pleonal segments is a more or less common feature within the i

Tanaidae in which the pleon is composed of up to five pleonites of which only three are fully developed.

n

Each of the three distinct pleonites bear one pair of pleopods. The evolutionary fate of the posterior two i

pleonites differs among the various genera of this family. For example, in phylogenetically primitive genera i

of the Tanaidae, the last two pleonites remain separate, but are distinctly smaller and never have pleopods. I

In more derived genera (e.g., Zeuxo) the last two pleonites are represented by two small tergites dorsally,

but by only a single sclerite ventrally. Furthermore, in some genera, one or both of the pleonites may be I

absent, resulting in a four-segmented (e.g., Tanais) or three-segmented (e.g., PancoIus) pleon. []

Typically, the pleonites are as wide asthe pereonites. In contrast, within the subfamily Anarthrurinae, I

the pleonites of the females are usually narrower than the pereonites. This is also the case in some species of i

the Akanthophoreinae.

i

Sexual dimorphism is commonly observed in the general habitus of most tanaidacean groups with •

the exception of the Tanaidae. Males often look quite different from the females. For example, the "swimming |

males", found in such families as the Paratanaidae, Anarthruridae, Typhlotanaidae, Pseudotanaidae, among

others, have a much more strongly developed pleon than the female and bears well-developed pleopods. []

Even in those taxa in which the females lack the pleopods, these appendages are still present in the |

corresponding males. In taxa that have an abbreviated post-marsupial development in the male, the subadult

males may look quite similar to the females, but can be distinguished by a stouter antenna 1 and somewhat i

more strongly developed pleon. I

Antenna 1

In most tanaidacean species, antenna 1 (Figure 3.2) is uniramous and uniform in structure. Major i

differences occur only between sexes.

Antenna 1 in the female typically is three- or four-segmented. The four-segmented condition is am

most likely plesiomorphic, and, apparently, the three-segmented state is the result of the fusion of the two i

proximal segments. This proposed transformation series is supported by the presence of one or two long

n

setae at the midlength of the first segment; these setae are thought to mark the original segmental suture of i

the plesiomorphic segments. The last segment typically bears one single distal aesthetase, i

i

Antenna 1 of the male is always seven-segmented in those groups that have "swimming males"

(e.g., Anarthruridae, Pseudotanaidae, among others). In these taxa, there is a three-segmented peduncle and i

a four-segmented flagelhtm. Each segment of the flagellum bears a large group of aesthetascs. Some long i

setae typically occur on the terminal segment.

I

Males of the Leptocheliidae are characterized by an antennal flagellum consisting of three or more •

segments. Each flagellar segment is not annular as in the "swimming males" of the other families but more I

elongate, and morphologically similar to a normal segment. Typically, they bear a group of at least 3-5

aesthetascs, i

184 i
I

!

| J /

!

I ,aesthetascs

I

I

I

2

I

I

!

I

I A.1 female A.1 male

Figure 3.2. Generalized first antenna of female andmate tanaidomorphan.

I In the Tanaidae, in which sexual dimorphism is weakly expressed, antenna 1 of the male is only

somewhat more elongate thanin the female, but otherwise is similar.

I Antenna 2 Antenna 2 (Figure 3.3) typically is six-segmented and does not show any great morphological

variability.

I The ftrst segment is small and in most cases semicircular in cross-section. Unfortunately, it is quite

often overlooked, resulting in an incorrect count of segments. The second and third segments are distinct

and each typically bears a distal seta on its outer border. Segment four is elongate and always bent ventrally.

i The distinctly 7-segmented antenna 2 of the members of the genus Heterotanoides Sieg, 1973

i 185
I

(Kudinova-Pasternak, 1977; Sieg, 1973, 1977b, 1986a) and the possible remnant of the segmental suture i

still present in many Akanthophoreinae species suggest that the fourth segment may be the result of the

fusion of two segments. Segment five is similar in length and shape to the fourth, while the sixth is always I

tiny and conical in the Tanaidomorpha. In some genera the sixth segment appears to be fused to the fifth, II

resulting in a "five-segmented" antenna 2.

The antenna 2 of apseudids differ from that just presented for tanaidomorphans. I

Mouthparts

Sexual dimorphism within the mouthparts is one of the characteristic features of the suborder !

Tanaidomorpha. The only exception to this general rule is the Tanaidae in which males and females have I

morphologically identical mouthparts. In all other tanaidomorphan families, adult males have strongly reduced

mouthparts (Sieg, 1984a).

Morphological reduction does not occur in the palpus of maxilla 1 and the epignath in i

tanaidomorphan species. Both are generally found to be functionally active in the respiratory chamber. In

some species, i.e., Siphonolabrumfastigatum Sieg, 1986, the maxilla 2 and remnants of the maxilla 1endite •

may also be present (Sieg, 1986a). In other species, mostly in "swimming males," only the maxilliped l

remains, while the other mouthparts axe fused to the labrum forming a conspicuous "epistome." Leptocheliidae

do not have a functional maxilliped, but instead have two small lamellae occasionally bearing some setae, i

Labrum !1

The labrum is always hood-shaped (a convex flap) and morphologically uniform throughout the

entire suborder Tanaidomorpha (Figure 3.3), except in Siphonolabrum Lang, 1973 (Figure 3.10). iIll[

Mandible

The detailed morphology of the mandible (Figure 3.3) of tanaidomorphans is an important diagnostic _!

character in distinguishing genera and species. For this reason, illustrations of this appendage is mandatory II

in taxonomic descriptions. Unfortunately, some structures, e.g., the laeinia mobilis or the pars molaris,

may look quite different depending upon the viewing angle, i

In all tanaidomorphan genera, each one of the mandibular pair is different (asymmetric) from the

other member and generally lacks a palpus. Each consists of the mandibular body (corpus mandibularis), a

mandibular projection (pars molaris), an articulated "cutting blade" (lacinia mobilis), and a fixed distal I

"cutting blade" (pars incisiva). Some setae located near the lacinia mobilis are present in members of the i

family Tanaidae; these setae are probably the remnants of the much better developed spine-row found in the

Apseudoidea. i

The asymmetry of the mandibles generally is expressed in the reduction of the lacinia mobilis of the

fight mandible. Within the Tanaidomorpha, the lacinia mobilis of the fight mandible is freely articulated

only in members of the Tanaidae, but is fused to the pars incisiva in the remaining families. This fusion i

results in a "two-pointed" pars incisiva. i

Within the Leptochetiidae and Typhlotanaidae these structures are not good generic or specific 1

discriminants because all species within a particular family seem to have nearly identical mandibles, i

i

On the other hand, the lacinia mobilis of the right mandible in the members of the Tanaidae shows

various degrees of morphological reduction and, therefore, is a valuable character in species identification •

(Sieg, 1980b). |

In other families, e.g., Anarthruridae and Pseudotanaidae, the morphology of the pars molafis is one

of the most important taxonomic characters atthe generic and specific levels. As first shown forPseudotanais, •

species that otherwise resemble each other in body shape and armature of appendages can easily be II

distinguished by the shape of the pars molaris (Sieg, 1977a). For many anarthrufidean genera, the general

shape of the pars molaris is characteristic; species can be distinguished from each other by small morphological i

186 I
I

I

I

I 6 endite

\

I

Mxl

I

I Mx2

I palpus

| 1 l'_

A.2

/ parsincisiva

i ,,/pars incisiva

I corpus taciniamobilis 1/

mandibutaris parsmolars,

I Md(r) Md(I)

I

Figure 3.3. Generalized tanaidomorphan second antenna, labrum, right and left mandible, first maxilla, and

I second maxilla.

i 187
!

differences at the apex of this structure. Within certain groups, e.g., Araphura Bird and Holdich, 1984, a !

thin, multi-pointed pars molaris is typical; the number and shape (blunt versus pointed) of the processes

(tips) on the apex are important specific discriminants (Sieg and Dojiri, 1989). Ill

Finally, within many genera of the subfamily Leptognathiinae and Anarthrurinae the pars molaris is II

more or less reduced and represented only by athin weak lobe (e.g., Tanaopsis G. O. Sars, 1882) or is totally

reduced as in Siphonolabrum Lang, 197t. I

First and Second Maxillae

Maxilla 1 (Figure 3.3) exhibts only slight variation among the various tanaidomorphan taxa. It I1

consists of a single endite and a one-segmented palpus. The number of spines on the endite is constant for all |

species within a specific genus, but may vary among the genera (Lang, 1967). So far the lowest number of

observed spines is five and the highest twelve. The palpus typically bears two long terminal setae, except in ib

the family Tanaidae in which a greater number of setae is present in the majority of species. I

Maxilla 2 (Figure 3.3) is oval in shape throughout the Tanaidomorpha and occasionally bears one or

two setae. I Paragnath (= Labium or Hypopharynx)

The labium (Figure 3.4) consists of two lobes that are medially fused in the proximal third. It I

consists of an inner and outer lobe with the outer lobe bearing a small one-segmented palpus distally in the •

plesiomorphic family Tanaidae. In all other tanaidomorphan families the palpus is absent and the outer lobe In

has undergone further reduction. In most species of the families Anarthruridae and Typhlotanaidae, the

outer lobe is represented by a convex protrusion, while it is totally missing in the Pseudotanaidae. !

Thoracopods

As in all malacostracans, the Tanaidacea have eight thoracic somites of which two, without exception, Ii

are fused to the cephalon forming a cephaiothorax. The first pair of thoracopods, the maxilliped, is specialized II

and serves as an additional pair of mouthparts. The second pair has been modified into a chellped. Therefore,

out of the ancestral eight thoracopods, only six pairs of walking legs or pereopods are present. 1

II Maxflliped and Epignath

The maxilliped (Figure 3.4) is the posteriormost appendage of the mouthparts. The coxa, if present, m

is small and situated at the base of the appendage. The basis is the largest segment and may be partly or 1

completely fused to the other member of the pair. The distal portion of the basis carries a pair of protrusions

or lobes referred to as the maxillipedal endites. The endopod consists of four segments and articulates with m

the basis. A free isehium is invariably absent and is probably integrated into the basis. The remaining I[

endopodal segments, collectively known as the palpus, are the merus, carpus, propodus, and dactylus. An

exopodite is never present throughout the entire order. n

Within the Tanaidomorpha, the progressive fusion of the basal region of this appendage is of some !

taxonomic value. The most plesiomorphic character state is represented by the Tanaidae. In this family, the

coxa is still present and the medial borders of the basis, as well as the maxillipedal endite, are not fused. In 1

all other families, the coxa is absent. In the Leptocheliidae, the basis remains unfused as in the Tanaidae, but |

the coxa is missing. One of the more derived character states is represented by the subfamily Pseudotanainae

in which the bases and the endites are completely fused, forming a plate-like structure and enclosing the Ill

posterior portion of the mouth chamber. Anarthruridae and Typhlotanaidae exhibit various degrees of fusion I_

along the medial margins of the basis and occasionally of the endite.

u

The distal margin of each endite usually bears one or two setae and some translucent hemispherical !

protrusions. In some taxa, the presence or absence and the spatial arrangement of these structures may be of |

taxonomic importance.

!

188 i
I

I

I

I _i°"°°

I

I

, 1

I

I Mxp

basis

!

!

!

I Figure 3.4. Generalized tanaidomorphan labium, maxilliped, and epignath.

I

i 189
!

The basis always bears at least one seta close to the articulation of the palpus. The number of setae I

on the patpus may play an important taxonomic role in distinguishing genera, especially within the family

Leptocheliidae in which some taxa have been described as having up to eight. -_

So far, the available information on the setation of different palpal segments is not reliable enough !1

to draw any definitive conclusions. The merus (first segment) usually does not bear any setae. The carpus

(second segment) usually is equipped with an outer seta which appears spiniform in some species. The inner I

margin of this segment bears several elements, variable in number as well as in structure: some are setiform, il

while others are highly sclerotized and spiniform, and sometimes barbed. The number and the combination

of spines and setae on this segment might be uniform within the genus or subfamily. The same holds true for i

the propodus (third segment), but this segment never bears an outer seta. The terminal (fourth) segment, the I

dactylus, is digitiform and typically bears one small outer seta and a group of inner setae variable in number

and size. m

Sexual dimorphism in the maxillipeds is common except in the Tanaidae. Even in families in which

the males lack mouthparts, remnants of the maxilliped and a normally developed epignath are present.

Typically, however, the maxilliped in the male is well-developed. Differences between the two sexes in I

these taxa are found in the setation and shape of the segments. In contrast, the maxilliped is almost completely II

reduced in the males of Leptocheliidae. Typically, it is only represented by a small plate-like structure that

bears one or two setae. In some leptocheliid males, the remnants of the palpus are represented by knoblike i

projections. II

Articulated to the maxilliped of all Tanaidomorpha is a posteriorly-directed epipadite, known as

the epignath. In the Tanaidae it is a broad, kidney-shaped structure, while in all other families it is small and n

elongate (sausage-like). Sometimes the tip is elongate and resembles a whiplike filament. IN

Cheliped lib

The general structure of the tanaidomorphan cheliped (Figure 3.5), especially the presence or absence i

of a coxa, has been a subject of controversy (Lang, 1971; Lauterbach, 1970; Sieg, 1980a, 1984a) and can

only be discussed in conjunction with a discussion of the carapace.

Tanaidomorphan taxa all have a well-developed dorsal carapace which curves ventrally to form a I

ventral chitinous plate that houses the branchial chamber into which the palpus of maxilla 1 extends and the

epignath is located.

| Lang (1971 ) stated that there exist some taxa that have a coxa and others that lack this segment. On

the other hand, Lauterbach (1970) in his paper on the cephalothorax of Tanais cavolinii Milne-Edwards,

1840 [= T. dulongii (Audouin, 1826)] believed the coxa to be absent and used the term side-piece

("Seitenstiick") in alluding to this structure. This side-piece, or plate, that lies near the insertion of the U

cheliped is probably a remnant of the stemite of the second thoracic segment and not the chelipedal coxa as

suggested by some authors. These sclerotized plates were referred to as the "coxa" by Lang (1971) and !

"pseudo-eoxa" by Sieg (1980b, 1984a); the latter author confirmed Lauterbach's opinion that a chelipedal 11

coxa is absent in all tanaidomorphan families.

The morphology of the cheliped is uniform throughout the Tanaidomorpha. The cheliped always I

consists of a well-developed basis which is articulated to the cephalothorax along its proximal margin or at W

a point slightly removed from this margin and indicated by a conspicuous projection. The ischium is generally

thought to be absent, except in the Neotanaidomorpha (Gardiner, 1975). The merus is small and more or i

less triangular having the carpal articulation along the tergal margin. The carpus is a very large segment

bearing two setae on the tergal border, usually one on the proximal and another on the distal areas; the

sternal border usually bears some longer setae. The propodus and dactylus together form the chela. The i

sternal part of the propodus is greatly expanded and is called the "fixed finger." The tergal margin of the n

fixed finger may be characteristically denticulated, while the sternal margin usually bears three setae. The

I

190 I
1

1

rostral __._" dactylus

I propodus distal

I _iniform tips

I c_

%.

1 _°" proximal

merus

I side-piece coxa

i

i basis _.-

proximal

!

i tergal

I rostral caudal

I

i sternal dactytus terminalspine

distal

I

Figure 3.5. Generalizedtanaidomorphan cheliped and pereopod.

I

I 191
!

fixed finger and dactylus have strongly sclerotized tips. Typically, there is a row of smaller setae, often I

referred to as the "comb," located on the rostral side near the articulation of the dactyhis.

Sexual dimorphism in the Tanaidomorpha is often primarily expressed in chelipedal differences.

This is a common feature, especially in the Leptocheliidae in which up to four different kinds of chelipeds Ii

can be distinguished in one species [e.g., Heterotanais oerstedti (Kr0yer, 1842); see Biickle-Ramirez, 1965;

Sieg, I984a]. Fortunately, none of the genera in this atlas has such a complicated post-marsupial development, l

On the other hand, "swimming males", which exhibit a general habitus different from that of their I

corresponding females, possess chelipeds morphologically similar to that of the female.

Pereopods i,

The remaining six pairs of thoracopods are called pereopods 1-6 (Figure 3.5). The cheliped has

been referred to as the first pereopod by some authors; this misnomer results in seven pairs of pereopods. am

However, since "pereopod" refers to a "walking leg" and Bate (1856) clearly restricted the term to the

appendages of"free" thoracic somites, we prefer the terminology used herein. This point has been discussed

in the past to some extent (Wolff, 1956; Lang, 1957; Sieg, 1980b; Messing, 1981).

All pereopods are styliform. Morphological adaptations of the pereopods to different kinds of i

sediments or habitats, observed in the Apseudoidea, are not found in the Tanaidomorpha. Typically, each

pereopod consists of a coxa, basis, ischium, merus, carpus, propodus, and dactylus which bears a strong I

terminal spine. An exopodite is never developed in the Tanaidomorpha. Within the family Tanaidae, the |

ischium is absent and is most probably fused with the basis.

The six pairs of pereopods can be separated into two functional groups. Females develop oostegltes i

on pereopods 1-4, or exclusively on pereopod 4. II

Pereopods 1 to 3. The first three pairs of pereopods are attached to the anteroventral comer of the

pereonites, are directed anteriorly, always have free coxae, and are used for spinning the silk tube. The coxa i

always has a bicondyle-type articulation which lies in a longitudinal (anterior-posterior) direction. The !1

basis also exhibits this type of articulation, but lateral in orientation. This "cardanic" articulation allows the

animal to move the leg in all directions. The terminal spine of these pereopods is never fused to the dactylus, i

Its morphological independence appears to be correlated with the special function of spinning a silk tube

which is produced by the thoracic glands,

The armature of each segment is important for distinguishing genera and identifying species. In I 

plesiomorphic taxa, pereopods 1-3 bear only setae on each segment. In more derived taxa, especially in the I

Anarthruridae, spines have secondarily developed on the merus and carpus, the number and position of

these spines are taxonomically useful characters, i

III

In most taxa, pereopod 1 is more slender than the next two pairs of pereopods and usually does not

have spines on the merus and/or carpus. m

Pereopods 4 to 6. The last three pairs of pereopods are attached to the posteroventral comer of the i

pereonite, are directed laterally, have a coxa fused with the sternite of the pereonite, and are used for clinging

to the tube.

am

The dactylus and terminal spine are often fused to form a claw. The fusion produces a large, I

functionally strong prehensile structure that is used to grasp the inner surface of the tube. This structure is

very characteristic for some taxa (e.g., Tanaidae). In the Anarthnu'idae the dactylus is uniquely modified: Im

the sternal border of the elongate dactylus is concave and the two resulting ridges are lined with setules U

(hairs). The basis can be quite massive in many Typhlotanaidae.

The fourth and fifth pair of pereopods are quite similar in appearance and armature. However, the I

propodus of pereopod 6 bears some additional distal setae or spines on the sternal border in most taxa. The I

armature of each segment of these pereopods is as taxonomically important as in the preceding three pairs.

!

192 I
!

I Marsupium

Three different types of marsupia or broad pouches exist within the Tanaidomorpha.

i The "normal- (= Heterotanais-) type" is represented by all families except Tanaidae and Pseudotanaidae. This type of marsupium is formed by four pairs of oostegites on pereopods 1-4. Each

oostegite develops from the coxa as a bilamellar sheet, containing some epithelial tissue within the

I interlamellar much space. After the animal moults, the oostegites start to grow. Each sheet of the oostegite becomes thinner and most of the epithelial tissue is lost or resorbed. The eggs tie between the oostegites and the

ventral surface of the body wall.

I The "Pseudotanais-type" is quite similar to the normal-type. However, only one pair of oostegites

are present; they occur exclusively on pereopod 4, and cover the genital opening.

The "'Tanais-type'" is unique. Although it also consists of only one pair of oostegites on pereopod 4

I as thePseudotanais-type, morphology oostegite very During early developmental

in the of the is different. the

stages, these oostegites are not sheet-like but more or less sac-like and filled with tissue. Later in the adult,

the eggs are laid or placed inside the oostegites, resulting in brood sacs or "avisaes" (Lang, 1960). How the

I inside the oostegites and how they fertilized is still unknown. The nutrition of the developing eggs get get

embryos was recently studied by Johnson and Attramadal (1982).

I PleopodsThe pleopods (Figure 3.6) are morphologically very uniform throughout alltanaidomorphan families.

The basis articulates with a one-segmented exapodite and a one-segmented endapodite. The outer border

I of (naked). the exopodite The outerbears border a rowof of themany, endopodite long, pinnate is similarsetae, to that whileof the the entire exopodite. innerThe border innerremains border unarmed bears at

least one distal seta, but it may have several setae in some species of the Tanaidae.

I Variation in setation is quite common and may be taxonomically valuable. Very often the outer margin of the exopodite bears a proximal "specialized" seta spatially separated by a distinct gap from the

other setae (Figure 3.6). This seta typically is slightly stouter than the others and is not pinnate, but pilose.

I Reduction in the number of setae is very common and, therefore, both pleopodal rami may bear only a few distal setae. Frequently, these setae may not have pinnae, consequently, are naked.

In the adults of some species, pleopodal setation is absent, but much more common is the complete

I loss of the pleopods. Since mane.a-stages and juvenUes of several species also tack pleopods, this character has to be used with caution, since the juveniles of one species could be confused with the adults of other

species.

I Sexual dimorphism is observed in allthose species in which "swimming males" reported.

have been

Often the females lack pleopods while the corresponding males have very large ones [e.g., Pseudotanais

forcipatus (Lilljeborg, 1864)].

I Uropods

The uropods (Figure 3.6), which articulate on the pleotelson, are the last pair of appendages. They

I are typically biramous structures consisting of a one-segmented basis, a short exopodite, and a longer .endopodite. In contrast to the Apseudoidea, the uropods within the Tanaidomorpha are not filiform, but are

styliform. The only tanaidomorphan species that have uropods morphologically similar to those of the

i Apseudoidea are some species of the Tanaidae. Reductions in the number of exopodal and endopodal segments is a common feature and has great

taxonomic relevance. Generally, the plesiomorphic families (e.g., Tanaidae and Leptocheliidae) typically

i have more than two endopodat segments, while the more derived ones (e.g., Anarthruridae, Pseudotanaidae, and Typhlotanaidae) never have more than two.

!

I 193
I

I

piloseseta

Uro 1

exopodil I

I

endopodite I

p, I

I

I

I

Uro l

._.--- pseudo-exopodte _ _ reducedexopodite I

endopodJte end°p°dite"_ t I

J ,, ore I

I

Figure 3.6. Generalized tanaidomorphan pleopod andrepresentative examples of uropods, i

194 I
!

I The exopodite also has undergone some morphological changes. In some taxa, e.g., the entire family

Tanaidae, the exopodite is absent. A much more common modification is the fusion of an exopodal remnant

I with theThe basis,uropod a feature mayfound be in some genera of the anarthruran subfamily Akanthophoreinae. sexually dimorphic is some taxa. The uropod in the female of Siphonolabrum

fastigatum Sieg, 1986 has the exopodite fused to the basis and the endopodite is two-segmented. The male,

I in contrast, has a small but free exopodite and the endopodite seems to be three-segmented (Figure 3.13).

i Internal Anatomy

The internal morphology of tanaids is still poorly known, especially that of the Tanaidomorpha. For

i detailed light-microscopical studies of the internal anatomy of tanaids, the reader is directed to Siewing (1953) and Lauterbach (1970). Only a brief account is provided herein.

The nervous system consists of a brain, a subesophageal mass, and a ventral chain. The eyes are

I msessi outh, le, aifstprese omacnht, (part and may of theorfomay regut)notwhi havech visual is dividedelements. into a filter The aandlimenta a masticatory ry tract cchamber, onsists ofa saynvceyntial tral

midgut, and a terminal anus. There axe two pairs of hepatopanereases, and one pair of maxillary glands

i fbase or exofcretio then.sixth The pair gonadofs

thoracic are paired; legs;thetheovaries testesaxaree coconne nnected ctedtototheoneoviducts or two which genitalopeconn laterall es on ytheatlast the

thoracic segments via the vas deferentia.

i l Development

I Sexual dimorphism is common, but present only when accompanied by hermaphroditism,

protogyny, and possibly protandry. Females produce several broods. Development of eggs takes place

I • inside a marsupium. The post-marsupial development of the order is only partly understood. In all cases,

i g development is direct and there is no planktonic larval stage. After hatching, the embryo, still contained

within the broodpouch, changes (without a molt) to the first post- embryonic stage (manca-I); the last pair of

I pereopod of these sappendages and all pleopo aredpresent. s have nInotaddition yet developed. to theseIndifferences, the second stage both man (manca- ca stages ff), thecanrudbeimdistinguished ents (anlage)

from the adult by the proportionately smaller pereonites. The next stage is termed "neuter" and represents

i the preparatory female/male which isfollowed by the adult. Post-embryonic instars and adult females resemble

each other closely, but usually differ markedly from the males, which often do not feed, having reduced

mouthparts, and a sealed anus.

I embryonicApseud development omorphansof arePagurapseudes probably all plargoensis rimarily gonocho reportedristbyie Messing (Sieg, 1984a (1979). ) anSexual d followdimorphism the postamong members of this suborder is only weakly developed, and is reflected in the first antenna and cheliped.

I On the other hand, members of the Tanaldomorpha exhibit a great variety of post-marsupial developmental pathways. A gonochoristic type of development similar to that of the Apseudomorpha is

found in the plesiomorphic tanaidomorphan taxa (e.g., Tanaidae). Reduction of the mouthparts in the male

I phase somefirnes leads to a highly complicated post-marsupial development expressed byup to four different types of males (e.g., Leptochellidae). Mainly, the secondary males may show strongly developed dimorphism

resulting in huge and striking chelipeds as well as totally different first antenna. More apomorphic families

I (e.g. for the , Ansearch arthruridae, of thePseu female dotanaidae, ("swimming and others) male").areHowever, also gonochoristic, since malesbuthave the sole not yet primary been male discovered is adaptedfor

many taxa, parthenogenesis may be the means of reproduction in these genera. Sexual dimorphism and

I protogyny are common in this suborder.

i 195
!

Ecology and Biology I

Tanaids normally build tubes by spinning silk produced by glands having their outlet on the dactylar IN

tips of the first to third pereopods. During construction, feces and detritus are added to the silk. These tubes i

can be found in the sediment or among algae, hydroids, and sponges. The presence of these tubes in f'me

sediment may reduce erosion, thus resulting in a more stable sandy bottom. When building a new tube, i

juveniles bore through the wail of the "mother tube" and build their own nearby (Biickle-Ramirez, 1965).

Therefore, tanaidaceans have patchy distribution with high population densities. High population densities

on the magnitude of 104to 105/ m-2can be observed in shallow waters. Consequently, they are an important i

part of the food web within marine ecosystems: tanaids serve as food for polychaetes, amphipods, decapods, II

fishes, and some water birds.

Analyses of the stomach and gut contents indicate that tanaids are usually scavengers or detritivores, I

although some may be raptorial carnivores. Their food normally consists of detritus or small algae, mainly m

diatoms. They may occasionally feed on nematodes and harpacticoids (Feller, 1978). Only the kailiapseudids

are filter feeders (Lang, 1956a) as indicated by the setal structure on their maxilliped and cheliped (Sieg, i

1984a).

Very little is known concerning habitat preference. Within the suborder Tanaidomorpha, there are i

some indications that several families occur in more or less regularly distinct habitats. For example, Tanaidae 1

are common in algal mats. If there is also plenty of sediment between these algal filaments, members of the R

family Paratanaidae are also found in this habitat, although they are more typical of sandy bottoms. If the

sand is less coarse or if the percentage of mud increases, members of the Leptognathiinae and Pseudotanaidae l[

would replace members of these other families. m

Zoogeographic Distribution I

Tanaids have a worldwide distribution, inhabiting the intertidal zone down to the hadal zone (nearly i

8,000 m depth). This order is not well known from tropical waters and in the deepsea, simply because the

group has not been extensively studied from these regions. It does, however, appear to be well represented

in the deepsea (e.g., Holdich and Bird, 1985; Sieg, 1983a, 1984a). Even though tanaids are not very diverse i

in shallow waters, the few species that are present in a specific locality may be very abundant there (Barnard, i

1970). As in other peracarids, tartaids also show an increase in species diversity with increasing depth

(Hessler et al., 1979). According to Wolff (1977), tanaids may comprise as much as 19%(by number) of the i

benthic macrofauna at 5,000 m depth, and quite often the order is found to be the second most abundant

I

peracarid group, next to amphipods, in ecological studies of the deep sea.

The zoogeography, and even the descriptive distributional pattems, of the Tanaidacea have been so i

far largely ignored by biologists. This is mainly a result of the difficulty in accumulating and analyzing the

m

information (data) contained in the scattered literature. Recent monographic revisions (Gardiner, 1975;

Sieg, 1977a, 1980b) and a bibliographic index listing all known literature pertaining to this order (Sieg, 1

1983b) have helped in the compilation of the published literature. In an attempt to help analyze distributional

patterns, an EDP-database was established at the Universi_t Osnabruck, Abteilung Vechta (Sieg, 1984). i

Initial use of this database aided in the recognition and understanding of the distributional patterns of a few i

families (Sieg, 1980b, 1983b) and species composition within a given area (Sieg and Heard, 1983). The

most comprehensive use of this database involved all the available information on the Tanaidacea and

allowed the study of worldwide distributional patterns of this order (Sieg, 1986b). 1

!

Data has revealed that the shallow water regions of the southern hemisphere, specifically around

Antarctica, may have been the "evolutionary center" for this order (Sieg, 1986b). A large number of tanaid 1

families and

genera, along with a very high degree of endemism, supports this contention, l

196 i
!

I The tanaid composition between the temperate North Atlantic and the temperate North Pacific

regions are very different. The North Atlantic has been studied more in relation to tanaids than the North

I Pacific, which may account, in part, for the huge difference between these two areas: the great dissimilarity between these areas may be more apparent than real. Since our tanaidacean fauna of the southern temperate

region is still only poorly known, especially in the southern part of South America and in the New Zealand

i area, comparisons among it and other zoogeographic areas may be preliminary at best. Our scanty information suggests that the northern temperate waters of the Pacific and the Atlantic Oceans share a more similar

tanaid composition with the tropical warm waters of these oceans than with the southern temperate regions

of these bodies of water.

I The tropical warm water areas of the Atlantic, Indian, Indo-West Pacific, and Pacific zoogeographic

subregions are more diverse than the northern or southern temperate zones and are different from each

I other, may have i.e., characterized by independent species compositions. In contrast to isopods and amphipods, tanaids

their greatest number of species within the tropics. The Atlantic tropical subregion appears to

have a more diverse tartaidacean fauna than the Pacific tropical subregion, but, as in the temperate waters,

i this may be an artifact of our past preference for studying the Atlantic Ocean. The tanaid fauna of the tropical Atlantic region, which includes the Caribbean Sea, Gulf of Mexico, and Mediterranean Sea, seems

to have a closer relationship to that of the tropical Pacific than to the Indian Ocean. The Indo-West Pacific

i fauna shares some species with both the tropical Pacific and Indian Oceans, a result consistent with its intermediate geographic position between these two oceans.

I Depth Distribution

I Several Tgeneralities he distributioconcerning n of tanaids in relation to the depth has been summarized (Sieg, 1983a, 1986b). the tanaid families have been noted. For example, the Apseudidae appears

to have a large bathymetric range, extending from shallow water (0-200 m) to nearly 6,000 m.

I The Neotanaidae (not treated in this atlas), the lone family within the suborder Neotanaidomorpha, is typically deep-sea. The depth-distribution of this family has been discussed by Wolff (1956) and Gardiner

(1975).

I Finally, the depth-distribution of the Tanaidomorpha has been reviewed by Sieg (1983a, 1986b). The plesiomorphic families (i.e., Tanaidae, Leptocheliidae, and Paratanaidae) are, with some exceptions,

shallow-water taxa. The more derived families (i.e., Anarthruridae, Typhlotanaidae, and Pseudotanaidae)

I ocLeptognathiinae, cur throughout a broad depth range from shallow water to hadal depths (below 6,000 m). The a subfamily of the Anarthruridae, exhibit a high species diversity at bathyal depths (200-2,000

m), which gradually decreases in abyssal (2,000-6,000 m) and hadal depths.

I In general, arelatively large number of tanaid species, contrary to most other crustaceans, to

appear

have broad depth ranges, ranging from shallow water to the deep sea. However, as the taxonomy of the

Tanaidacea becomes better known, this perception may be modified.

I

i Materials and Methods

The specimens examined were part of an extensive voucher collection of Crustacea collected for

i the MMSISanta Maria Basin and Western Santa Barbara Channel Study and Atlas, supplemental MMS

material from the bulk collection obtained from Battelle, a collection of Tanaidacea housed at the Los

Angeles County Museum of Natural History, and specimens collected by the Biology Laboratory,

Environmental Monitoring Division, Bureau of Sanitation, Department of Public Works of the City of Los !

I 197
!

Angeles. Selected specimens were measured and dissected in glycerol. The appendages were mounted on i

slides according to the procedure described by Sieg (1973).

and SantaAllBarbara type andMuseum non-typeof Natural specimensHistory, have been exceptdeposited in the National Museum of Natural History I those on loan from the Los Angeles County Museum

of Natural History.

!

Aknowledgements

We thank Charles (Tony) Phillips, Environmental Monitoring Division (EMD, City of L.A.), Roy I

Kropp, formerly of Battelle Ocean Sciences - Ventura Operations, Paul Scott, Santa Barbara Museum of

Natural History, and Hans Kuck, formerly of the Los Angeles County Museum of Natural History for I providing specimens and collection data. Jim Roney (EMD, City of L.A.) and Isabelle Williams (ENSR,

Woods Hole) are acknowledged for their critical review of the manuscript and their helpful suggestions.

Final acknowledge ly, we thank Jim Blake (ENSR, Woods Hole), for asking us to undertake this study. We gratefully I the Universit_t Osnabriick for providing travel funds to the first author (MD) for his trip to

Germany where the initial part of this study was done.

Abbreviations Used in Keys and Figures I

A. 1 = first antenna Mxp = maxilliped I

A.2 = second antenna Epi = epignath

L = labrum Che = cheliped I

Md(r) = right mandible P. 1-P.6 = pereopods 1-6

Md(1) = left mandible P1.l-P1.5 = pleopods 1-5 I

Mx. 1 = first maxilla Pit = pleotelson

Mx.2 = second maxilla Uro = uropod I

La = labium

I

I

I

I

I

I

198 I
!

I Glossary

" The following terms are defined with respect to their application to the Tanaidacea.

I Abdomen: See pleon. Caudal: Term introduced by Lang (1968) to

describe a specific surface or border of body

I Aesthesclerotized), tase: A setiformcylindrical element, throughout thin-walledentire(not sposterior. omite or appendage; synonymous with

length, with blunt tip, probably

chemosensory, and occurring exclusively on Cephalon: Anteriormost part of body, bearing

I flagellum of antenna 1. eyes (when present), mouth, two pairs of antennae and mouthparts, excluding

Antenna: Second pair of cephalic appendages maxilliped.

i (Antenna 2). Cephalothorax: Anterior tagma resulting from

Antennule: First pair of cephalic appendages fusion of cephalon and first two

(Antenna 1). thoracomeres.

I Alimentary tract: Tubular structure between Chela: Distal end of cheliped formed by propodus

mouth and anus, consisting of foregut, and dactylus.

i midgut, hepatopancreas. and hindgut, and associated with Cheliped: Second pair of thoracic appendanges,

Basis: Second segment (article) of thoracic and adjacent and posterior to maxilliped. lit

I abdominal appendages; sympodal segment Claw: Fusion of dactylus and terminal spine in

III immediately distal to coxa and carrying the pereopods.

rami. Coxa: First segment (article) of thoracic and

I Biramous: A term used to describe an appendage abdominal appendages; sympodal segment

having two rami (branches), an exopodite proximal to basis.

i and endopodite. Daetylus: Seventh segment (article) of thoracic Branchial chamber: Cavity between carapace appendages.

and body wall into which the maxillular

Endite: Medially-directed projection of coxa or

I palp with and the epignath extend; synonymous respiratory chamber, basis.

Brood pouch: See marsupium. Endopodite (endopod): Inner ramus of thoracic

I and abdominal appendages. Brood sac: Specialized marsupium of the family

Tanaidae; each sac-like oostegite ("ovisac" Epignath: Epipodite of maxilliped.

I sensu Lang, 1960) houses eggs within its Epipodite: Exite of coxa with branchial function;

cavity, exclusively found on maxilliped.

i Carapace: Outgrowth of the maxillary segment Epistome: Portion of cephalothorax of male, which is fused with tergites of first and posterior to labrum and consisting of fused

second thoracomere; it is curved ventrally, remnants of mouthparts.

covered on its inner surface with a

I Exopodite (exopod): Outer ramus of thoracic and

respiratory epithelium,

and encloses

respiratory chamber, abdominal appendages.

I Carpus:appendages, Fifth segment (article) of thoracic Flagellofum:antenna Multisegmented 1 or 2. distal portion (ramus)

Fixed f'lnger: Distally-directed projection of

I chelipedal propodus.

I 199
!

Foregut: Anterior part of alimentary tract, Manca(-stage): Post-embryonic developmental

consisting of esophagus and stomach, stage, lacking sixth pereopod and pleopods

(Manca-1) or having rudiments of these _l

Genital cones: Conical projection on sternite of appendages (Manca-ID.

sixth pereonite of male; this projection bears

one or two gonopores. Mandibles: Pair of appendages immediately

posterior to labrum; functions in masticating I

Gonochoristie: Sexes separate; condition in food. ==

which individuals remain the same sex

throughout their entire life cycle. Marsupium: Ventral chamber in female formed 1

by oostegites in which eggs and embryos are I

Grinding area: Flat distal end of processus brooded.

molaris which typically is surrounded by a

slightly crenulated wall; often reduced or Maxillae: Third pair of mouthparts (maxilla 2). I

lacking. Maxillary gland: Excretory organ situated in

Hepatopancreas: Caeca located at border of fore- maxillary somite with its opening on II

and hindgut and mostly directed posteriorly; maxilla. |

occasionally one pair is directed anteriorly.

Maxilliped: First pair of thoracic appendages;

Hermaphroditism: Condition in which specialized for feeding. R

individuals develop functional male and

Maxillules: Second pair of mouthparts (maxilla 1). female genital organs - either

simultaneously or sequentially. Merus: Fourth segment (article) of thoracic I

Hypopharynx: See labium, appendages.

I

Hyposphaenium (pl. hyposphaenia): Process Molar process: See pars molaris. 1

(hook) on stemite of pereonite or pleonite. Neuter: Post-embryonic stage with full set of

Incisor process: See pars incisiva, pereopods and pleopods, but without

developed gonads ("juveniles"). 1

Isehiuln: Third segment (article) of thoracic

appendages. Oeellus: A simple eye or pigment spot functioning

as light receptor, tm

Juveniles: Post-embryonic stages that show Oostegite: Normally thin, flat plates arising from |

neither sexually defined morphological

structure nor have developed functional inner margin of coxae of pereopods 1 - 4 or

gonads. only 4. I

u

Labium: Flat, bilobed structure situated posterior Ovary: Female gonad.

to mouth, with or without a one-segmented Oviduct: Tube in female for passage of ova from •

distal palp; not homologous with a ovary to gonopores. il

segmental appendage; lower lip.

Ovisaes: Marsupium of Tanaidae; saclike m

Lahrum: Bulbous, hood-like structure situated structure within which eggs are brooded. l

anterior to mouth; upper lip.

Palpus (or palp): Segmented or unsegmented

Laeinia mobilis: Articulated "cutting blade" of structures of different origin in mandible,

mandible;developed mainly on left maxilla 1, labium, and maxilliped.

mandible, very often absent on right

mandible. Paxagnath: See labium. •

II

Lower lip: See labium. Pars ineisiva: Fixed (non-articulated) distal

"cutting blade" of mandible; synonymous

with incisor process. •

I

2oo |
!

I Pars molaris: Medial projection of mandible Somite: Body segment, usually covered with

originally used for grinding; synonymous sclerotized tergite and sternite.

I with molar process. Sternal: Term introduced by Lang (1968) to

Peduncle: First three segments of antenna 1 and describe a specific surface of body somite or

first five segments of antenna 2. appendage; synonymous with ventral.

I Pereon: Free thoracomeres 3-8; anterior portion Sternite: Ventral sclerotized surface of body.

of trunk, excluding maxillipedal and

chelipedal somites. Subadult males: Second juvenile stage of males

I that are sexually mature.

Pereonite: Somite belonging to the pereon. Swimming male: Highly transformed male

i Pereopod: Walking leg; appendage belonging to having a large pleon with well-developed one of six free thoracomeres (3-8). pleopods and a small pereon; mouthparts

reduced, except maxilliped.

Pleon: Posteriormost body tagma, consisting of

I pleonites and pleotelson. Syrnpodite (adj. sympodal): First two segments (coxa and basis) of thoracic or abdominal

Pleonite: Somite belonging to pleon, appendages; synonymous with protopod(ite).

I Teison: Posteriormost part of body, beating anus;

Pleopod: Appendage

of pleonite,often natatory.

Plcotelson: Structure resulting from fusion of at in Recent tanaidaceans always fused to sixth

i least sixth pleomere (abdominal somite) and pleonite to form pleotelson.

telson. Tergal: Synonymous with dorsal (Long, 1968);

Pcopodus: Sixth segment (article) of thoracic see sternal.

I Tergite: Sclerotized dorsal surface of body.

appendages.

Protandry (adj. protandrous): Hermaphroditism Testes: Male gonads.

i in which functional male phase precedes female phase. Thoracic glands: Compound glands lying in first

three pereortites and secreting silk-like

Protopodite (protopod): First two segments substances that are transported via a channel

I (coxa and basis) of thoracic or abdominal appendages; synonymous with sympodite to end of dactylus and used for constructing tubes; restricted to Tanaidomorpha.

(adj. sympodal).

I Protogyny (adj. protogynous): Hermaphroditism Thoracofastmeand re: second Thoracicthoracomere somite; in tanaidaceans are fused with

in which functional female phase precedes cephalon and bear specialized

i Pseudobi maleramphase, ous: Condition in which two rami appendages, respectively. maxilliped and cheliped,

are present in uropods, but one is fused to Uniramous: A term used to describe an

I proximal segment (basis). appendage having only one ramus (branch),

Pseudo-coxa: Posterior part of cephalothorax to an exopodite or endopodite.

I which cheliped articulates. Upper lip: See labrum.

Rostral: Synonymous with anterior (Long, 1968); Uropods: Styliform abdominal appendages of the

see caudal, pleotelson; usually biramous, but often

I Sclerite articulated side uniramous or pseudobiramous. Side-piece:

on one to

carapace and on other side to area posterior Vas deferentia: Tubes in male for passage of

i to a distal projection of chelipedal basis; sperm from testes to gonopores.

formerly called coxa.

I 201
!

List of Species I

Suborder Apseudomorpha Sieg, 1980

Family Apseudidae Leach, 1814 _l

Genus Carpoapseudes(Lang, 1968) M

Carpoapseudes caraspinasus Dnjiri and Sieg, new species

Suborder Tanaidomorpha Sieg, 1980 1

Family Tanaidae Dana, 1849 II

Genus ZeuxoTempleton, 1840

Zeuxo maledivensis Sieg, 1980 1

Family Leptocheliidae Lang, 1973 |

Genus Leptochelia Dana, 1849

Leptochelia dubia (Krcyer, 1814) •

Family Paratanaidae Lang, 1949 I

Genus Paratanais Dana, 1849

Paratanais intermedius Dojiri and Sieg, new species II

Family Anarthruridae Lang, 1971 II

Subfamily Anarthrurinae Lang, 1971

Genus Siphonolabrum Lang, 1972 •

Siphonolabrum cahforniensis Dojiri and Sieg, new species U

Subfamily Akanthophoreinae Sieg, 1986

Genus Paraleptognathia Kudinova-Pasternak, 1981 •

Paraleptognathia cf. gracilis (KrOyer, 1842) II

Paraleptognathia bisetulosa Dojiri and Sieg, new species

Genus ScoIoura Sieg and Dojiri, 1991 Inn

Scoloura phiUipsi Sieg and Dojiri, 1991 Ii

Genus Chauliopleona Oojiri and Sieg, new genus

Chauliopleona dentata Dojiri and Sieg, new species 1

Genus Araphura Bird and Holdich, 1984 |

Araphura breviaria Dojiri and Sieg, new species

Araphura cuspirostris Dojiri and Sieg, new species Ill

Genus TanaelIa Norman and Stebbing, 1886

|

Tanaella propinquus Dojiri and Sieg, new species

Genus Incertae Sedis

Male species 1 I_

u

Male species 2

Male species 3 1

Subfamily Leptognathiinae Sieg, 1973 m

Genus Leptognathia G.O. Sars, 1882

Leptognathia cf. breviremis (Lilljeborg, 1864)

Genus Tanaopsis G.O. Sars, 1896 1

Tanaopsis cadieni Sieg and Dojiri, 1991

Family Typhlotanaidae Sieg, 1984

Genus TyphIotanais G.O. Sars, 1882 !

Typhlotanais williamsae Dojiri and Sieg, new species

HI

Typhlotanais crassus Dojiri and Sieg, new species

Family Pseudotanaidae Sieg, 1973 I!

Genus Pseudotanais G.O. Sars, 1882 I1

Pseudotanais makrothrix Dojiri and Sieg, new species

Pseudotanais cah'forniensis Dojiri and Sieg, new species I

202 I
!

I Key to Families

I 1A. A.1 biramous; Md with palp (suborder Apseudomorpha) ................. Carpoapseudes caraspinosus lB. A.1 uniramous; Md without palp (suborder Tanaidomorpha) ........................................................ 2

I 2A. Mouthparts (except maxillipeds in some families) reduced to lobiform structures (generally males)

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

I 2B. Mouthparts not reduced (females and some males) ........................................................................ 3

3A. P. 1 - P.6 without ischium; Uro uniramous; Mxp with coxa and with basis not fused medially; last 2

I smaller than 3 with 3 4 with 3 of pleonites preceding ones (occasionally only pleonites), pairs or

pleopods ............................................................................................................................... Tanaidae

i 3B. P. I - P.6 with ischium; Uro typically biramous, sometimes pseudobiramous; Mxp without coxa,

basis fused or unfused medially; typically with 5 pleonites of equal size ...................................... 4

I 4A. Endopodite of Uro at most with 2 segments ................................................................................... 5

4B Endopodite of Uro with more than 2 segments (Mxp not fused medially; with eyes) .....................

i ................................................................................................................................... Leptocheliidae

5A. Endite of Mxp enlarged and laterally expanded ........................................................... Paratanaldae

I Mxp enlarged nor laterally expanded ........................................................................

5B. Endite of not 6

I 6A. Merus cross-section, of P.1 andwithboth1 tergmargins al spine; covered dactylwithus fine of character setules),isticnot fused shapewith(eloterminal ngate, semic spineircu............. lar in

................................... Anarthruridae (subfamily Akanthophoreinae and subfamily Anarthrurinae)

I 6B. Merus of P.1 with setae dactylus of typical shape (not elongate), usually fused to terminal

only;

spine ................................................................................................................................................. 7

I 7A. Pars molaris of Md broad, grinding area surrounded by crenulated wall ................ Typhlotanaidae

7B. Pars molaris of Md reduced, small, grinding area without crenulated wall, and terminating as an

I attenuate process or several points, or bearing several spiniform elements ................................... 8

8A. Carpus of P.2 with 1 tergal spine; pereonites 1 and 2 extremely small, about 5-7x broader than long;

I marsupium formed by only 1 pair of oostegites ....................................................... Pseudotanaidae

8B Carpus of P.2 with only setae; pereonites I and 2 of typical shape, mostly about 3-4x broader than

i long; marsupium formed by 4 pairs of oostegites ........ Anarthruridae (subfamily Leptognathiinae)

9A. With eyes ....................................................................................................................................... 10

I 9B eyes ..................................................................................................................................

Without 11

!

i 203
!

10A. Mxp reduced to lobiform structure, lacking palpus; A.1 with more than 7 segments; Che much I

larger than and different in shape from that of female ................................. Leptocheliidae (males)

10B. Mxp not reduced to lobiform structure, but modified from that of female, palpus always present;

A. 1with 7 (?) segments, first 2 segments distinctly broader than remaining ones; Che of similar size

and shape to that of female ............................................................................... Paratanaidae (males)

!

11A. Che articulated to cephalothorax along proximal margin of basis ....................................................

............................................................................. Anarthruridae (males - subfamily Anarthrurinae)

11B. Che articulated to cephalothorax by a "side-piece" along distal lateral margin of basis .............. 12 I

12A. Merus of P. 1 with tergal spine ..................... Anarthruridae (males - subfamily Akanthophoreinae) I

12B. Merus of P.1 with only setae ......................................................................................................... 13

13A. Last 3 segments of A.1 of similar length .................................................... Typhlotanaidae (males) I

13B. Last 2 segments of A. 1 of similar length ..........................................................................................

........................... Pseudotanaidae (males) and Anarthruridae (males - subfamily Leptognathiinae) I

Key to Genera and Species i

Note: Genera enclosed within brackets are those that have been reported from California, but were not roll

represented in the present study. Species enclosed within brackets indicate males that were not collected. I

1A. With eyes ......................................................................................................................................... 2 1

lB. Without eyes .................................................................................................................................. 12 I

2A. P.1 - P.6 without ischium ................................................................................................................ 3 I

2B. P.1 - P.6 with ischium ..................................................................................................................... 7

m

3A. Pleon with more than 3 pleonites .................................................................................................... 4 I

3B. Pleon with 3 pleonites ...................................................................................................... [Pancolus] I

II

4A. Uro long and slender, each segment more than twice as long as broad ....................... [Synaptotanais]

4B. Uro short, each segment twice as long as broad ............................................................................. 5 I

5A. Pleon with 4 pleonites ..................................................................................................... [Sinelobus] •

5B. Pleon with 5 pleonites (last 2 pleonites distinctly smaller than preceding ones) ............................ 6 |

segment of A. 1 twice as long as second segment .................................................. [Anatanais] I

6A. First

lilt

6B. First segment of A. 1 2.5x (nearly 3x) longer than second segment (genus Zeuxo) ..........................

............................................................................................................................ Zeuxo maledivensis I

|
!

I 7A. A. 1 with more than 3 segments ....................................................................................................... 8

7B. A. 1 with 3 segments ...................................................................................................................... 11

I 8A. A. 1 with more than 4 segments (exclusively males) ....................................................................... 9

8B. A. 1 4-segmented (genus Paratanais) ........................................................... Paratanais intermedius

!

9A. A. 1with more than 7 segments; all mouthparts reduced to lobiform structures; Um endopodite with

more than 2 segments (genus Leptochelia) .......................................................... Leptochelia dubia

I 9B. A.1 with 6-7 segments; mouthparts except Mxp reduced to tobiform structures; Uro endopodite

with 2 segments ............................................................................................................................. 10

I 10A. A.1 7-segmented (genus Pseudotanais, in part) ........................... [Pseudotanais makrothrix, male]

10B A.1 6-segmented (genus Paratanais) .............................................. [Paratanais intermedius, male] !

11A. Endopodite of Um with more than 2 segments; basis of Mxp not fused medially (genus Leptochelia)

I .............................................................................................................................. LeptocheIia dubia 1lB. Endopodite of Um 2-segmented; basis of Mxp fused medially (genus Pseudotanais) ....................

................................................................................................................... Pseudotanais makrothrix

!

12A. A.1 7-segmented, first _md second segments laterally compressed, flagellar segments with many

i 12B. aestheta A.1 3- toscs; 5-segmented, mouthparts,first except and second Mxp, reduced segmentstocylindrical, lobiform structures only 1terminal (exclusively aesthetasc; males)mouthparts ......... 26

not reduced .................................................................................................................................... 13

I 13A. A.1 3- or 4-segmented ................................................................................................................... 14

i 13B. A.1 5-segmented (exclusively subadult males of Chauliopleona, Paraleptognathia, and possibly Scoloura) ....................................................................................................................................... 19

I 14A. A.1 4-segmented ............................................................................................................................ 15 14B. A.1 3-segmented ............................................................................................................................ 24

I 15A. Che articulated to cephalothorax by a "side-piece" along distal lateral margin of basis .............. 16

15B. Che articulated to cephalothorax along proximal margin of basis (genus Siphonolabrum) .............

I ............................................................................................................ SiphonoIabrum caIiforniensis

16A. Uro with exopodite fused to basis (pseudobiramous) ................................................................... 17

I 16B. Uro with exopodite not fused to basis (biramous) ........................................................................ t9

I 17A. Exopodite of Uro represented by a well-developed projection, occasionally reaching length of first endopodal segment (genus Araphura) .......................................................................................... 18

17B. Exopodite of Uro only represented by a tiny, knobtike projection on basis (genus Tanaella) .........

I .......................................................................................................................... Tanaella propinquus

i I 205
!

18A. Pleotelson short, with lateral borders relatively straight, and without knoblike process on posterior I

margin ................................................................................................................. Araphura breviaria

18B. Ple posterior otelsonmargin long, subquadrate, with slightly concave lateral borders, and with knoblike process on I ............................................................................................. Araphura cuspirostris

19A. Pleotelson with large, ventrolateral, triangular process (genus Scoloura) ............ Scofoura phillipsi I

19B. Pleotelson without such a process ................................................................................................ 20

20A. Last pleonite with a large, midventral, caudally-directed spiniform process that reaches more than half I

the length of the pleotelson (genus Chauliopleona) ........................................ Chauliopleona dentata

20B. Last pleonite without such a process ............................................................................................. 21 I

21A. Merus of P. 1 with exclusively setae or unarmed .......................................................................... 22

I

2lB. Merus of P.1 with a tergal spine .......................................................... Paraleptognathia cf. gracilis

22A. Merus of P.1 with 2 long, slxong setae, both as long as carpus ........................................................ I

.............................................................................................................. Paraleptognathia bisetulosa

22B. Merus of P.I unarmed or with only short setae, never as long as carpus ..................................... 23 I

23A. Coxa of P.1 with well-developed, rostrally-directed, triangular process; merus of P. 1unarmed (genus m

Tanaopsis) ............................................................................................................ Tanaopsis cadieni |

23B. Coxa of P.1 without such a process; merus of P. 1 with 1 seta (genus Leptognathia) ......................

............................................................................................................... Leptognathia cf. breviremis !

24A. Pereonites 1 and 2 extremely small, typically both pereonites together as long as third pereonite; m

molaris reduced to various degrees (genus Pseudotanais G.O. Sars, 1882) .............................

pars

............................................................................................................... Pseudotanais californiensis

24B. Pereonites 1 and 2 of normal size, both pereonites together longer than pereonite 3; pars molaris not

reduced, with broad grinding area surrounded by a crenulated wall (genus Typhlotanais) ......... 25 II

im

25A. Lateral borders of pereonites nearly parallel; Uro with exopodite and endopodite indistinctly 2- •

segmented; exopodite only slightly longer than first endopodal segment; sternal border of propodus I

in P.2 with 2 setae of equal size ................................................................. TyphIotanais wiIliamsae am

25B. Lateral borders of pereonites convex; Uro with endo- and exopodite 1-segmented, exopodite slender, I

reaching about 2/3 endopodal length; sternal border ofpropodus in P.2 with 1 short and 1long setae

(exceeding combined length of dactylus and terminal spine) ......................... TyphIotanais crassus I

26A. Che articulated to cephalothorax by a "side-piece" along distal lateral margin of basis; Mxp endite

with short distal seta ...................................................................................................................... 27

Ii 26B. Che articulated to cephalothorax along proximal margin of basis; Mxp endite with long distal seta

(genus SiphonoIabrum) ......................................................... [Siphonolabrum californ&nsis, male]

I
!

i 27A. Merus of P.1 exclusively with seta(e) or unarmed ........................................................................ 30

27B. Merus of P. 1 with at least I tergal spine (subfamily Akanthophoreinae) ..................................... 28

I

28A. Uro with endopodite 3-segmented; caudo-distal part of chelipedal carpus not expanded; P.1 - P.3

with tergal spines of meres and carpus short, reaching only 113length of following segment .... 29

I 28B. Uro with endopodite 2-segmented; caudo-distal part of chelipedal carpus expanded; P.1 - P.3 with

tergal spines of merus and carpus long, reaching about midlength of following segment (probably

i genus Chauliopleona) ................................................................................................. male species 1

29A. Ischium of P.6 with 2 small setae; cephalothorax with anterior part distinctly smaller than posterior

I part; P1with outer margin of exopodite denticulated ................................................ male species 3

29B. Ischium of P.6 with 1 spine and 1 small seta; cephalothorax with anterior part not distinctly smaller

than posterior part; P1 lacking denticles on outer margin of exopodite ..................... male species 2

I

30A. A.1 with segment 3 about twice as long as broad (genus Pse_dotanais) .........................................

I .................................................................................................. [Psemtotanais caIiforniensis, male] 30B. A.1 with segment 3 annular (much shorter than long) (genus Tanaopsis) .......... Tanaopsis coxlieni

I

I Species Descriptions

Suborder Apseudomorpha Sieg, 1980

I Family Apseudidae Leach, 1814

I Genus Carpoapseudes Lang, 1968 Carpoapseudes caraspinosus Dojiri and Sieg, new species

i Figure 3.7

Type Material. Specimen (holotype, with P. 1 and Che missing; USNM) from cruise 3-4, Sta. R-7

(rep. 2), 34°52.90'N 121°10.30'W, 565 m, off Pufisima Point, Santa Maria Basin, California, originally

I as Apseudes graciIis; specimen (paratype, dissected; SBMNH) BSR-28,

identified from Sta. 35°04.22'N

121°19.65'W, 603 m, southwest of Pt. San Luis, Santa Maria Basin, California, originally identified as

Apseudes gracilis.

II

I Description. (Figure 3.7). Body, 5.5 mm, about 5x longer than broad, and with 5 pairs of oostegites

(cheliped with 1 pair of reduced oostegites). Cephalothorax trapezoid in general outline; midlaterally separated

I by indentation and 1 small thorn; anterior half also equally rounded; rostrum large, spear-shaped; eyelobes large, pointed, without visual elements. Pereonites 1 and 2 lacking spines, very often with welt-developed

coxal spines of P.1 and P.2 mistaken for pereonal spines), 3.5-4.0x broader than long; pereonites 4-6 with

i posteriorly expanded protrusions atinsertion of pereopods; coxa with small thorn (often mistaken for pereonal

spines); pereonite 3 about 1.5x broader than long, with long anteriorly, slightly curved thorns; pereonites 4

and 5 about 1.25x longer than broad, with laterally-directed thorns; pereonite 6 also 1.5x broader than long,

i laterally_ thorn small. All 5 pleonites of equal size, withwell-developedpleurites,2.5× broader than long.

i 207
!

First antenna with of 4-segmented peduncle, 8-segmented inner flagellum, and about 20-segmented I

outer flagellum; peduncle 1about twice as long asremaining peduncular segments combined; outer flagellum

with about 5 aesthetascs. II

Mouthparts of typical apseudoidean shape. Maxillipeds not fused; coxa short, no seta; basis with i

lateral margin expanded, no setae; palpus 4-segmented, with inner palpal margin with 1 long strong seta,

outer margin with 1 short, but strong seta; segments 2-4 bearing typical setation; endite with specialized 1

spine welldeveloped. II

Cheliped slender; ischium absent; semicircular coxa of adult females with reduced oostegite; basis

2.5x longer than broad, exopodite welldeveloped, 3-segmented, bearing 4 long pinnate setae terminally,

and group of distal setae on sternal margin; merus bent tergally, with sternal margin covered with setae;

carpus 2.5x longer than broad, with 2 short distal setae on tergal margin and 3 long and 4 short setae along

sternal margin; propodus combined with fixed finger about twice as long as broad, bearing 6 setae sternally, HI

spiniform element at its tip, 1 row of membranous structures (chemoreceptors?) along tergal margin, and 2

caudal setae near articulation of dactylus; dactylus with spine curved caudally, carrying 2 long and 1 short

setae. I

Pereopod 1 slender, non-fossorial; coxa with large anteriorly-directed projection; basis 4.8x as long

as broad, no setae; exopodite 2-segmented, with 6 pinnate setae; ischium small; merus as long as carpus or

slightly longer, 2.5x longer than broad, with 3 distal setae on tergal border, 1 row of setae along sternal I

margin, and 1 distal pinnate spine; carpus 3.7x longer than broad, with 1 row of setae along sternal and g

tergal margins and 2 pinnate spines distally; propodus short, 1.4x longer than broad, with row of setae along

sternal and tergal margins, 1 row of 5 pinnate spines distally, 1 short specialized spine near articulation of I

dactylus; dactylus half as long as propodus, bearing sternal row of setae (each setal loci consisting of 2

II

setae) and 3 tergal setae; short terminal (dactylar) spine not fused to dactylus; terminal spine with 2 small

setae at base on sternal margin. I

II Pleopods 1-5 of similar shape, biramous; coxa fused with sternite; basis elongate; exo- and endopodite

1-segmented; inner and outer margins covered with pinnate setae.

Pleotelson tong, 2.6x longer than broad. I

Uropods only partially known; basis 2.75x longer than broad, with inner margin equipped with

several terminal setae; endopodite missing; exopodite 15- to 17-segmented, with 3 long setae and 1 short •

seta. |

Remarks. Carpoapseudes caraspinosus represent the only species of the suborder Apseudomorpha

Sieg, 1980 collected during the MMS Soft-Bottom Benthic Fauna of the Santa Maria Basin and Western I

Santa Barbara Channel study.

Carpoapseudes and Leviapseudes, both belonging to the subfamily Leviapseudinae within the

Whitelegiidae, are closely related to each other. However, both genera are easily separated by the shape of I

the pleonites. In Carpoapseudes the pleonites are broader than long, while those ofLeviapseudes are triangular

and typically longer than broad. Additionally, in Carpoapseudes the pleonites show well-developed pleurites,

while these are small in Leviapseudes. In Carpoapseudes the carpus is about 2.5x longer than broad, but in In

Leviapseudes it is almost 4.5x longer than broad.

Carpoapseudes caraspinosus may be distinguished from all its congeners by a combination of

characters. So far, all known members of Carpoapseudes lack a lateral thorn anterior to the carapace chamber,

which is characteristic for species of the genus Leviapseudes, while the new species is the only species roll

within its genus that possesses such a small thorn. Carpoapseudes auritocheles Kudinova-Pasternak, 1975

and C. caraspinosus have anterolateral thorns on pereonite 6. But, these species may be distinguished by the

shape of pereonite 2: C. caraspinosus lacks anterolateral thorns, while C. auritocheles has well-developed I

ones.

!

208 I

!

Finally, the shape of the specialized propodal spine close to the articulation of the dactylus on P.1 I

may be characteristic. However, as in many other cases, morphological details are not always examined and

reported. Because of these shortcomings, the present taxonomic value of these morphological characters is I

reduced and do not allow detailed morphological comparisons to be made among the species of this genus. |

Etymology. The specific name caraspinosus is a combination of the Greek words cara, meaning

head or top, and spinosus, thorny, and alludes to all the spines on the head and body. 1

J Type Locality and Type Specimens. Santa Mafia Basin, California. Types deposited in the National

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

"Type Material." I

Distribution. This species is so far known only from its type locality.

Ill

Suborder Tanaidomorpha Sieg, 1980 I

Family Tanaidae Dana, 1849 I

Genus Zeuxo Templeton, 1840

Zeuxo maledivensis Sieg, 1980 I

Figure 3.8

Ill

Material Examined. 1neuter (dissected) from Sta. 53, 34°37.96'N 120°50.38'W, 196 m, northwest I

of Pt. Axguello, Santa Maria Basin, California; 2 neuters from Sta.56, 34030.32'N 121°01.02'W, 900 m,

southwest of Pt. Arguello, Santa Maria Basin, California; 1 male (dissected) from Sta. BRA-6, 35°20.88'N I

120°59.62'W, 109 m, between Pt. Estero and Pt. Buchon, Santa Maria Basin, California. I

Description (male) (Figure 3.8). Body 3.8x longer than broad. Cephalothorax about as long as

broad, tapered at anterior end. Pereonite 1 shortest of pereonal segments; pereonites 1-5 similar in width; •

pereonites 4 and 5 longest and pereonite 6 narrowest of all pereonites. Pleonites 1-5 divisible into 2 groups: II

first group consists of pleonites 1-3, with each pleonite about 4x broader than long, bearing pleopod; second

group composed of pleonites 4 and 5, with each pleonite about 10x broader than long, lacking pleopod. R

First antenna 3-segmented, bearing numerous aesthetascs terminally.

Mandible with broad, fiat, crushing pars molaris; 8 hairs (setules) situated at insertion of tacinia

mobilis. I

Maxilliped with small coxa, each bearing 1 naked seta; basis not fused to opposite member, each

bearing single distal seta. Distal margin of each endite with specialized seta, scattered setules, and several I

curved rows of tiny spinules. Palpus 4-segmented; first segment with lateral seta and a few rows of minute I[

spinules; second segment with approximately 8 medial and 1 lateral setae; third segment with numerous

medial setae, many pinnate at tips; inner margin of fourth segment with 7 setae (5 pinnate, 2 naked); outer

margin with 2 pinnate setae terminally, 1 naked seta medially. !

Cheliped with sternal protuberance articulating with side-piece. Meres small, with small protuberance

on sternal distal margin, bearing 2 sternal setae. Carpus stout, about as long as broad, expanded at tergal 1

proximal comer, bearing 1 tergal seta at midlength, 2 distal tergal setae, small rounded sternal protuberance, I

and 4 sternal setae. Fixed finger broad and bluntly rounded terminally, tipped with small stout spine; tergal

and sternal borders with numerous setae. Dactylus with subterminal seta and numerous small, stout spinules m

on inner (sternal) margin. I

I

210 I
I

Figure 3.8. Zeuxo maledivensis Sieg, 1980 male: C_, general habitus of male; Md, mandible; Mxp, maxilliped;

m _general Che, habit chelipe u d; P.1, pereopod 1; P.6, pereopod 6; PI, pleopod; Uro 0 _, uropod of male; neuter, s of neuter; Uro, uropod of neuter.

i

m 211
!

Pereopod 1 with coxa bearing a slight protuberance on sternal distal comer near 2 naked setae; basis I

with proximal hair and setae and sternal distal seta; ischium absent; merus with small tergal distal seta;

carpus with 1 long sternal seta; propodus with 4 setae; dactylus slender, with 1 setule; terminal spine not II

fused to dactylus. Pereopod 6 with carpus bearing 2 naked setae, 3 stout bifid spines, and 2 stout spines; l

propodus with 2 sternal setae, 3 terminal setae, 1terminal hair, and several spatulate spines; dactylus clawlike,

armed with row of translucent spinules on each side. i

Pleopods 1-3 with basis carrying 3 outer pinnate setae and 1 inner pinnate seta; exopodite with n

numerous outer pinnate setae; endopodite with several outer pinnate setae and 1 inner pinnate seta.

Pleotelson, 2.3x broader than long, with irregular border, i

Uropod unlramous; basis with 5 setae. Exopodite absent. Endopodite 4-segmented; fin-stsegment

with 1 feathered hair; second segment with 3 setae; third segment with 3 setae; fourth segment with 7 setae

and 1 feathered hair. i

u

l)eseription (neuter) (Figure 3.8). Body about 4x longer than broad. Cephalothorax about as long

as broad. Pereonites tapering posteriorly. Pleonites 1-3 expanded slightly relative to preceding pereonites 5

and 6. i

Uropod with endopod only 3-segmented.

Remarks. Zeuxo maledivensis Sieg, 1980 is recorded for the first time in California waters. It had i

previously been recorded only from the Maldives, Japan, and Florida, but it's occurrence in California was

predicted by Sieg and Wirm (1981:341). So far, a total of eight species belonging to the family Tanaidae

have been recorded from the California coast: Synaptotanais notabilis Sieg and Winn, 1981, Sinelobus n

stanfordi (Richardson, 1901), Pancolus caIiforniensis Richardson, 1905, Anatanais pseudonormani Sieg

and Winn, 1981, Zeuxo normani (Richardson, 1901), Z paranormani Sieg, 1980, and now Z. maledivensis

Sieg, 1980. i

Sinelobus stanfordi and Pancolus californiensis are easily recognized by having 4 and 3 pleonites,

respectively. Synaptotanais notabilis also has 5 pleonites visible dorsally. But, this species differs from

Zeuxo in having the two reduced pleonites still f_ee and not fused sternally to the pleotelson. Finally, the

totally different shape of the uropodal segments makes the distinction between Synaptotanais notabilis and U

Zeuxo species quite easy. Anatanais pseudonormani may be recognized by the relatively elongate

cephalothorax and the large triangular protrusion of the coxa in pereopod 1. i

Within Zeuxo, the newly recorded species Z. maIedivensis is characterized by having a slightly

reduced lacinla mobilis on the right mandible. This is also true for Z. normani and Z. paranormani, but Z.

maledivensis can be distinguished from these two congeners by the pleopodal endopodite which bears only i

1 seta, instead of 2 or 3, on the inner border. Finally, the coxal protrusion on pereopod 1 in Z. maledivensis i

is only slightly developed, while it is distinctly larger in Z. normani and Z. paranormani.

Juveniles (neuters) of Z. maledivensis have only 4-segmented uropods, as does Z. coralensis Sieg, i

1980, another species predicted by Sieg and Winn (1981) to inhabit California waters. It can be distinguished

n

from adults of this species (juveniles ofZ. coraIensis have a 3-segmented uropod) by the shape of the coxal m

protrusion in pereopod 1 and additionally by the armament of the second segment of the maxillipedal i

palpus. In Z. maledivensis there are only 2 relatively long setae, while adults ofZ. coralensis have mostly 4

u

(seldomly 3) setae in addition to the 3 pitose spines. i

Type Locality and Type Specimens. Holotype and paratypes deposited in Senckenberg Museum/ i

Frankfurt (SMF 8690 and SMF 8691), Zoologisches Museum Berlin (ZMB 15951) and National Museum

of Natural History (USNM).

Distribution. Zeuxo maledivensis Sieg, 1980 is known from scattered records from the Maldives, i

Japan, Florida, and the Santa Maria Basin, California.

!

212 i
!

I Family Leptocheliidae Lang, 1973

I Genus Leptochelia Dana, 1849

Leptoehelia dubia (Krcyer, 1842)

I Figures 3.9, 3.10

Material Examined. Santa Maria Basin, phase I: 1 male from Sta. 6, 35°20.88'N 120°59.62'W,

I 190 between Pt. Estero and Pt. Buchon, October 1984; 1 female from Sta. BRA-16, 35°12.23'N

m,

121°16.29'W, 591 m, off Pt. Buchon, March 1985; 1 female and I neuter from Sta.-73, 34°28.21'N

120°36.80'W, 98 m, between Pt. Conception and Pt. Arguello, February 1985. Santa Mafia Basin, phase II:

I 1 neuter from cruise 1-3, Sta. R-4 (rep. 1), 34°43.01'N 120°47.39'W, 93 m, off Purisima Pt., May 1987; 1

neuter from cruise 3-4, Sta. R-4 (rep. 3), 34°43.01'N 120°47.39'W, 93 m, off Purisima Pt., May 1989; 1

manca-I from cruise 2-4, Sta. R-4 (rep. 2), 34°43.01'N 120°47.39%V, 93 m, off Purisima Pt.; 1 female from

I c

c

ruruise ise 3-4, 1-3, Sta. Sta. R-8 R-4 (rep. (rep. 1), 2), 34°55.30'N 34°43.01'N 120°45.87'W, 120°47.39'W, 9093m,m,offoffPt.Purisima Sal, MayPt.,1987. May I989; 1 female from

From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring

I Division, of Los 10 females, 1 from Sta. B8, 33°53'48"N 118°28'27"W, 45 11 City Angeles: neuter m,

January 1990; 7 females from Sta. D4, 33°51'07"N 118°31'30"W, 80 m, 11 January 1990; 1 female and 1

male from Sta. C4(2), 33°58'17"N 118°34'00"W, 60 m, 12 July 1990; 3 females and 1 male from Sta. B3,

I 34°00'25"N 118°35'46"W, 45 09 1990; 4 females, 1 male, and 8 neuters neuter from

m, July (1 dissected)

Sta. D2, 33°53'40"N 118°35'20"W, 79 m, 08 January 1990; 1male from Sta. 4C, 33°50'40"N 118°26'22"W,

77 m, 24 July 1986; 3 females and 1 male from Sta. C1(2), 33°59'49"N 118°42'50"W, 60 m, 09 July 1990;

I five females and 3 males (1 female and 1 male dissected) from Sta. B8, 33°53'48"N 118°28'27"W, 45 19

m,

July 1990; 4 females and 2 males from Sta. D4, 33°51'07"N 118°31'30"W, 80 m, 03 September 1987.

i From Los Angeles County Museum of Natural History: 3 females, 1 male, and 2 neuters (LACM

59-147), 205 m, rocks, 2.45 mi, 249.5 ° T (true north) from Gull Island, Santa Cruz Channel, Santa Barbara

County, California, 33°56.10'N 119°52.28'W, R/V Velero IV(AHF 6806-59), 22 December 1959; 3 females

(LACM 59-147), 205 m, rocks, 2.45 mi, 249.5 ° T from Gull Island, Santa Cruz Channel, Santa Barbara

I County, California, 119°52.28'W, /V (AHF 6806-59), 1959;

33°56.10'N R/V Velero 22 December 6 females

(LACM 59-149), 201 m, rocks, 2.3 mi, 246.5 ° T from Gull Island, Santa Cruz Channel, off south side of

Santa Cruz Island, California Channel Islands, California, 33°56.05'N 119°52.05%V, R/V Velero IV (AHF

I 6805-59), 22 December 1959; 1 female (LACM 59-161), 86 sand, 2.4 rni, 248 ° T from Kinton Point,

m,

Santa Cruz Island, California Channel Islands, California, 33°59.53'N 119°55.92%V, R/V Velero IV (AHF

6803-59), 22 December 1959; 1 female and 1 neuter (LACM 55-35), 73 m, mud, 9.9 mi, 203 ° T from Santa

Monica Pier Light, Los Angeles County, California, 33°51.37'N 118°34.7'W, R/V Velero IV (AHF 2998-

55), 06 February 1955; 1 female (LACM 58-60), 35 m, green mud, 2.1 mi, 231 ° T from Santa Monica Pier

Light, Los Angeles County, California, 33°59.2'N 118°32.0'W, R/V Velero IV(AHF 5732-58), 15 May

I 1958; 2 females (LACM 55-67), 119 m, sand and mud, 7.4 mi, 229 ° T from end of E1Segundo Pier, Los Angeles County, California, 33°50.00'N 118°32.38%V, R/V Velero IV (AHF 3385-55), 23 August 1955; 1

female (LACM 53-125), 229 m, sand, 6.4 mi, 239° T, end of Redondo Beach Pier, Los Angeles County,

I California, 33°47.05'N 118°30.12'W, R/V Velero 1V (AHF 2789-53), 08 July 1953; 2 females (LACM 54- 38), 11 m, mud with stones, 4.1 mi, 331 ° T from Palos Verdes Point, Los Angeles County, California,

33°50.00'N 118°28.00'W, R/V Velero IV (AHF 2725-54), 08 May 1954; 1 female (LACM 53-128), 183 m,

I sand, V Velero 3.4 mi 301 ° T from Pt. Vicente Light, Los Angeles County, California, 33°44.13'N 118°25.80'W, R/

IV(AHF 2357-53), 08 July 1953; 1 female and 1 male (LACM 57-102), 20 m, coarse black sand,

3.35 mi, 112° T from Pt. Vicente Light, Los Angeles County, California, 33°43.30'N 118°20.88%V, R/V

I

I 213
!

Velero IV (AHF 5102-57), 24 May 1957; 2 females (LACM 60-100), 219 m, 4.7 mi, 349.5" T from Pt. i

Fermin Light to midpoint, Los Angeles County, California, 33°37.80'N 118°16.73'W, R/V Velero IV (AHF

7161-60), 08 October 1960; 1 male (LACM 60-101), 214 m, mud, 3.8 mi, 163.5 ° T from Pt. Fermin Light, •

Los Angeles County, California, 33°38.60'N 118°16.27'W, R/V Velero IV (AHF 7174-60), 09 October l

1960; 2 females and 1male (LACM 53-126), 75 m, sand, 6.3 mi, 184" T from L.A. Breakwater Lighthouse,

Los Angeles County, California, 33°36.23'N 118°15.50_V, R/V VeIero IV (AHF 2355-53), 11 July 1953; 8 •

females and 1fragment (LACM 52-51), 88 m, 9.2 mi, 156.5 ° T from L.A. Lighthouse, Los Angeles County, I

California, 33°34.08'N 118°10.68'W, R/V Velero IV (AHF 2126-52), 25 June 1952; 1 female (LACM 60-

108), 16 m, silt, 0.2 mi, 254.5 ° T from base of Newport Pier, Newport Canyon, Orange County, California, •

33°36.40'N 117°55.90'W, R/V Velero IV(AHF 7031-60), 05 May 1960; 1 female (LACM 60-184), 139 m, |

sand, 45 mi, 037.5 ° T from Point La JoUa, San Diego County, California, 32°52.38'N 117°15.5'W, R/V

Velero IV(AHF 7043-60), 07 May 1960; 2 females and 1 male (LACM 60-185), 84 In, sand, 1.4 mi, 027.5 ° i

T from Point La Jolla, San Diego County, California, 32°52.35'N 117°15.45'W, R/V Velero IV (AHF 7044- i

60), 07 May 1960.

Description (female) (Figure 3.9). Body, 3-5 mm long, about 7x longer than broad. Cephalothorax, I

1.4x longer than broad, with curved lateral borders and tapered anterior end. Pereonite 1 shortest of pereonites; II

pereonites 2-6 quadrangular with relatively straight, parallel, lateral borders; pereonite 4 longest; pereonite

5 almost as long as broad. Pleonites 1 and 5 narrowest of pleonites and pleonites 2-4 broadest; each pteonite •

about 5x broader than long. l

First antenna 3-segmented; first segment long and slender, 3.5x longer than broad; third segment

with 2 subterminal setae and slight constriction giving appearance of a fourth segment; formula for armature: •

4 setae + 7 feathered hairs, 2 setae + 1 feathered hair, 4 setae + 1 aesthetasc.

Second antenna 6-segmented with armature formula as follows: 1 small seta, 2 stout spines, 1 stout

spine, 3 setae + 3 feathered hairs, 2 setae, and 5 setae. I

i

Mandible with pars motaris broad; crushing area well-developed; 113 of surrounding wall

denticulated.

Maxilliped without coxae; basis not fused medially and bearing 5 setae near insertion of palpus, i

Endites each lobiform with 3 hyaline bulbs and 1 seta. Palpus 4-segmented; distal margin of first segment

slanting toward inner border, unarmed; second segment trapezoidal, inner border longer than outer, outer I

border with 1 seta, inner border with 4 setae and row of small setules; third segment longest, with 7 pinnate |

setae and 2 naked setae; fourth segment with 9 pinnate setae and 1 naked seta.

Cheliped carpus slender, 2.5x longer than broad, with 2 tergal setae and 3 sternal setae. Palm "comb" •

with 3 small pinnate setae and 1naked seta; 3 rows of minute spinules situated near comb. Fixed fmger with l

spiniform tip and 5 indentations; tergal border with 3 setae and 1seta (dotted seta in figure) near insertion of

dactylus; sternal border with 4 setae. Dactylus with 1 proximal seta. •

Pereopod 1 with merus beating 2 tergat setae and 1 sternal seta; carpus with 5 setae; propodus with l

1tergal and 3 sternal setae; dactylus and terminal spine longer than propodus; dactylus with 1 seta. Pereopod

6 with carpus bearing 2 setae and 2 spines; propodus with 2 tergal spines and 1 long and 4 shorter pinnate •

setae; dactylus and terminal spine forming a stout curved claw.

Pleotelson, 2.5x broader than long, with protuberant posteromedian area.

i

Uropod biramous. Exopodite 1-segmented, 2.8x longer than broad, with 3 setae. Endopodite 5- !

segmented; first segment 1.5x longer than broad, with 1 small distal seta; second segment 2.8x longer than an

broad, with 1 feathered hair; third segment 3x longer than broad, with 1 distal seta; fourth segment 4x i

longer than broad with 1 long seta and 1 feathered hair; fifth segment about 5.7x longer than broad, with I I

subterminal and 4 tong terminal setae.

i

!

214 i
I

Figure 3.9. Leptochetia dubia (Krc_yer, 1842) female: 9, general habit-as of female; A.1, first antenna; A.2,

second antenna; Md, mandible; Mxp, maxilliped; Che, cheliped; P.1, pereopod 1; P.6, pereopod 6;

I Uro, uropod.

I

I

i 215
I

Figure 3.10, Leptochelia dubia (Kr_yer, 1842) male: C_, general habitus of male; A.1, first antenna; C_1, flagellar I

segments of first antenna (variant); A.2, second antenna; Mxp, maxilliped; Che, cheSped; P.1, pereopod 1

1; P.6, pereopod 6; Uro, uropod.

I

I

I

2,6 I
!

I Description (male) (Figure 3.10). Body, 2 - 3.5 mm long, 6x longer than broad. Cephalothorax

1.3x longer than broad, with narrow anterior end andbroad posterior end. Pereonite 1 shortest of pereonites

and pereonite 4 longest; all pereonites similar in width. Pleonites 1-5 slightly broader than pereonites, each

I about 5-5.5x broader than long.

First antenna with 3-segmented peduncle and 5- or 6-segmented flagellum; first segment long,

I sbroad; lender,remaining about 4.5xsegments longer than comparatively broad; secondshort;segment last one3.2xconsiderably longer thansmaller broad;than thirdpresegment ceding 2xones. longer Fomula than

for armature as follows: 1 seta + 2 feathered hairs, 2 setae + 2 feathered hairs, 2 setae, 3 aesthetascs, 3

I aesthetascs, 4 aesthetascs, 4 aesthetascs, 3 aesthetascs, 4 setae + 1 aesthetasc. Second antenna 6-segmented; first segment semicircular in cross-section; second and third segments

short, about as long as broad; segment 4 long and slender, 6.7x longer than broad; segment 5 about 5.7x

I lo1nger than broad; segment 6 minute, as long as broad. Armature formula as follows: 1seta, 1 seta + 1 spine, spine, 2 setae + 3 feathered hairs, 1 seta (?), and 5 setae (?).

Maxilliped sexually dimorphic; basis reduced to unarmed plates; palpus reduced to rounded lobes

I without setae.

Cheliped carpus slender, 2.4x longer than broad. Palm "comb" with numerous spinules. Fixed

finger curved with 1 proximal tergal process and 1 tergal process near midlength; 1 tergal seta located

I between and insertion of and 3 setae at midlength; sternal border

proximal process dactylus near process

with 3 setac. Dactylus slender, curved, with 1 proximal tergal seta and several setules along sternal margin.

i with Pereopod 1 more slender than in female; merus with 1 tergal seta; carpus with 6 setae; propodus

5 setae; dactylus and terminal spine as long as propodus; dactylus with 1 seta. Pereopod 6 carpus with

stout spines and 2 sternal setae; propodus with 2 tergal spines and 1 long and 7 shorter pinnate setae.

I Pleotelson Uropod about 2.2x broader than long. biramous with basis bearing 2 setae. Exopodite 1-segmented, 3x longer than broad, with 3

setae. Endopodite 5-segmented, with armature as follows: 1 feathered hair + 1 setule, 2 setae + 1 feathered

I hair, 1 seta, 2 setae + 1 feathered hair, 5 setae + 1 feathered hair. Remarks. This species can be distinguished from all other southern California tanaidaceans by the

5-segmented endopodite of the uropods in both the adult female and male. The endopodite of the neuter of

1 this species is 3-segrnented (Figure 3.10). Several characters in combination also are helpful in the diagnosis 1

of L. dubia: (1) the spines on the second and third segments of the second antenna of the female, (2) the

structure of the pars molaris of the mandible, (3) the unfused maxillipedal basis, (4) 3-5 setae on the distal

I end of the maxillipedal basis, (5) the armature of the maxillipedal endite, and (6) 2 setae on the merus of pereopod 1. The males can be distinguished from the females by the large chelipeds and the handcuff-like

propodusldactylus combination. Some males with a different number of flagellar segments (Figure 3.10,

I O_1) were present in the collection.

This species inhabits a wide range of substrates from rocks and sand to mud and silt.

i Typ DistriebuLotioncalit. y Leptochelia and Type Sdubia peciesoccurs . See KrCger in tropical (1842). and subtropical shallow waters throughout the

world; it is known from Santa Maria Basin, California south to La Jolla, San Diego County, California.

I

!

!

i 217
I

Family Paratanaidae Lang, 1949 I

Genus Paratanais Dana, 1849 1

i

Paratanais intermedius Dojiri and Sieg, new species

I

Figure 3.11 I

Type Material. 1 female hototype (USNM) from Sta. BRA-20, 35°15.72'N 121°04.68_W, 396 m, m

off Pt. Buchon, Santa Maria Basin, California, June 1985, originally identified as Paratanaidae sp. A; I 1

female paratype (SBMNH)(dissected) from Sta. BRC-1, 35°27.86'N 121005.33%V, 98 m, off Pt. Estero, i

Santa Mafia Basin, California, June 1985, originally identified as Paratanaid_e sp. A.; 1 manca-ll (paratype; i

SBMNH) from Sta. BRA- 1, 35°12.23'N 121°16.29'W, 591 m, southwest of Pt. Buchon, Santa Mafia Basin, 1

California, originally identified as Tanaidacea sp. A; 1 manca-II (paratype; USNM) from Sta. 6, 35°20.88'N i

120°59.62'W, 109 m, between Pt. Estero and Pt. Buchon, Santa Mafia Basin, California, October 1984,

originally identified as Tanaidacea sp. A. H

i

II

Description (female) (Figure 3.11). Body, 2.16 mm long, about 6x longer than broad. Cephalothorax,

1.2x longer than broad, with posterior margin broader than anterior. Pereonites 1 and 6 similar in length; iiim

pereonite 1 broadest; pereonite 4 longest; all pereonites with convex lateral borders. Pleonites 1-5 similar 1

lengths and widths; 4.4x broader than long.

First antenna 4-segmented; first segment about 2.5x longer than broad; second segment as long as •

broad; third segment smallest, annular; fourth segment 2.6x longer than broad; formula for armature: 1 seta 1

+ 5 feathered hairs, 1 seta + 4 feathered hairs, 1 seta, and 4 setae + 1 aesthetasc + 1 feathered hair.

Second antenna 6-segmented; segment 1 small and unarmed; armature formula of segments 2-6 as •

follows: 2 setae, 1 stout spine, 1 seta + 5 feathered hairs, 1 seta, and 5 setae.

Mandible with broad pars molafis; crushing area well-developed with serrations.

m

Maxilliped without coxae; basis fused to about midlength, bearing 1 distal seta near insertion of 1

palpus. Endites each with 2 hyaline bulbs and 1 medial seta. Palpus 4-segmented; first segment stout and

i

unarmed; second segment with 1 naked seta on outer border and 1 naked seta, 1 pinnate stout seta, and 1 m

denticulated spine on inner border; third segment with 3 pinnate setae and 1 naked seta; and fourth segment I

with 5 pinnate setae and 1 naked seta.

ml

Cheliped carpus stout, 1.3x longer than broad (length measured along tergal margin), with 2 small 1

tergal setae (1 proximal, another distal) and 2 sternal setae. Fixed finger with spiniform tip and bilobed !

protuberance (indicated by arrow in figure); tergal border with 3 setae and 1 seta (dotted seta in figure) near

I

insertion of dactylus; sternal border with 1seta. Dactylus stout with 1small tergal seta and 1 sternal spiniform

element. I

Pereopod 1 with merus almost as long as carpus, bearing 1 tergal seta; carpus with 1 tergal seta and

2 sternal setae; propodus with 3 sternal setae; dactylus and terminal spine longer than propodus; dactylus •

with 1 seta. Pereopod 6 with carpus beating 2 stout tergal spines, 1 small sternal seta, and 1 large clawlike 1

spine; propodus with 2 stout tergal spines, 3 sternal pinnate spines, and 1 sternal feathered hair; dactylus

with 2 minute sternal setules, and several curved rows of minute spinules; terminal spine relatively stout •

and curved. I

Pleotelson, 2.5x broader than long, with rounded posterior margin.

Uropod biramous. Exopodite 1-segmented, 3.7x longer than broad, with 1 subterminal and 2 terminal I

setae. Endopodite 1-segmented with constriction at about midlength, approximately 2.4x longer than broad,

and bearing 2 feathered hairs near constriction, and 5 setae at tip.

I

218 I
I

I

i •_ Che

I / _,_.

/

l

| J "

l Md

| v __

I _ "_ Uro

I _

I _

P.6

I

,

!

i Figure 3.11. Paratanais intermedius Dojiri and Sieg, new species, female: _, general habitus of female; A. 1, first antenna; A.2, second antenna; Md, mandible; Mxp, maxilliped; Che, cheliped; P.1, pereopod 1;

P.6, pereopod 6; Uro, uropod.

I

I

i 219
!

Remarks. The genus Paratanais is readily identified by a combination of the following characters: I

a broad pars molaris, a pair of eyes, strong spine on the third segment of the second antenna, and a 2-

segmented uropodal endopodite, i

Paratanais intermedius n.sp. appears to be morphologically related to P. impressus Kussakin and l

Tzareva and P. spinanotandus Sieg, but can be distinguished from these two species by the length of the

merus relative to the carpus (about equal in length in P. intermedius) and by the serrated stout spine on the •

second palpal segment of the maxilliped.

This species was identified as Paratanaidae sp. A and Tanaidacea sp. A in the MMS voucher

collection, i

Etymology. The specific name intermedius alludes to the species morphological similarities to two

species, Paratanais impressus and P. spinanotandus.

i

Type Locality and Type Specimens. Santa Maria Basin, California. Types deposited in the National 1

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

"Type Material."

Distribution. So far known only from its type locality, Santa Maria Basin, California. 1

Family Anarthruridae Lang, 1971 i

Subfamily Anarthrurinae Lang, 1971 I

Genus Siphonolabrum Lang, 1972

I

Siphonolabrum californiensis Dojiri and Sieg, new species

Figures 3.12, 3.13 I

WID

Type Material. 1 female (holotype; USNM), 2 paratypes (2 females; 1dissected; SBMNH), and 1

female (dissected) from Sta. 079-BSR-01-TX, 34°24.12'N 120°28.32'W, 98 m, off Pt. Conception, Western •

Santa Barbara Channel, California, originally identified as Leptognathia sp. F. 1

Material Examined. Santa Maria Basin and Western Santa Barbara Channel, phase I: 1 male from

Sta. 79, 34°24.12'N 120°28.32'W, 98 m, off Pt. Conception, 09 November 1983, originally identified as 1

Cryptocope sp. D; 2 neuters and 2 fragments from Sta. 39, 34°49.53'N 120°56.85aN, 294 m, off Pt. Sal,

October 1984, originally identified as Leptognathia sp. F; 1 neuter from Sta. 59, 34°33.65'N 120°47.18'W,

216 m, off Pt. Arguello, October 1984, originally identified as Leptognathia sp. D. Santa Maria Basin, i

phase II: 4 neuters and 1 fragment from cruise 1-1, Sta. PJ-17 (rep.2), 34°56.56'N 120°48.98'W, 126 m,

originally identified asAraphura sp. C; 4 neuters from cruise 2-3, Sta. R-4 (rep.1), 34°43.01'N 120°47.39'W,

92 m, off Purisima Pt., originally identified as Araphura sp. C; 7 neuters and 1 fragment from cruise 2-4, •

Sta. R°2 (rep. 1), 35°05.50'N 120°53.40'W, 161 m, offPt. San Luis; 5 females from cruise 1-2, Sta. PJ-23

(rep.1), 34°56.33'N 120°49.90%V, 143 m, originally identified asAraphura sp. C; 6 females, 1 neuter, and 1

manca-II from cruise 1-1, Sta. PJ-7 (rep. 1), 34°55.79'N 120°48.60'W, 123 m, originally identified as •

Leptognathia sp. F; 11 females and 2 manca-II from cruise 1-1, Sta. PJ-1 (rep. 1), 34°55.79'N 120°49.91'W,

145 m, originally identified as Leptognathia sp. F; 11 females and 3 manca-lII from cruise 1-1, Sta. R-2

(rep. 3), 35°05.30'N 121°00.90'W, 409 m, off Pt. San Luis, originally identified as Leptognathia sp. F; 4 II

females and 1 manca-II from cruise 1-1, Sta. PJ-2 (rep. 2), originally identified as Leptognathia sp. F; 9 1

females, 1 neuter, and 1 manca-II from cruise 1-3, Sta. PJ-8 (rep. 3), 34°56.87'N 120°49.91a_V, 142 m,

originally identified asAraphura sp. C; 8 females from cruise 1-3, Sta. R-4 (rep. 2), 34°43.01'N 120°47.39'W, I

220 I
!

I 92 m, off Purisima Pt., originally identified asAraphura sp. C; 7 females and 2 neuters from cruise 1-3, Sta.

PJ-7 (rep. 3), 34°55.79_1 120°48.60'W, 123 m, originally identified as Araphura sp. C; 5 females and 3

i manca-l identified/fromas Leptognathia cruise 1-1, Sta. sp. F;R-69 females (rep. 3), and 34°41.40'N 3 neuters120°57.90_V, from cruise 410 1-3, m, Sta.offPJ-1Purisima (rep. 2),Pt.,34°55.79'N originally

120049.91'W, 145 m, originally identified asAraphura sp. C; 10 females from cruise 1-3, Sta. R-1 (rep. 3),

i 35°05.83'N neuter, and 3120°49.16_V, manca-II from 91cruise m, off 1-3,Pt.Sta.SanPJ-23Luis, (rep.originally 2), 34°56.33_ identified 120°49.90 as Araphura N_r, 143 m, sp.Araphura C; 6 females, sp. C. 1

From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring

I Division, January 1989. City of Los Angeles: 1 fragment (neuter) from Sta. E2, 33°58'39"N 118°39'16"W, 150 m, 05

Deseriptlon (female) (Figure 3.12). Body, 1.74-2.32 mm long, about 7x longer than broad.

I Cephalothorax, all pereonites 1.2x longer than broad, with lateral margins convex. Pereonite 1 shortest ofpereonal segments; similar in width. Pleonites 1-5 similar in lengths and widths, approximately 3-3.7x wider than

long.

I First antenna 4-segmented; first segment about 2.4x longer than broad; second segment 1.3x longer than broad; third segment smallest, as long as broad; fourth segment 3.3x longer than broad; formula for

armature: 1 seta + 4 feathered hairs, 2 setae + 3 feathered hairs, 2 setae, and 6 setae + 1 aesthetasc + 1

I feathered hair.

Labrum with large patch of setules.

i Mandible Maxillipedwithout withoutparscoxa; molaris; basis pars incisiva tipped with several teeth. fused. Distal margin of endites each with 1 long seta. Palpus 4-

segmented; first segment stout and unarmed; second segment with 3 medial setae; third segment with 4

I setae; fourth segment with 4 inner and 1 small outer setae. Cheliped with basis lacking enlarged protuberance (projection); proximal basal margin articulating

with ventral surface of cephalothorax. Merus slightly enlarged and beating protuberance on sternal surface

I immediately distal to single seta. Carpus stout, about 1.3x longer than broad, bearing I proximal tergal seta, 1 distal tergal seta, and 2 proximal sternal setae. Fixed finger with spiniform tip, bicuspid tooth (indicated

by arrow in figure) near tip, and 1 single tooth near midlength of f'mger; tergal border with 3 setae near

I midlength midlength. and 6 setae near insertion of dactylus; sternal border with 1 seta. Dactylus with 1 large seta near

Pereopod 1 with merus about as long as carpus and bearing no setae or spines; carpus with 3 setae;

I propodus with 1 seta; dactylus with long whiplike seta. Pereopod 6 with carpus bearing 3 stout plumose

spines and 1 sternal setule; propodus with 2 plumose tergal spines and 1 plumose sternal spine on distal

margin; dactylus slender.

I Pleotelson, long, convex posterior margin.

1.6x broader than with

Uropod "pseudobiramous." Exopodite fused to basis with 1 subterminal and 2 long setae and 1

i terminal feathered hair. Endopodite 2-segmented; first segment 2.5x longer than broad and carrying 1 seta and 2 feathered hairs; second segment 1.9x longer than broad and bearing 5 setae and 1 feathered hair.

Description (male) (Figure 3.13). Body 4.8x longer than broad. Cephalothorax 1.5x broader than

I long,; pereonites broadest in3-5posterior similar,half. 2.2-2.5x Pereonites broader1,than 2, andlong. 6 similar Pleonitesin length 1-5 approximately and width, 3.3-3.5x 3x broaderbroader than long. than

First antenna with 2-segmented peduncle and 4- or 5-segmented flagellum. Formula for armature as

I follows: aesthetascs, 1 setanumerous + 3 feathered hairs, 2 setae + 2 feathered hairs, 2 setae + I feathered hair, numerous (~10) (~10) aesthetascs, 5 setae + 1 aesthetasc.

!

I 221
I

Figure 3.12. S_ahonolabrum californiensis Dojiri and Sieg, new species, female: _), general habitus of female;

A. 1, first antenna;L, labrum; Md, mandible; Mxp, maxilliped; Che, cheliped; P. 1, pereopod 1; P.6 i

pereopod 6; Uro, uropod. •

I
i

Figure 3.13. Siphonolabrum californiensis Dojiri and Sieg, new species,, male: C_, general habitus of male;

A.I, first antenna; Mxp, maxilliped; Che, cheliped; P.1, pereopod 1; P.6, pereopod 6; P1, pleopod;

t Pit, pleotelson; Uro, uropod.

i 223
!

Maxilliped basis fused medially and bearing 1 seta near insertion of each palpus. Endites each with I

1 seta. Palpus 4-segmented; first segment unarmed; second segment with 3 distal setae; third segment with

3 setae; and fourth segment with 6 setae. II

Cheliped [] basis as in female. Merus with 1 sternal seta at about midlength. Carpus about 1.3x longer

than broad, bearing 1proximal tergal seta and 1 distal tergal seta and 2 sternal setae. Propodus with 1rostral

seta, 2 longer caudal setae, and comb consisting of several (about 10) setae. Fixed finger with rugose (irregular)

tergal margin and bearing 3 setae near spiniform tip; sternal margin with 1 seta at about midlength. Dactylus I

with 1 large caudal seta near midlength.

Pereopod 1 more slender than in female; merus unarmed; carpus with 1distal tergal seta; propodus i

with small distal tergal seta; dactylus with 1 seta. Pereopod 6 merus with 2 stout tergal spines; carpus with m

3 stout spines; propodus with 3 spines and 1 seta; propodus and dactylus with row of spinules (or

denticulations). !

Pleopod biramous; exopodite and endopodite about 2x longer than broad and bearing numerous

pinnate setae. Is

Pleotelson about 1.4x broader than long, with protuberant posteromedial margin; tip of posteromedial I

protuberance with 2 small setae.

Uropod biramous. Exopodite l-segmented, 3x longer than broad, with 3 (?) setae. Endopodite 3- •

segmented, with armature as follows: 5 feathered hairs, 1 seta + 2 feathered hairs, and 5 setae.

Remarks. The genus Siphonolabrum can be easily recognized from the other tanaid genera inhabiting

the Santa Mafia Basin and Western Santa Barbara Channel by the proximal margin of the chelipedal basis

articulating directly with the cephalothorax; in other genera the cheliped articulates with the cephalothorax III

via a sclerotized plate known as the "side-piece."

The large patch of setules on the labrum, the absence of the pars molaris, the presence of long setae I

on the maxillipedal endites, the humplike process on the sternal surface of the chelipedal merus of the ;il

female, and the bicuspid tooth on the tergal border of the fixed finger of the cheliped of the female distinguishes

this species from all known taxa.

Etymology. The specific name alludes to the state from which the new species was collected.

Type Locality and Type Specimens. Santa Maria Basin and Western Santa Barbara Channel, i

California. Types deposited in the National Museum of Natural History (USNM) and the Santa Barbara II

Museum of Natural History (SBMNH); see "Type Material."

Distribution. Reported from its type locality, Santa Mafia Basin and Western Santa Barbara Channel, 1_

California south to Santa Monica Bay and San Pedro, California.

Subfamily Akanthophoreinae Sieg, 1986

Genus Paraleptognathia Kudinova-Pasternak, 1981

II

Paraleptognathia ef. gracilis (Kr_yer, 1842)

Figure 3.14 I

Material Examined. 1 neuter (dissected on slides) from phase II, cruise 1-1, Sta. PJ-2 (rep. 1),

34°55.30'N 120°45.90'W, Santa Maria Basin, California. !

l)eseription (neuter) (Figure 3.14). Body, 1.5 mm long, about 6x longer than broad. Cephalothorax,

approximately 1.3x longer than broad, with convex lateral margins. Pereonites with convex lateral borders, i

and decreasing in width posteriorly with pereonite 1 broadest and pereonite 5 narrowest; pereonite 6 shortest •

224 II
i Figure 3.14, Paraleptognathia cf. gracilis (Kr_yer, 1842), neuter: neuter, general habitus of neuter, A.1, first

antenna; Md, mandible; Che, cheliped; P. 1, pereopod 1; P.6, pereopod 6; Plt, pleotelson; Uro, uropod.

I

225
!

and slightly broader than pereonite 5. Pleonites 2-5 similar in size, each about 4-5x broader than long; I

pleonite 3 widest of pleonites.

First antenna 4-segmented with armature as follows: 1 seta + 4 feathered hairs, 1 seta + 2 feathered 1

hairs, 2 setae, and 5 setae + 1 aesthetasc, respectively.

Mandible with pars molaris bearing several (9?) pointed processes at tip.

Cheliped carpus moderately stout, about 1.5x longer than broad (length measured along tergal I

border); tergal border without pointed protrusion proximally; sternal border without pIatelike projection.

m

Propodus with crenulations on distal end of tergal margin near insertion of dactylus. Fixed finger with 3 m

spiniform processes ("teeth") and sclerotized tip; tergal border with 3 setae and 1 seta near articulation of HI

dactylus; sternal border with 2 setae. Dactylus with crenulations along tergal border.

u

Pereopod I with merus bearing 1 large spine; carpus with 1 tergal and 1 sternal spines; propodus Ill

with row of minute setules along tergal border and tergal spine at tip; dactylus with 1 seta. Pereopod 6 with |

carpus equipped with 3 pilose spines and 1 small naked seta; propodus with row of minute setules, 2 tergal

pilose spines and 3 sternal naked setae. m

Pleotelson, 1.5x broader than long, with slightly convex lateral borders, each with 1 seta at about m

midlength; posterior margin with protuberant median area bearing 2 small setae.

Uropods biramous. Exopodite 2-segmented, slightly shorter than first endopodal segment; first _l

segment, about 2x longer than broad, naked; second segment, with proportion similar to that of first segment,

having 2 setae at tip. Endopodite 2-segmented; first segment, 3.8x longer than broad, with 2 feathered hairs

and 1 seta; second segment, 4x longer than broad, with I feathered hair, 1 subterminal seta, and 2 small and Ill

3 long terminal setae.

Remarks. This single specimen was originally identified as Leptognathia sp. C. The original

description ofLeptognathia graciIis by Kr0yer (1842) is not adequate for comparative purposes. This specimen 1

may represent a new species, but should be carefully compared to specimens positively identified as

Jl

Paraleptognathia gracilis (Kr0yer, 1842) before a f'mal decision on its taxonomic status is made.

species can be distinguished from Paraleptognathia bisetulosa new species, described below, 1

This

by the shape of the cephalothorax, the presence of crenulations on the tergal border of the propodus of the

i

cheliped, and the number of setae on the merus of pereopod 1. m

Distribution. In addition to its type locality, this species has so far only been collected from the I

Santa Maria Basin at Sta. PJ-2, California.

Paraleptognathia bisetulosa Dojiri and Sieg, new species I

Figure 3.15 I

Type Material. 1female holotype (on slides; USNM) from cruise 1-3, Sta.R-6 (rep. 1), 34°41.40'N I

120°57.90'W, 410 m, off Purisima Pt., Santa Mafia Basin, California, May 1987, collected by Battelle. d

Description (female) (Figure 3.15). Body, 2 mm long, about 6x longer than broad. Cephalothorax, I

slightly longer than broad, with nearly straight parallel borders, and markedly tapered at anterior end.

1

Pereonites 2-4 with convex lateral borders; pereonite 6 narrowest of pereonites, with concave lateral borders. m

Pleonites 1-5 similar in length and width to each other, and about as broad as pereonites 2-4. II1

First antenna 4-segmented with armature as follows: 2 setae + 4 feathered hairs, 2 setae + 4 feathered

hairs, 2 setae, and 4 long and 1 short setae + 1 feathered hair + 1 aesthetasc. •

Mandible with pars molaris beating several pointed processes at tip. 1

I

226 I
II

Figure 3.15, Paraleptognathia bisetulosa Dojiri and Sieg, new species, female: Q, general habitus of female;

A.1, first antenna; Md, mandible; Che, cheliped; P.1, pereopod 1; P.6, pereopod 6; Plt, pleotelson;

i Uro, ttropod.

i 227
!

Cheliped carpus about 1.8x longer than broad (length measured along tergal border), with small I

platelike projection on sternal border; tergal border without pointed protrusion proximally; palm bearing

"comb" with 1 long and 3 short setae on caudal surface. Fixed finger with slender spiniform tip and 3 ll

smaller "teeth;" tergal border with 1 seta near insertion of dactylus and 3 setae near midlength; sternal |

border with 2 long setae. Distal part of propodus and dactylus with slight crenulations along tergal border.

Pereopod 1 with merus bearing 2 long, strong setae instead of spines; carpus with 1 tergal and 1 I

sternal setae; propodus with spiniform sternal comer and 1 tergal spine. Pereopod 6 with carpus carrying 3 I!

pilose setae and 1 small sternal seta; propodus with 2 tergal pilose setae and 3 sternal pilose setae.

setae. Pleotelson, 1.5x broader than long; posterior margin with protuberant median area bearing 4 minute i

Uropods biramous. Exopodite 2-segmented and slightly shorter than first endopodal segment; first

segment, about 3x longer than broad, with 1 long slender seta; second segment longer than first, about 4.5x i

longer than broad, bearing 1 long and 1 short setae. Endopodite 2-segmented; first segment 6x longer than

broad, and having 1 seta and 2 feathered hairs; second segment 8x longer than broad, as long as first

segment, but more slender, and carrying 1 subterminal and 3 long slender setae and 2 feathered hairs, l

I

Remarks. This species can be distinguished from Paraleptognathia cf. gracilis by the characters

outlined above in the Remarks section for the former species. The presence of two setae on the merus of

pereopod 1 distinguishes P. bisetulosa from its congeners. In addition, the single seta on the first exopodal il

segment of the uropod helps to separate this species from P. cf. gracilis.

Etymology. The specific name refers to the two setae on the merus of pereopod 1. l/

Type Locality and Type Specimen. Santa Mafia Basin, California. Types deposited in the National I

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

"Type Material." i Distribution. This species is so far known only from its type locality, Santa Maria Basin, California.

Genus Scoloura Sieg and Dojiri, 1991 i

Scoloura phiUipsi Sieg and Dojiri, 1991 II

II

Figure 3.16

Scolouraphillipsi Sieg and Dojiri, 1991: 1495-1501, figs. 1-4. /

Ill

Material Examined. Phase I: 4 females from Sta. 50, 34°37'48'N 121°0 l'40"W, 591 m, off Purisima

Pt; 1 neuter from Sta. 79, 34°24'07"N 120°28'19"W, 98 m, off Pt. Conception, Western Santa Barbara _l

Channel; 1 female from Sta. 73, 34°28'13"N 120°36'48"W, 98 m, between Pt. Conception and Pt. Arguello; 1

1 fragment from Sta. 3, 35°27.07'N 121°10.20'W, 291 m, off Pt. Estero, Santa Maria Basin, California.

From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring I

Division, City of Los Angeles: 1 female from Sta. 7B, 33°54'45"N 118°35'25"W, 138 m, 18 January 1977; l

1 female from Sta. 7B, 33°54'45"N 118°35'25"W, 180 m, 12 July 1984; 1 female from Sta. 8C, 33°58'47"N

118°36'17"W, 125 m, 19 January 1977; 1 female from Sta. E2, 33°58'39"N 118°39'16"W, 150m, 05 January E

1989; 2 females from Sta. 8C, 33°58'47"N 118°36'17"W, 120 m, 22 July 1977; 2 females from Sta. 4C,

33°50'40"N 118°26'22"W, 77 m, 24 July 1986; I female from Sta. 4C, 33°50'40"N 118°26'22"W, 08 January

1987; 1 female from Sta. 7B, 33°54'45"N 118°35'25"W, 172 m, 29 July 1985; 3 females (1 dissected) from 1

Sta. 4C, 33°50'40"N 118°26'22"W, 76 m, 13 January 1977; 1female from Sta. 7B, 33°54'45"N 118°35'25"W, II

11 July 1985; 1 female from Sta. 7B, 33°54'45"N 1I8°35'25"W, 190 m, 09 July 1982.

I

228 I
!

I From Los Angeles County Museum of Natural History: 5 females and 1 neuter (LACM 54-54) 7.8

mi 298° T (true north) from Palos Verdes Pt., 33°49'59"N 118°34'06"W, 165 m, sandy blue-gray mud, R/V

I Velero IV(AHF 2789-54), 22 May 1954; 2 females (LACM 60-100) 4.7 mi 349.5 ° T from Pt. Fermin Light to midpoint, 33°37'48"N 188°16'44"W, 219 m, R/V Velero IV (AHF 7161-60), 08 October 1960; 1 female

(LACM 55-67) 7.4 mi 229° T from end of E1 Segundo Pier, 33°50'00"N 118°32'23"W, 119 m, sand and

i mud, R/V Velero IV(AHF 3385-55), 23 August 1955; 2 females and 1 fragment from 6.3 mi 184° T from L.A. Breakwater Lighthouse, 33°36'14"N 118°15'30"W, 75 m, sand, R/V Velero IV (AHF 2355-53), 11

July 1953; 1 neuter (LACM 52-51) 9.2 mi 156.5° T from L.A. Lighthouse, 33°34'05"N 118°10'41 "W, 88 m,

i RIV Velero IV(AHF 2126-52), 25 June 1952; 2 females (LACM 54-40) 6.4 mi 304° T from Palos Verdes Pt., 33°50'00"N 118°32'00"W, 121 m, green sticky mud, R/V Velero IV (AHF 2727-54), 08 May 1954; 2

females and 2 neuters (LACM 54-38) 4.1 mi 331° T from Palos Verdes Pt., 33°50'00"N 118°28'00"W, 11

m, mud with stones, R/V Velero IV (AHF 2725-54), 08 May 1954; 1 female (LACM 53-125) 6.4 mi 239 ° T,

I end of Beach 33°47'03"N 118°30'07 229 R/V Velero IV 08

Redondo Pier, IIW, m, sand, (AHF2789-53), July

1953; 1 neuter CLACM60-83), 172 m, sandy mud, Coronado Canyon, 3.7 mi 013 ° T from N. Coronado Id.,

San Diego County, California, 32°30.27_ 117°16.80%V, R/V Velero IV (AHF 6845-60), 01 February 1960,

I EX_AHF [60-83].

Description (female) (Figure 3.16). Body, 3.60-5.00 mm long, about 7x longer than broad.

i Cephalothorax 1.3x longer than broad; anterior 1/3narrowed, posterior 2/3 with parallel borders; posterolateral corners with a pair of rounded dorsal lobes. Pereonites 1, 5, and 6 quadrangular; pereonites 2-4 hexagonal

with lateral margins noticeably convex in dorsal view. First pereonite 1.9× broader than long and narrowed

i posteriorly. Pereonite 2 about 1.2x broader than long with broadest area in anterior third. Pereonites 3 and 4 about as long as broad, and broadest at about midlength of each somite. Pereonite 5, narrowest of pereonal

somites, subrectangular, about as long as broad, and widest in posterior region. Pereonite 6 short, 1.8x

i broader than long, widest across posterior margin. Pleonites with 5 tergites visible dorsally; sternites with keel-like mid-longitudinal ridge; all somites of similar size, broader than pereonites and each about 4.5x

broader than long.

I + 2 feathered First antenna hairs; 24-segmented setae + 1 feathered with armature hair; andof7segments setae + 1asaesthetasc, follows: 1respectively. seta + 4 feathered hairs; 2 setae

Mandible with pars molaris terminating in 9-10 pointed processes.

I Cheliped palm bearing row of crenulations along tergal border and caudal surface with comb consisting of 5 setae. Fixed finger with strongly sclerotized tip; tergal border with 3-4 setae, 3 pointed teeth,

and I seta near articulation of dactylus; sternal border with 2 setae; rostral surface of fixed f'mger with row

I of blunt protuberances. Dactylus curved, bearing a sclerotized tip; tergal border crenulated.

Merus of pereopod 1 subtriangular, wider in distal region, and bearing 1 spine and 1 seta distally.

Dactylus of pereopods 4-6 elongate, with longitudinal groove lined with 2 rows of minute setules.

I Pleotelson bearing pair of large, ventrolateral, triangular-shaped processes; caudal point protuberant

and bent ventrally.

I first endopodal Uropod segment; biramous endopodite with very slender also 2-segmented. rami; exopodite 2-segmented and extending about 1/2 length of

Description (neuter) (Figure 3.16). Total length less than that of adult female. Pereonites with

I lateral margins not as convex and pteonites not as expanded as those of female. Remarks. This species was described by Sieg and Dojiri (1991) as a new genus and species based

on specimens obtained from this study and from the Southern California Association of Marine Invertebrate

I Taxonomists (SCAMIT) who identified these specimens originally as Leptognathia sp. C. It can be

distinguished from all known species by the presence of a pair of ventrolateral triangular-shaped processes

on the pleotelson.

i This to inhabit substrates of mud and silt.

species appears consisting sandy sandy

i 229
Figure 3.16. Scoloura Md, mandible; phillipsiChe,Siegcheliped; and Dojiri, P. 1, 199 pereopod 1, female: 1; P.6, _), pereopod general habitus 6; Uro,ouropod. f female; Neuter: A. 1, firstN,ante general nn; !

habitus of neuter.

i

230 I
!

I Type Locality and Type Specimens. Holotype female (LACM 81-134.1) and 4 female paratypes

(LACM 81-134.2; 1 dissected) from Sta. 4C, 33°50'40"N, 118°26'22"W, 89 m, sandy silt, Santa Monica

I the BayNatural , California, Historycollected Museumby the of Los Biology Angeles Laborat County. ory, Hyperion Treatment Plant, 22 July 1981, deposited in

Distribution. Pt. Sal (Santa Maria Basin) to Coronado Island (Mexico), 11-519 m (Sieg and Dojiri,

I 1991).

Genus Chauliopleona Dojiri and Sieg, new genus

I Type species (here designated): Chauliopleona dentata Dojiri and Sieg, new species.

II Diagnosis. Pleonite 5 with prominent, posteriorly-directed, sternal spiniform process.

I[ Remarks. Leptognathia armata, L. hastata, L. amdrupii, and the new species described below are

morphologically related, but generic transfer of the threeLeptognathia species into this new genus is postponed

I until detaileddetailed studyrede is completed. scriptions axedone for these species. A generic diagnosis can only be determined after a

Etymology. The generic name is a combination of the Greek words chaulios, meaning prominent,

I and pleo, meaning swim, and alludes to the prominent tooth on the pleon.

Chauliopleona dentata Dojiri and Sieg, new species

I Figure 3.17

t 1 femaleTanyped 1Masuteria badultl. ma 1 female le dissecte holodtype ) fro(USNM) m Sta. 4C,an33d 6°5p0aratypes '40"N 118°26'22"W, (SBMNH; 5 females, 76 m, Santa 1 suba Modnica ult male; Bay,

California, collected by the Biology Laboratory, Environmental Monitoring Division, 08 January 1987.

I 100 m, off MatePurisima rial Exami Pt., nSanta ed. Phase Maria I:Basin, 1femaleCalifornia, and 1 sub09adJanuary ult male 1984; from Sta. 2 females 42, 34°and 48.04'N 2 neuters 120°47.5 from0aSta. N,

79, 34°24.12'N 120°28.32'W, 98 m, off Pt. Conception, Western Santa Barbara Channel, California. Phase

I II: m, 2offfemaPurisima les, 1 suPt., badSanta ult male, Maand ria Basin, 4 neutersCalifornia, from cruisecollected 1-2, Sta. byRBattelle -4 (rep. 3), in January 34°43.0171987 q 12on0°47. board 39%VM,/92V

Aloha; 8 females, 4 neuters, and 1 manca-l] (dissected) from cruise 2-3, Sta. R-4 (rep. 1), 34°43.01'N

on120board °47.39M'W,/V 92Aloham,;off5 females PurisimaandPt.,2Santa neuterMaria s fromBasi cruise n, Calif 1-3,oSta. rnia,R-4collected (rep. 3),by34Battel °43.01'Nle in 120°47.39'W, January 1988

92 m, off Purisima Pt., Santa Maria Basin, California, collected by Battelle on May 1987 on board M/V

I From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring

A/oha.

Division, City of Los Angeles: I female and I subadult male from Sta. 4C, 33°50'40"N I 18°26'22"W, 76 rn,

I, • san Julydy1986; silt, 131female Januaryfrom197Sta. 7; 56B, females 33°52'27"N from Sta. 118°4C, 34'I2"W, 33°50'40"N 75 m, 30118°26'22"W, July 1986; 1asubadult pproximamtelyale from 76 m,Sta. 24

6B, 33°52'27"N 118°34'12"W, 79 m, 07 February 1986; 3 females, 1 subadult male, and 1 neuter from Sta.

I D2, 80 rn,33°1I53'4January 0"N 1181990; °35'210"W, female80 from m, 08Sta. JanuaryD3, 33199°51'47"N 0; 2 females 118°from 35'15"W, Sta. D4, 80 m, 33°517l'0July 7"N 1989; 118°31'3 1 female 0"W,

from Sta. 4C, 33°50'40"N 118°26'22"W, 76 m, 08 January 1987; 2 females from Sta. D2, 33°53'40"N

I 118 199°035'2 ; 2 females 0"W, 80from,m03Sta.September D4, 33°51'198 07"N7; 1118female °3 l'30fr"W, om Sta. 80 m, 134, 0333°51'07"N September 11198°31'3 87; 1 female 0"W, 8fr1omm,Sta. 20 July 6B,

33°52'27"N 118°34'12"W, 79 m, 21 January 1987; 2 females from Sta. IM, 33°51'07"N 118°31'30"W, 80

i m, 26 July 1990.

l 231
I

Figure 3.17. Chauliopleona dentata Dojiri and Sieg, new species, female: _, general habitus of female; _ A.1, !_

first antenna of female; Md, mandible; Che, cheliped; P.1, pereopod 1; (_ 1, terminal segment of

HE

pereopod 1 of neuters and some variant females; Uro, uropod. Subadult Oz, general habitus of In

subadult male; subadult O_A. 1,first antenna of subadult male. I

232 I
!

I From Los Angeles County Museum of Natural History: 1 female (LACM 59-147), 205 m, rocks,

2.45 mi, 249.5 ° T from Gull Island, Santa Cruz Channel, Santa Barbara County, California, 33°56.10'N

I 273 119°°52.28'W, T from baseR/Vof Velero El Segundo IV (AHF Pier, Lo6806-59), s Angeles22County, December California, 1959; 133°53.00 neuter (LA 'N 115°30.63_, CM 55-46), 53R/Vm,Velero 4.25 mi, IV

(AHF 2994-55), 05 February 1955; 1female (LACM 55-67), 119 m, sand and mud, 7.4 mi, 229 °T from end

I of 55),E123Segundo AugustPier, 1955;Los1 Angeles County, California, 33°50.00'N 118°32.38'W, R/V Velero IV (AHF 3385- female (LACM 54-38), 11 m, mud with stones, 4.1 mi, 331° T from Palos Verdes

Pt., Los Angeles County, Califorrfia, 33°50.00'N 118°28.00'W, R/V/Velero IV (AHF 2725-54); 1 female

i (LACM California,54-54), 165 m, sandy blue-gray mud, 7.8 mi, 298 ° T from Palos Verdes Pt., Los Angeles County 33°49.98'N 118°34.10'W, R/V Velero IV (AHF 2789-54), 22 May 1954; 2 females (LACM 60-

100), 219 m, 4.7 mi, 349.5 ° T from Pt. Fermin Light to midpoint, Los Angeles County, California, 33°37.80'N

i 118°16.73'W, R/V Velero IV(AHF 7161-60), 08 October 1960; 1female (LACM 60-86), 949 m, mud over sand, 14.95 mi, 264° T from Pt. La Jolla, San Diego County, California, 32°49.62'N 117°35.20'W, R/V

Velero IV (AHF 7049-60), 07 May 1960; 1female and 1 neuter (LACM 60-110), 585 m, mud, 31.5 mi, 254 °

T from China Point Light, Tanner Canyon, San Clemente Island, California Channel Islands, California,

I 32°39.55'N 119°01.40'W, RN Velero IV (AHF 6834-60), 29 January 1960; 1 female (LACM 60-84), 225

m, mud, 34.1 nil, 246 ° T from China Point Light, San Clemente Island, Tanner Bank, California Channel

Islands, California, 32°34.47'N 119°02.70'W, R/V Velero IV (AHF 6837-60), 29 January 1960; 1 female

I (I.,ACM 60-183), m, muddy sand, mi, Norte, Coronados, Baja

108 3.85 024 ° T from Coronado Islas de Los

California, Mexico, 32°30.30'N 117°16.07'W, R/V Velero IV (AHF 6846-60), 01 February 1960.

I Cephalothorax, Descriptio1.3xn (female) (Figure 3.17). Body, 2-4 mm long, about 6.7x longer than broad. longer than broad, with relatively straight parallel lateral margins, abruptly tapered

anteriorly. Pereonites 1-4 similar in width; pereonite 5 narrowest; pereonite 6 shortest. Pleonites 1-5 similar

I in size, 4-5x First broader antenna than 4-segmented long; pleonite with armature 5 with large, as follows: psteriorly-directed, 1 seta + 3 feathered sternal,hairs, spiniform 1 seta +process. 2 feathered

hairs, 2 setae, and 6 setae + 1 feathered hair + 1 aesthetasc.

I Mandible with pars molaris bearing approximately 10 pointed processes at tip.

Cheliped carpus about 1.7x longer than broad; palm with comb bearing 2 small rows of spinules, 4

short setae, and 1 longer seta. Fixed finger with sclerotized tip and 3 small teeth; tergal border with 3 long

I and 1 shorter setae near articulation of dactylus; sternal border with 2 setae. Dactylus with crenulations along tergal margin.

Pereopod 1 with merus bearing 1 pinnate spine; carpus with tergal row of spinules, 1 tergal spine,

I and 2 setae; propodus with 1tergal row of hyaline spatulate spinules in females with marsupia, other females

and neuters with attenuate spinules (see female 1 in figure), curved rows of minute setules, 1 stout pinnate

spine, and 1 slender seta.

I Pleotelson, 1.7x broader than long, with protuberant posteromedian area.

Uropods biramous. Basis with 3 small setae near insertion of endopodite. Exopodite 2-segmented,

I distinctly seta; secondshorter segment than 4xfirstlonger endopodal than broad, segment;andfirst bearisegment ng I longaboutand2.7x1 short longersetaethanatbroad, tip. Endopodite with 1 distal2-

segmented; first segment 4.5× longer than broad, and equipped with 1 seta and 2 feathered hairs; second

I segment Des 6.5x longer than broad, with 1 subterminal and 3 long setae and 2 feathered hairs terminally. cription (subadult male) (Figure 3.17). Body smaller than that of adult female, but proportions

of body somites similar. Cephalothorax similar in shape, about 1.25× longer than broad.

I First antenna 5-segmented with armature formula as follows: 1 seta, 2 setae, 2 setae, 0, 6 setae + 1

aesthetasc.

i

i 233
!

Remarks. The posteriorly-directed spiniform process on the ventral (sternal) surface of the last I

pleonite helps distinguish Chauliopleona dentata from all other tanaidacean species collected in the Santa

Mafia Basin and Western Santa Barbara Channel. This species appears to be closely related to Leptognathia _l

armata, L. hastata, and L. amdrupii, which may eventually be transferred to Chauliopleona (see generic !1

"Remarks" section above). Unfortunately, the descriptions of these three species are not adequate for a

detailed comparison with C. dentata. The crenulation along the chelipedal dactylus helps distinguish C.

dentata from the above three species. I!

Specimens collected by the Environmental Monitoring Division were previously identified as

"'Leptognathia sp. E" of SCAMIT. I

Etymology. The specific name refers to the numerous spatulate spinules on the propodus of pereopod

1 in females with rnarsupia.

Type Locality and Type Specimens. Santa Monica Bay, California. Types deposited in the National !

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

IIW

"Type Material." R

Distribution. This species has been collected from Santa Mafia Basin south to Coronado Norte, 1

Islas de Los Coronados, Baja California, Mexico.

Genus Araphura Bird and Holdich, 1984 I

Araphura breviaria Dojiri and Sieg, new species I

Figure 3.18

Type Material. 1 female holotype (USNM) and 3 paratypes (1 female, 2 neuters; SBMNH) from i

Sta. 4C, 33°50'40"N 118°26'22"W, 76 m, Santa Monica Bay, California, 08 January 1987. II

Material Examined. Santa Mafia Basin, California, phase I: 2 females from Sta. 54, 34°36'34"N

120°52'01"W, 396 m, between Pt. Arguello and Purisima Pt., November 1984; 2 females from Sta. 61, l[

34°33'01"N 120°48'53"W, 345 m, off Pt. Arguello, November 1984. Phase II: 20 neuters, 3 manca-I, 4

u

manca-ll, and 6 fragments from cruise 2-4, Sta. R-4 (rep. 2), 34°43'01"N 120°47'23"W, off Purisima Pt.; 14 m

neuters, 4 manca-ll, and 1manca-I from cruise 1-3, Sta. R-4 (rep. 3), 34°43'01 "N 120°47'23"W, offPurisima ll

Pt. u

From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring I

Division, City of Los Angeles: 2 females from Sta. El, 33°59'03"N 1I8°42'50"W, 150 m, 09 July 1990; 1 .l_

female and 1 neuter from Sta. B5, 33°58'00"N 118°31'45"W, 45 m, 11 August 1987; 2 females from Sta.

2B, 33°56'25"N 118°29'14"W, 37 m, silt; 1 female from Sta. 4C, 33°50'40"N 118°26'22"W, 76 m, sandy Ill

silt;3 females (2 dissected) from Sta. 3B, 33°54'45"N 118°28'29"W, 40 m, silt with organic debris; 2 females •m

from Sta. OC, 34°01'17"N 118°36'00"W, 29 m, clay silt.

From LACM: 3 females and 1 neuter (LACM 55-35), 9.9 mi, 203° T from Santa Monica Pier Light, II

Los Angeles County, California, 33°51'22"N 118°34'42"W, 73 m, mud, R/V Velero IV (AHF 2998-55), 06 |

February 1955; 2 females (LACM 55-36) 14.3 mi., 345 ° T from Aero Beacon, San Nicolas Island, Ventura

County, California, 33°28'04"N 119°34'41 "W, 732 m, rock, sand, and mud, R/V Velero IV(AHF 3031-55), II

07 May 1955; 1 female (LACM 52-51) 9.2 mi., 156.5 ° T from L.A. Lighthouse, Los Angeles County, l

California, 33°34'05"N 118°10'41"W, 88 m, R/V VeIero IV(AHF 2126-52), 25 June 1952.

Description (female) (Figure 3.18). Body, 2.45-3.64 mm long, about 7x longer than broad. Ill

Cephalotborax, stout, 1.1× longer than broad, anterior half narrowed, posterior half with parallel sides. II

Pereonites subquadrangular; pereonites 2-5 with irregular, slightly convex lateral borders. First pereonite

1.6x broader than long and tapered posteriorly. Pereonite 2 about 1.2x broader than long. Pereonites 3-5 tll

II

234 |
!

I

f

Che

A.1 "-

t

i f

I _ oro I

I i, _', \

| / ,

i

FiN 3.18. Ar_ph_ brev_ia DojM and Sieg, new species, female: (_, general habitus of female; Md,

i mandible; Che, cheliped; P.1, pereopod 1;P.6, pereopod 6; Pit, pleotetson; Uro, uropod.

i 235
!

about as long as broad. Pereonite 6 shortest of pereonal somites, 1.4x broader than long. Pleonites 1-5 I

similar in size, about as broad as pereonites 1 and 2, and each about 5.6x broader than long. II

First antenna 4-segmented with armature as follows: 1 seta + 5 feathered hairs; 1 seta + 2 feathered /

hairs; 2 setae + 1 feathered hair; and 8 setae + 1 aesthetasc, respectively. I

Mandible with pars molaris terminating in 7 pointed processes.

Cheliped palm bearing "comb" of 5 short setae and 1 longer seta on caudal surface. Fixed finger •

with strongly sclerotized tip; tergal border with 3 setae and 1 seta near articulation of dactylus; sternal II

border with 2 setae. Dactylus with 1 sternal (or 2 sternal ?) setae.

Pereopod 1 with merus bearing 1 seta and 1 spiniform seta tergally; carpus with 1 tergal spiniform

seta and 1-2 sternal setae; propodus with 1 tergal seta. Pereopod 6 with carpus bearing 2 barbed spines il

tergally and 2 barbed spines and 1 seta sternally; propodus having several rows of minute spinules along

sternal border and bearing 2 barbed spines tergally and 3 barbed spines sternally; dactylus with longitudinal

groove lined with 2 rows of minute setules. |

Pleotelson short, 1.6x broader than long, with relatively straight lateral borders.

Uropod "pseudobiramous;" exopodite fused with basis. Fused exopodite reaching beyond midlength l_

of first endopodal segment and bearing 2 long lateral setae and 1 short subterminal seta. Endopodite 2-

m

segmented; fn'st segment 4.3x longer than broad, with 1 long seta and 2 feathered hairs; second segment I

about 1/2 length of first, 2.5× longer than broad, and beating 1 long subterminal seta and 1 feathered hair, 2 •

short setae, and 3 long setae at tip. m

Remarks. Araphura breviaria is described from MMS specimens and from specimens in the i

SCAM1T collections. These specimens were originally identified as "'Araphura sp. A" and "'Leptognathia |

sp. A". It can be distinguished from A. cuspirostris new species by the short pleotelson, straight lateral

borders of the pleotelson, and absence of the conical knob on the posterior border of the pleotelson.

This species has been collected from various types of sediments (e.g., sandy silt, silt, and silty clay) I

ranging in depth from 29-732 m.

Etymology. The specific name is from Latin meaning abridged or shortened and alludes to the I

short pleotelson of this species. II

Type l.,aeality and Type Specimens. Santa Monica Bay, California. Types deposited in the National

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see I

"Type Material." U

Distribution. Santa Mafia Basin south to Los Angeles Light, Los Angeles County. I

41

Araphura euspirostris Dojiri and Sieg, new species

Figure 3.19 !

Type Material. 1 female holotype (USNM), 13 female paratypes (2 females dissected on slides;

SBMNH) from Sta. 55, 34°33'40"N 120°56'19"W, 590 m, west of Pt. Arguello, Santa Mafia Basin, California,

September 1984. II

Material Examined. Santa Maria Basin, phase I: 4 females, 7 neuters from Sta. 59, 34°33'39"N

120°47'11 "W, 216 m, southwest of Pt. Arguello, October 1984. Phase II: 7 neuters, 1 manca-I from cruise

1-3, Sta. R-4 (rep. 3), 34°43'01 "N 120°47'23"W, 92 m, southwest of Purisima Point; 8 females, 4 neuters, 3

II

manca-I, 3 manca-II, 3 fragments from cruise 2-4, Sta. R-4 (rep. 2), 34°43'01 "N 120°47'23"W, southwest of

Purisima Point; 2 neuters from cruise 1-3, Sta. R-4 (rep. 3), 34°43'01 "N 120°47'23"W, 92 m, southwest of

Purisima Point. I

From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring i

Division, City of Los Angeles: 1 female from Sta. E3, 33°58'21"N 118°36'44"W, 149 m, 10 July 1989. I

236 I
!

I From LACM: 1female (LACM 59-264) 8.75 mi., 235° T from Santa Monica Pier Light, Los Angeles

County, California, 33°55'29"N 118°38'32"W, 457 m, sand, RN Velero IV(AHF 6779-59), 19 December

i County, 1959; 3 females California, and33°55'30"N 3 neuters (LACM 118°35'55"W, 55-96) 7.1293mi., m, mud 226°andT from Santa Monica Pier Light, Los Angeles silt, R/V Velero IV (AHF 3180-55), 05 July

1955; 2 females (LACM 54-55) 5.5 mi., 287.5 ° T from Palos Verdes Point, Los Angeles County, California,

i 33°48'00"N 118°32'00"W, 459 m, blue-grey mud, R/V Velero IV (AHF 2793-54), 22 May 1954; 1 female (LACM 53-127) 8.45 mi., 184° T from L.A. Breakwater Lighthouse, Los Angeles County, California,

33°34'07"N 118°15'44"W, 366 m, clay, R/V Velero IV (AHF 2354-53), 02 July 1953; 1 female (LACM 60-

101) 3.8 nil., 163.5 ° T from Pt. Fermin Light, Los Angeles County, California, 33°38'36"N 118°16'16"W,

I 214 m, mud, R/V Velero IV(AHF 7174-60), 09 October 1960; 1 female (LACM 55-101) 11.7 mi., 257 ° T

from end of E1Segundo Pier, Los Angeles County, California, 33°52'08"N 118 °39'18"W, 457 m, mud, R/V

Velero IV (AHF 3399-55), 23 August 1955; 1 female (LACM 60-108) 0.2 mi., 254.5 ° T from base of

I Newport Pier, Newport Canyon, Orange County, m,

California, 33o36'24"N 117°55'54"W, 16 silt, R/V

Velero IV (AHF 7031-60), 05 May 1960; 1 female (LACM 60-124) 4.8 nil., 019 ° T from Port Hueneme

Light, Ventura County, California, approximately 34°03'55"N 119°13N¢, 428 m, mud, sand, and pebbles,

I R/V Velero IV (AHF 6899-60), 11 March 1960; 4 females and 1 neuter (LACM 53-128) 3.4 mi., 301 ° T

from Pt. Vicente Light, Los Angeles County, California, 33°44'08"N 118°25'48"W, 183 m, sand, R/V VeIero

IV(AHF 2357-53), 08 July 1953; 1 female (LACM 53-125) 6.4 mi., 239 ° Tend of Redondo Beach Pier, LOs

I Angeles County, California, 33°47'03"N 118°30'07"W, 229 m, sand, R/V Velero IV (AHF 2789-53), 08

July 1953; 1 female (LACM 60-86) 14.95 mi., 264° T from Pt. La Jolla, San Diego County, California,

32°49'37"N 117°35'12"W, 949 m, mud over sand, R/V Velero IV(AHF 7049-60), 07 May 1960; 1 female

I (LACM 59-147) 2.45 mi., 249.5 ° T from Gull Island, Santa Cruz Channel, Santa Barbara County, California, 33°56'06"N 119°52'17"W, 205 m, rocks, R/V Velero IV (AHF 6806-59), 22 December 1959.

i Description (female) (Figure 3.19). Body, 2.73-3.49 mm long, about 9.2x longer than broad. Cephalothorax, slender, 1.4x longer than broad, and tapered anteriorly. Pereonites subquadrangular; pereonites

I and 6 similar in length and width, about 1.2x broader than long. Pereonites 2-5 similar in length, but

pereonites 4 and 5 narrower than preceding somites. Pleonites 1-5 similar in size, each about 4x broader

I than long.

First antenna 4-segmented with armature as follows: 1 seta + 4 feathered hairs; 1 seta + 4 feathered

I hairs; 2 setae; and 6 setae + 1 aesthetase, respectively.

| Mandible with pars molaris bearing 6 pointed processes at tip.

Chetiped palm bearing "comb" of 3 short and 1 long setae on caudal surface. Fixed finger with

I strongly sclerotized tip; tergal border with 3 long and 1 shorter setae near articulation of dactylus; sternal border with 2 long setae. Dactylus with 1 caudal seta and 1 sternal seta.

Pereopod 1 with merus bearing 1 seta and 1 spiniform seta tergally; carpus with 1 tergal spiniform

i seta and 1-2 sternal setae; propodus with 3 setae. Pereopod 6 with bearing 2 barbed spines tergally

carpus

and 2 barbed spines and 1 seta sternally; propodus having several rows of minute spinules on sternal border

and beating 2 barbed spines tergally and 3 barbed spines sternally; dactylus with longitudinal groove lined

I with 2 rows of minute setules.

Pleotelson subquadrate, 1.2x broader than long, with slightly concave lateral borders, and bearing

small conical or digitiform process on posterior border.

I Uropod "pseudobiramous." Fused exopodite reaching about 3/4 length of proximal endopodal

segment and carrying 3 setae. Endopodite 2-segmented; first segment slender, 5.3x longer than broad, with

i 1 long seta and 1 feathered hair distally; second segment about 1/2 length of first, with 1 long subterminal seta, 1 feathered hair, 2 short setae, and 3 long setae at tip.

Description (neuter). Total length less than that of adult female; habitus as in figure. Lateral borders

i of pleotelson slightly concave.

i 237
Figure 3.19. Araphura cuspirosUis Dojiri and Sieg, new species, female: _, general habitus of female; A.1, I

first antenna; Md, mandible; Che, chelipexi; P. 1, pereopod 1; P.6, pereopod 6; Plt, pleotelson; Uro,

uropod. Neuter, general habitus of neuter, i

238 i
!

I Remarks. Araphura cuspirostris is described from specimens originally obtained for MMS and from specimens belonging to SCAM1T. These specimens were originally identified as Araphura sp. B and

Leptognathia sp. D. It can be distinguished from A. breviaria by the general shape (subquadrate) of the

i ple process otelsoonn, the the poste slightrilyorcoborder ncave lateral of the bpleotelson. orders of the pleotelson, and the presence of the conical or digitiform

Araphura cuspirostris has been collected from a wide variety of sediments including rocks, pebbles,

I sand, mud, muddy silt, and silt.

Etymology. The specific name is a combination of the Latin words cuspis, meaning point, and

rostrum, meaning beak, and alludes to the small conical process on the pleotelson.

I Type Locality and Type Specimens. Santa Maria Basin, California. Types deposited in the National

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

I "Type Material." Distribution. Santa Maria Basin south to Pt. La Jolla (San Diego County).

i Genus Tanaella Norman and Stebbing, 1886

I TanaeUa propinquus Dojiri and Sieg, new species Figure 3.20

I Type Material. 1 female holotype (USNM) and 4 paratypes (SBMNH; 3 females and 1 neuter) from Sta. E7, 33°54'44"N 118°34'17"W, 150 m, Santa Monica Bay, California, collected by Biology

Laboratory, Environmental Monitoring Division, 16 July 1990.

I Material Examined. Santa Maria Basin and Western Santa Barbara Channel, phase I: 1 female and 7 neuters from Sta. 84, 34°13'32"N 120°31'11 "W, 394 m, south of Pt. Conception, Western Santa Barbara

Channel, California, February 1985; 2 females from Sta. 73, 34°28'13"N 120°36'48"W, 98 m, between Pt.

I Conception and Pt. Arguello, Santa Maria Basin, California, January 1985. Phase I/: 7 females, 6 neuters, and 21 manca-I from cruise 3-1, Sta. R-4 (rep. 3), 34°43'01"N 120°47'23"W, 92 m, off Purisima Pt., Santa

Maria Basin, California; 7 females, 4 neuters, 4 fragments, and 58 manca-I from cruise 2-3, Sta. R-4 (rep.

I 1), 34°43'01 "N 120°47'23"W, 92 m, off Purisima Pt., Santa Maria Basin, California.

From Santa Monica Bay, California, collected by Biology Laboratory, Environmental Monitoring

Division, City of Los Angeles: 2 females from Sta. E2, 33°58'39"N 118°39'16"W, 150 m, 04 January 1989;

I 2 females from Sta. E3, 33°58'21"N 118°36'44"W, 149 m, 10 July 1989; 1female from Sta. 7B, 33°54'45"N 118°35'25"W, 168 m, 11 February 1986; 2 females from Sta. 7B, 33054'45"N 118°35'25"W, 21 January

1987; 1 female from Sta. E8, 33°54'32"N 118°35'49"W, 150 m, 17 January 1989; 2 females and 1 manca-I

i from Sta. E8, 33°54'32"N 118°35'49"W, 150 rn, 20 August 1987; 1 female from Sta. E7, 33°54'44"N 118°34'17"W, 148 m, 17 July 1989; 1 female from Sta. E8, 33°54'32"N 118°35'49"W, 149 m, 17 July 1989;

2 females from Sta. E3, 33°58'21"N 118°36'44"W, 150 m, 11 August 1987; 1 specimen from Sta. E8,

I 33°54'32"N 118°35'49"W, 150 m, 08 January 1990; 1 female from Sta. E8, 33°54'32"N 118035'49"W, 151 m, 25 January 1988; 4 females from Sta. E7, 33°54'44"N 118°34'17"W, 150 m, 14 July 1988; 3 specimens

from Sta. 8C, 33°58'47"N 118°36'17"W, 116 m, 08 January 1987; 1 female from Sta. C7(2), 33°53'35"N

I 118°32'15"W, 59 m, 16 July 1990; 1 female from Sta. E2, 33°58'39"N 118°39'16"W, 152 m, 28 January 1988; 4 females from Sta. El, 33°59'03"N 118°42'50"W, 150 m, 09 July 1990.

From LACM: 4 females (LACM 55-67) 7.4 mi, 229 ° T (true north) from end of E1Segundo Pier,

I 33°50'00"N 118°32'23"W, 119 m, sand and mud, R/V Velero IV (AHF 3385-55), 23 August 1955; 1 female (LACM 54-40) 6.4 mi, 304° T from Palos Verdes Pt., 33°50'00"N 118°32'00"W, 121 m, green sticky mud,

RIV Velero IV (AHF 2727-54), 08 May 1954; 2 females (LACM 53-78) 1.55 mi, 002 ° T from Long Pt.

I Light, Santa Catalina Island, California Channel Islands, 33°25'57"N 118°21 '54"W, 320 m, clay mixed

i 239
!

with broken shell sand, R/V Velero 1V (AHF 2423-53), 29 September 1953; 9 females and 3 neuters I

(LACM 54-54) 7.8 mi, 298 ° T from Palos Verdes Pt., 33°49'59"N 118°34'06"W, 165 m, sandy blue

gray mud, R/V Velero IV (AHF 2789-54), 22 May 1954; 3 females and 1 neuter (LACM 55-96) 7.1 mi, 8

226 ° T from Santa Monica Pier Light, 33°55'30"N 118°35'55"W, 293 m, mud and silt, R/V Velero IV(AHF I

3180-55), 05 July 1955; 4 females and 5 neuters (LACM 53-125) 6.4 mi, 239 ° T from end of Redondo

Beach Pier, 33°47'03"N 118°30'07"W, 229 m, sand and mud, R/V Velero IV (AHF 2361-53), 08 July 1953; IBm

13 females and4 neuters (LACM 60-101), 3.8 mi, 163.5° T from Pt. Fermin Light, 33°38'36"N 118°16'16"W, I

214 m, mud, R/V Velero IV (AHF 2361-53), 08 July 1953. From San Diego County: 3 females (LACM 60-

83) 3.7 mi, 013° T from N. Coronado Island, 32°30'16"N 117°16'48"W, 172 m, sandy mud, R/V Velero IV I 

(AHF 6845-60), 01 February 1960. From Mexico: 3 females (LACM 60-183) 3.85 mi, 024 ° T from Coronado I

Norte, Islas de Los Coronados, Baja California, 32°30'18"N 117°16'04"W, 108 m, muddy sand, R/V Velero IV

(AHF 6846-60), 01 February 1960. mR

Description (female) (Figure 3.20). Body, 2.78-3.31 mm long, about 7.9x longer than broad.

Cephalothorax, 1.4x longer than broad, narrowed anteriorly. Pereonites with convex lateral borders; pereonites

1, 2, and 3 similar in width; pereonite 3 longest and broadest of pereonites; pereonite 6 shortest, with small i

ventrally-directed spiniform process on sternite; pereonites 5 and 6 similar in width and narrowest of I

pereonites. Pleonites 1-5 similar in size, each about 5x broader than long, and bearing ventral hyaline keel

along its length. II

First antenna 4-segmented with armature as follows: 1 seta + 4 feathered hairs; 1 seta + 2 feathered HI

hairs; 2 setae; and 6 setae + 1 aesthetasc, respectively.

Mandible with pars molaris beating several pointed processes at tip. I

u

Cheliped carpus relatively slender; palm bearing comb of 4 short and 1 long pinnate setae on caudal

surface. Fixed finger with strongly sclerotized tip; tergal border with 3 long and 1shorter setae near articulation m

of dactylus; sternal border with 2 long setae. Dactylus with 1 caudal seta and 1 sternal seta. i

Pereopod 1 with merus beating 1 seta and 1 barbed spine; carpus with 1 seta and 2 barbed spines;

propodus slender, with 2 setae and 1 barbed spine. Pereopod 6 with propodus bearing several rows of ii

minute spinules and 4 spiniform setae; dactylus with longitudinal groove lined with 2 rows of minute setules. I

Pleotelson, 1.7× broader than long, with convex lateral borders; posterior margin concave on either

side of very slightly protuberant median area which bears 2 small setae; posterior margin with posteriorlydirected spiniform process. |

Uropods curved inwards, giving "bow-legged" appearance; 4.4x longer than broad. Exopodite

presumably represented by 3 naked setae at distal end of basis. Endopodite 2-segmented; first segment 1.9x

longer than broad, with 1 seta and 2 feathered hairs; second segment 2.2x longer than broad and beating 1 II

short and 4 long setae.

Remarks. Tatmellapropinquus is described from specimens originally obtained for MMS, SCAMIT, n

and the Los Angeles County Museum of Natural History. SCAMIT specimens were identified as Leptognathia U

sp. B and later simply as TanaeIla sp. It can be distinguished from all other tanaidaceans from the MMS

survey by the "bow-legged" appearance of the paired uropods. This species has been collected from a i

variety of substrates (e.g., sandy mud, clay mixed with broken-shell sand, silty mud, and mud). I

Etymology. The specific name is from the Latin word meaning "near" and alludes to its I

morphological similarity to other species of its genus. UU

Type Locality and Type Specimens. Santa Monica Bay, California. Types deposited in the National

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see am

"Type Material." |

Distribution. This species has so far been collected from Santa Maria Basin south to Coronado

Norte, Islas de Los Coronados (Baja California). I

|
I

I

[

I

I

#

i tt

I

I _gure 3.20. Tanaellapropinquus Dojiri andSieg,new species, female: _), generalhabitusof female; A. 1,first

antenna;Md,mandible;Che, cheliped;P.1,pereopod1;P.6,pereopod6;Pit,pleotelson;Uro,uropod.

i

i 241
!

Genus Incertae Sedis m

II

Male species 1

Figure 3.21 m

Material Examined. 1 male from, Sta. 42 (phase I), 34°48.04'N 120°47.50'W, 100 m, west of

Purisima Pt., Santa Maria Basin, California, 06 January 1984, originally identified as Cryptocope sp. C. I

Description (male) (Figure 3.21). Body about 4x longer than broad. Cephalothorax, about as long U

as broad, narrowed anteriorly. Pereonites 1-3 about 4-5.5x broader than long; pereonites 4 and 6 approximately

3x broader than long; pereonite 5 longest of pereonites, 2.5x broader than long. Pleonites 3-3.5x broader m

than long, except pleonite 4 about 2.6x.

First antenna 7-segmented with numerous aesthetascs on segments 4-6; terminal segment with 5

setae and 1 aesthetasc.

u

Maxilliped with fused basis bearing 1 distal seta near insertion of palpus. Endites each with 1 distal

seta. Palpus 4-segmented; first segment unarmed; second segment with 2 setae; third segment with 4 setae; im

and fourth

segment bearing 6 setae, m

Chelipedal carpus robust, about 1.6x longer than broad, distally inflated (segmental margin indicated

by dotted line and arrow in figure; area above dotted line depicts inflated portion), and bearing t tergal and •

1 sternal setae. Fixed finger with spiniform tip; tergal border with 3 setae; area near insertion of dactylus II

with 1 small seta; "comb" with 3 relatively long setae (or spines) and a row of several spines; sternal margin

with 2 setae. m

Pereopod 1 with merus bearing 1 spine and 1 seta; carpus with 2 spines and 1 seta; propodus with

several rows of minute spinules. Tergal spines of merus and carpus of pereopods 1-3 long, reaching to about

midlength of following segment. Pereopod 6 with long slender dactylus bearing long rows of spinules.

II Pleopods without serrations along outer margin of exopodite.

Pleotelson, 1.2x broader than long, somewhat triangular in outline. n

Uropod biramous. Exopodite 2-segmented; first segment, 2.3x longer than broad, bearing 1 long I

seta; second segment 3.6x longer than broad, carrying 1 long and 1 shorter setae. Endopodite 2-segmented;

first segment longer than entire exopodite, about 4x longer than broad, and bearing 1 seta and feathered

hairs; second segment, 4.3x longer than broad, with 1 subterminal and 4 terminal setae and 1 feathered hair. |

Remarks. Mate species l differs from the other two unidentified males (species 2 and 3) by having

2-segmented rami of the uropod (exopodite 1-segmented and endopodite 3-segmented in species 2 and 3), ...IN

an inflated distal margin of the chelipedal carpus, and tergal spines of the merus and carpus of pereopods 1- l

3 reaching to about midlength of following segment. This male may belong to the new genus ChauliopIeona.

Distribution. Santa Maria Basin, California. I

Male species 2

Figure 3.22 m

Material Examined. 1 male from Sta. 42 (phase I), 34°48.04'N 120°47.50'W, 100 m, west of •

Purisima Pt., Santa Maria Basin, California, October 1984, originally identified as Cryptocope sp. C. II

Description (male) (Figure 3.22). Body about 4x longer than broad. Cephalothorax, 1.3x broader

than long, constricted at about midlength, anterior portion narrower than posterior. Pereonites 1-6 similar in m

length/width ratios, ranging from 3.5x to 4.1x broader than long. Pleonites 1-5 similar to each other. Ill

First antenna 7-segmented with numerous aesthetascs on segments 4-6; terminal segment with 5

setae and a feathered hair. m

242 m
I

I

i

"1 i •/'

• A.1

I _

i I : " .

I { '

,

!

J

I 01 P!

I

Uro •

Pit

I , P,1

| L R6

I

I

!

Figure 3.21. Genus incertae sedis, male species 1: O_ 1, general habitus of male; A.1, first antenna; Mxp,

t maxilliped; Che, cheliped; P. 1,pereopod 1;P.6, pereopod 6; P1,pleopod; Pit, pleotelson; Uro, uropod.

i 243
!i

Figure 3.22. Genus incertae sedis, male species 2: O_2, general habitus of male; A.1, first antenna; Mxp, maxilliped; I1

Che, cheliped; P. 1, pereopod 1; P.6, pereopod 6; P1, pleopod; Pit, pleotelson; Uro, uropod.

!
!

I Maxilliped similar to that of male species 1; palpus with third segment bearing 3 setae.

Cheliped as in male species 1 except inflated area on distal margin of carpus lacking.

i Pereopods 1-3 with tergal spines on merus and carpus short, not reaching to midlength of following segment. Pereopod 6 with i spine and 1 seta on ischium and rows of spinules along length of dactylus.

Pleopods as in male species 1. II

I Pleotelson 1.4x broader than long, with posterior portion triangular in outline.

Uropod biramous. Exopodite 1-segmented, 2.3x longer than broad, and bearing 3 setae. Endopodite

I 3-segmented; than broad, withfirst1segment seta and 1.3x a feathered longer than hair; broad, third segment with several 3x longer featheredthanhairs; broad,second with 1segment subterminal 2.5x longer seta, 4

terminal setae, and feathered hair.

I Remarks. Male species 2 can be distinguished from male species 1 by the 1-segmented exopodite and 3-segmented endopodite of the uropod and from male species 3 by the shape of the cephalothorax and

pleotelson, the absence of denticles on the outer margin of the pleopodal exopodite, and the presence of 1

I spine and 1 seta on the ischium of pereopod 6. Distribution. Santa Mafia Basin, California.

I Male species 3

Figure 3.23

I Material Examined. 1 mate from Sta. 42 (phase I), 34°48.04'N 120°47.50'W, 100 m, west of

Purisima Pt., Santa Maria Basin, California, 06 January 1984, originally identified as Cryptocope sp. C.

I Description (male) (Figure 3.23). Body about 3.8x longer than broad. Cephalothorax, 1.2x broader than long, with distinctly narrower anterior portion. Pereonite 1 with concave rostral (anterior) margin;

pereonites 2-4 about 5x broader than long; pereonites 5 and 6 about 3.7x broader than long. Pleonites

i approximately 4x broader than long, except pleonite 5 about 3.2x broader than long.

First antenna 7-segmented with numerous aesthetascs on segments 4-6; terminal segment with 3

setae.

I Maxilliped similar to those of male species 1 and 2; palpus with second segment bearing 3 setae,

third segment with 4 setae, and fourth segment with 1 subterminal and 4 terminal setae.

I Cheliped similar to male species 2 with carpus about 1.6x broader than long.

• Pereopods 1-3 similar to those in male species 2: tergal spines on merus and carpus short. Pereopod

6 with rows of spinules along length of dactylus.

I with exopodite serrated along outer margin. Pleopods

Pleotelson 1.2x broader than long; posterior projection triangular in outline.

I 3-segmented; Uropodfirst biramous. segmentExopodite 2x longer 1than -segmented, broad with3.2xseveral longerfeathered than broad, hairs; andsecond bearingsegment 3 setae. E2.7xndopodite longer

than broad, with 1 seta and a feathered hair; third segment 2.7x longer than broad, with 5 setae and a

I featheredRehair. marks. Male species 3 can be distinguished from male species 1 by the 1-segmented exopodite

and 3-segmented endopodite of the uropod. Male species 3 differs from both male species 1 and 2 by the

I narrow serrate outer anteriormargin portionof theof the pleopodal cephalothorax, exopodite.the attenuate posterior projection of the pleotelson, and the

Distribution. Santa Mafia Basin, California.

l

i 245
I

i

--r--/j i

_., ,.__: II - I

Uro I

__ _ ,_ -_ J I

4: -___ ,__ J_l _ _" . I

- P.6 "

/_ i

Mxp I

R

I

Figure 3.23. Genus incertae sedis, male species 3: O_3, general habims of male; A.I, first antenna; Mxp, maxilliped;

Che, cheliped; P.1, pereopod 1; P.6, pereopod 6; PI, pleopod; Pit, pteotelson; Uro, uropod. I

_46 I
!

I Subfamily Leptognathiinae Sieg, 1973

I Genus Leptognathia G.O. Sars, 1882 Leptognathia cf. breviremis (Lilljeborg, 1864)

i Figure 3.24

Material Examined. Santa Maria Basin, 2 neuters from phase II, cruise 1-3, Sta. PJ-1, (rep. 3),

34°55.79'N 120°49.91'W, 145 m, off Point Sal; 7 females from cruise 2-4, Sta. R-2 (rep. 1) 35°05.50'N

I 161 southwest of Pt. San Santa Maria California; 1 female from cruise 2-4, 120°53.40'W, m, Luis, Basin,

Sta. R-4 (rep. 2), 34°43.01'N 120°47.39'W, 93 m, off Purisima Point, Santa Maria Basin, California.

i C Description (female) (Figure 3.24). Body, 0.84 - 0.90 mm long, about 4.3x longer than broad. ephalothorax, 1.2x broader than long, with convex lateral margins. Pereonites 1-3 similar in length and

width, 2.6x broader than long; pereonites 4-6 about 2.4x broader than long. Pleonite 1 longest, about 4.3x

i broader than long; pleonite 2 broadest, 6.3x broader than long; pleonites 3 and 4 about 5.8x broader than long; pleonite 5 narrowest, 4.8x broader than long.

First antenna 4-segmented; first segment 1.4x longer than broad; second segment about as long as

I broad; third segment 2.4x broader than long; fourth segment 1.8x longer than broad; formula for armature: 1 seta + 3 feathered hairs, 1 seta + 2 feathered hairs, 2 setae, and 5 setae + 1 aesthetasc.

Mandible with attenuate pars molaris with bifid tip.

I Chelipedal !.9x longer than broad, with 2 small tergal setae (1 proximal, another distal) and

carpus

2 sternal setae. Palm comb consisting of 2 small setae with 1 additional larger seta. Fixed finger with

spiniform tip and several denticles; tergal border with 3 setae; sternal border with 1 long seta. Dactylus

i curved and stout.

Pereopod 1 with merus bearing 1tergal seta; carpus with 1 spiniform tergal seta, with 1 long spinfform

seta and 1 small seta on sternal margin; propodus with 1 tergal spiniform seta and 2 sternal setae; dactylus

i spine as long as propodus. Pereopod carpus bearing spines seta; propodus

and terminal about 6 with 3 and 1

with 2 tergal and 3 sternal spines; dactylus and terminal spine shorter than propodus.

Pleotelson, 1.4x broader than long; posterior margin with protuberant median area bearing 2 pairs

I of setae.

Uropod biramous. Exopodite 1-segmented, 1.8x longer than broad, with 2 terminal setae. Endopodite

I 2-segmented; first segment 1.3x longer than broad, naked; second segment 1.9x longer than broad, bearing

5 setae and 1 feathered hair.

Remarks. The two specimens ofLeptognathia in the MMS voucher collection are both neuters and

I are additional in poormaterial, conditiona. positive Both specimens appear to be related to L. breviremis (Lilljeborg, 1864), but, without identification and detailed description are not possible. We have retained the

original identification of the material as Leptognathia cf. breviremis.

I This species can be distinguished from the other MMS/Santa Maria Basin and Western Santa Barbara Channel tanaidaceans by the combination of a stout body, stout 4-segmented first antenna, attenuate pars

molaris of the mandible, one long and one short sternal setae on carpus of pereopod 1, and an unarmed stout

I first endopodal segment of the uropod.

Type Locality and Type Specimens. Type material was collected in Norway.

i Distribution. The specimens described here were collected from the Santa Maria Basin, California.

l

i 247
I

!

.. I

/ I

I

\ I

/

S I

1

• I

I _ \ \

Pit I

I

I

Figure 3.24. Leptognathia cf. breviremis (Lilljeborg, 1864), female: 9, general habitus of female; A.1, first I

antenna; Md, mandible; Che, cheeped; P.1, pereopod 1; P.6, pereopod 6; Plt, pleotelson; Uro, uropod.

i
!

I Genus Tanaopsis G.O. Sars, 1896

Tanaopsis cadieni Sieg and Dojiri, 1991

I Figures 3.25, 3.26

Tanaopsis cadieni Sieg and Dojiri, 1991:1501-1511, figs. 5-10.

I Material Examined. From Santa Maria Basin and Western Santa Barbara Channel, phase h 9

females, 1 male, 6 neuters, 3 manca-II, 2 fragments from Sta. 67, 34°30'17'N 120°45'30"W, 282 m, off Pt.

I Arguello; 2 neuters, 11manca-II manca-IIfrom Sta. 73, 34°28'13"N 120°36'48"W, 98 m, between Pt. Conception and Pt. Arguello; from Sta. 59, 34°33'39"N 120°47'11"W, 216 m, offPt. Arguello; 5 females from Sta.

71, 34°29'02"N 120°44'01 "W, 306 m, southwest of Pt. Arguello. Phase II: 3 neuters, 9 manca-I from cruise

I Sta. 1-3, Sta. PJ-13PJ-7(rep.(re2), p. 3), 34°56'01"N 34°55'48"N120°49'55"W, 120°48'36"W,144123m,mOctober , May 1987; 1986;21nemanca-I uters, 1from manca-I cruisefrom2-3,cruise Sta. 1-1, R-8

(rep.2), 34°55'18"N 120°45'52"W, 90 m, west of Pt. Sal, January 1988.

I From Santa Moniea Bay, California, collected by Biology Laboratory, Environmental Monitoring Division, City of Los Angeles: 1 male from Sta. B5, 33°58'00"N 118°31'45"W, 45 m, 11 August 1987; 6

females from Sta. E7, 33°54'44"N 118°34'17"W, 150 m, 16 July 1990.

I From LACM: 1 female (LACM 53-125) 6.4 nil., 239° T end of Redondo Beach Pier, 33°47'30"N 118°30'07"W, 229 m, sand and mud, R/V Velero IV (AHF 2361-53), 08 July 1953. From Orange County,

California: 1 female from B44, 33°34'34"N 118°05'22"W, 242 m, Newport Canyon, off Huntington Beach,

i collected Sanitation Districts of 23 September 1983; 3 females and 1 male (1

by County Orange County,

female and 1 male dissected on slide) from Sta. B39, 33°33'18"N 117°58'27"W, 200 rn, Newport Canyon,

off Newport Beach, collected by County Sanitation Districts of Orange County, 18 July 1983; 2 manca-I

I from Sta. 32, 33°34'43"N 117°57'42"W, 59 m, Newport Beach, collected by County Sanitation Districts of Orange County, 21 July 1982; 2 females from Sta. B24, 33°33'36"N 118°01'03"W, 200 m, off Huntington

Beach, collected by Sanitation Districts of Orange County, 03 August 1984; 1 female from Sta. B44,

I 33°34'30"N 118°05'22"W, 242 m, Newport Canyon off Huntington Beach, collected by County Sanitation Districts of Orange County, 23 September 1983; 1 neuter (LACM 62-172) 73 mi 192.5 ° T from Dana Point,

33°20'30"N 117°44'52"W, 719 m, mud, R/V Velero IV (AHF 7728-62), 10 April 1962. From San Diego

I County, California: 1 female CLACM60-85) 1.6 mi, 354.5 ° T from Pt. La Jolla, 32°52'48"N 117°16'32"W, 104 m, muddy silt, R/V Velero IV (AHF 7038-60), 06 May 1960.

Description (female) (Figure 3.25). Body, 3.01-3.97 mm long, about 5.5x longer than broad.

I Cephalothorax about as long as broad; widest in posterior region; posterior border curved. Lateral margins of all pereonites rounded. First pereonite 3.8x broader than long, with rostral border concave. Pereonite 2

about 2.6x broader than tong. Pereonites 2 and 6 about 1.6x broader than long, similar in shape. Pereonites

I 4 and 5 about 1.3x broader than tong. All pereonites similar in width. Pteonites alt of similar size, similar in width to pereonites.

First antenna 4-segmented with armature of segments as follows: 2 setae + 7 feathered hairs; 2 setae

I + 4 feathered hairs; 2 setae + 1 feathered hair; setae, 1 hair,

6 feathered 1 aesthetasc.

Mandible with pars molaris an unornamented spiniform process.

I with 6 spin Firesst.maxilla with strongly curved endite equipped with 3 short rows of minute setules and tipped

Cheliped palm with comb consisting of 1 long and 3 shorter setae. Fixed f'mger with usual seta near

I base of dactylus, terminating in bifid spiniform process; tergal border with 3 setae and 1 spiniform process having a small accessory process at its base; sternal border with 2 setae; rostral surface with longitudinal

ridge extending from articulation point of dactylus to near base of spiniform process. Dactylus crenulated

i along tergal margin, bearing 1 proximal seta, and terminating in heavily sclerotized spine.

i 249
B

Figure 3.25. Tanaopsis cadieni Sieg and Dojiri, 1991, female: Q, general habitus of female; A.1, first antenna; I1

Md, mandible; Mx. 1, first maxilla; Che, cheliped; P. 1, pereopod 1; P.6, pereopod 6; Uro, uropod.

!

250 n
!

Figure 3.26. Tana_psis cadieniSieg and Dojiri, 1991,male: O_, general habitus of male; A. 1,first antenna;Che,

i cheliped; P.I, pereopod 1; P.6, pereopod 6; Uro, uropod.

i 251
!

Pereopod 1 with coxa bearing rostrally-directed triangular process; merus unarmed; propodus N

elongate, about 3.7x longer than broad with 1 tergal and 2 sternal setae; dactylus armed with 1 long seta and

not fused to terminal spine. Dactylus and terminal spine of pereopods 1-3 longer than propodus. Pereopods

4-6 with dactylus and terminal spine combined shorter than propodus. Dactylus without long proximal seta. |

Pleotelson, 1.5x broader than long, with 1 pair of setae at caudal comers, 1 pair subapically, and 2

pairs (1 long, 1 short)apically.

Uropod biramous with stout mmi. Exopodite 2-segmented and extending slightly beyond midlength U

of endopodite. Endopodite 1-segmented, but with constriction (segmental suture ?) at midlength, 3× longer

than broad. I

Deseription (male)(Figure 3.26). Body, 1.49-2.18 mm long, stout. Cephalothorax 1.5x broader

than long, with relatively straight posterior margin. Pereonites and pleonites similar in appearance; pereonites

3-6 slightly longer and a bit narrower than remaining body somites.

m

First antenna 6-segmented with curved rows of numerous aesthetascs on segments 4 and 5; terminal

segment with 4 setae, 1 feathered hair, and 1 aesthetasc. II

Mandible as in female. U

Cheliped palm with comb composed of 11 stiff setae (spines ?) and 2 longer setae. Fixed finger

terminating in simple spine; tergal border serrate. Dactylus with 2 small spines on sternal border. B

I

Pereopod 1 with propodus missing 1 tergal seta; long seta on dactylus absent. Pereopod 6 with

dactylus and terminal spine combined as long as propodus.

as in female, except caudal point more slender and 1 pair of feathered hairs. !

Pleotelson

Uropod as in female, except curved row of feathered hairs near base of endopodite.

Remarks. This species was described by Sieg and Dojiri (1991) based on specimens collected for

the present study and from the Southern California Association of Marine Invertebrate Taxonomists U

(SCAMIT). The specimens had been previously identified by MMS as Cryptocope spp. A and B and by

SCAMIT as "'Leptognathia sp. B" and later as Tanaopsis sp. A. This species can be distinguished from all •

other known California tanaidaceans by its stout body shape, short uropod, and by the detailed morphology II

of the mandible, first maxilla, cheliped, triangular process on the coxa of pereopod 1, and absence of elements

(spines and/or setae) on the merus of pereopod 1. n

This species appears to inhabit substrates consisting of mud and muddy silt.

Type Locality and Type Speeimens. Female holotype (LACM 60-173.2) and 13 female paratypes

(LACM 60-173.3) 2.25 mi 156.5°T (true north) from Catalina Head, Santa Catalina Island, California Channel •

Islands, 33°2Y12"N, t18°29'38"W, 86 m, mud, R/V Velero IV (AHF 6823-60), 27 January 1960. Types m

deposited in the Natural History Museum of Los Angeles County. im

Distribution.

to La Jolla (San Diego), California, 45-719 m (Sieg and Dojiri, i991), n

santa Mafia Basin

1IF

!

!

!

!

252 N
!

I Family Typhlotanaidae Sieg, 1984

I Genus Typhlotanais G.O. Sars, 1882

Typhlotanais wiUiamsae Dojiri and Sieg, new species

i Figure 3.27

I 4 fragments, Type andMater 3 manca-I) ial. 1 female fromhocruise lotype 1-t, (USNM) Sta. PJ-15 and 34(rep. para1), types34°55.80'N (20 females120°50.60'W, [4 ovigerous]155 , 7 m, neutMers, /V

Aloha, October 1986;

I 98 m, originally Material identified Examined.as Santa Leptognathia Maria Basin, sp. H;phase 1 neuter I: 2 neuters from Sta. from58,Sta. 34°73, 34.35'N 34°28.21 120°45.18 'N 120°36.80'W, "_V,99 m,

November 1984; 1 female (ovigerous) from Sta. 71, 34°29.04?4 120°44.01'W, 306 m, originally identified

i asSta.Leptognathia 54, 34°36.57'Nsp. H; 1202°52.02 neuters"_V, from396Sta. m, originally 42, 34°48.40'N identifi120 ed °as47.50'W, Leptognathia 100 m;sp. 2 females H, Marchand1985; 1 neuter vial from with

label reporting 2 females from Sta. 42, 34°48.04'N 120°47.50'W, 100 m, with 1 dried specimen. Santa

I Maria 120°47.40'W, Basin, phase 92 m,II:M42/V neuters, Aloha, May 3 mancas, 1987;and32 females 8 fragments and from 6 neuters cruisefrom1-3,cruise Sta. R-41-2,(rep. Sta.3),PJ-734°43.0 (rep.0'N2),

34°55.79'N 120°48.60'W, 123 m, originally identified as Typhlotanais sp.; 96 females, 4 neuters, 4 mancaI II, as and Typhlotanais 1 manca-Isp.; from25crufemales ise 1-2,andSta.2PJ-7 neuters(rep.from3), 34°55.79'N cruise 1-1, Sta. 120°48.6 PJ-40'W, (rep. 123 2), m, originally originallyidentified identifiedas

Leptognathia sp. H; 16 females, 1 neuter, and 1 manca-I from cruise 1-1, Sta. PJ-6 (rep. 1), 34°54.71'N

I manca-I 120°49.91%V, from 148 cruisem, originally 1-1, Sta. PJ-8 identified (rep. as2),Leptognathia 34°56.80'N 120°49.91'W, sp. H; 14 females, 142 1m,neuter, originally 1 manca-l identiI,fiedandas4

Leptognathia sp. H; 27 females and 2 neuters from cruise 1-1, Sta. PJ-2 (rep. 3), originally identified as

i

• Leptognathia 120°49.90'W, sp. 143H;m,54originally females, 3identified neuters, asandTyphlotanais 1 manca-I fromsp.; cruise 28 females, 1-2, Sta. 1 neuter, PJ-23 and (rep.72),manca-I 34°56.33'N from

cruise 1-2, Sta. PJ-6 (rep. 3), 34°41.40'N 120°57.90'W, 410 m, originally identified as Typhlotanais sp.; 28

i females, 20 neuters, and 1 manca-I from cruise 1-1, Sta. PJ-8 (rep. 1), 34°56.87'N 120°49.91'W, 142 m, originally identified asLeptognathia sp. H; 26 females from cruise 1-1, Sta. PJ-4 (rep. 3), originally identified

asLeptognathia sp. H; 32 females and 7 neuters from cruise 1-2, Sta. PJ22 (rep. 3), 34°55.25'N 120°49.93'W,

I 143 m, oFrom riginaLACM: lly identified 1 female as Ty(LACM phlotana54-94), is sp. 457 m, gravel and mud, 13.1 mi, 301 ° T from North Light,

Santa Barbara Island, Channel Islands, California, 33°35.98%1 119°15.18'W, R/V Velero IV (AHF 2969-

I 54), Pier 31Light, October Los Angeles 1954; 1 female County,andCalifornia 1 neuter, (LACM 33°51.37'N55-35), 11873°34.70 m, 'mud, W, R9.9/V mi, Velero 203 °IVT (AHF from Santa 2998-55), Monica06

February 1955; 1 female (LACM 55-36), 732 m, rock, sand, and mud, 14.3 mi, 345 ° T from Aero Beacon,

i San Nicolas Island, Ventura County, California, 33°28.07'N 119°34.68'W, R/V Velero IV (AHF 3031-55), 07 May 1955; 3 females (LACM 60-77), 415 m, coarse muddy sand, Tanner Canyon, 35.3 mi, 249°T from

China Point Light, San Clemente Island, California Channel Islands, California, 32°35.80'N 119°04.92'W,

I RIV VeIero IV (AHF 6836-60); 1neuter CLACM60-84), 225 m, mud, 34.1 mi, 246 ° T from China Pt. Light, San Clemente Island, Tanner Bank, California Channel Islands, California, 32°34.47'N 119°02.70'W, R/V

Velero IV(AHF 6837-60), 29 January 1960; 1 neuter (LACM 60-186), 814 m, mud, 265 nil, 204°T from

Ribbon Rock, Santa Catalina Island, California Channel Islands, California, 33°22.78'N 118°36.20'W, R/V

i Velero IV (AHF 6829-60), 28 January 1960.

!

I 253
!

Description (female) (Figure 3.27). Body, 1.40-1.62 mm long, about 7x longer than broad. N

Cephalothorax, 1.3x longer than broad, with lateral margins nearly parallel (only slightly convex). Pereonites

1-6 subquadrangular; pereonites 1 and 6 with similar length/width ratio: 1.8x broader than long; pereonites an

2, 3, and 5 about 1.3x as wide as long; pereonite 4 as long as broad. Pleonites 1-5 about 4-5x broader than II

long.

First antenna 3-segmented; fh-stsegment about 2.5x longer than broad; second segment smallest, as D

long as broad; third segment more than 3x longer than broad; formula for armature: 1 seta + 8 feathered

hairs, 2 setae, and 5 setae + 1 aesthetasc + 1 feathered hair.

Mandible with truncate pars molafis; crushing area with crenulated wall. I

u

Maxilliped with basis fused and bearing 1 distal seta each near insertion of palpus. Endites each

with 2 distal setae and 2 bulbs. Palpus 4-segmented; first segment unarmed; second segment with 1 lateral am,

and 3 medial setae (2 of them spiniform); third segment with 3 medial setae; fourth segment with 1 medial •

and 5 terminal inner setae.

Chelipedal carpus slender, about 3x longer than broad (length measured along tergal margin), with iB

2 small tergal setae (1 proximal, another distal), and 3 sternal setae. Fixed finger with spiniform tip; tergal l

border with 3 setae; area near insertion of dactylus comb with 1 small seta and row of spinules; sternal

margin with 2 setae. on

Pereopod 1 with merus longer than carpus and bearing 2 tergal setae; carpus with 2 tergal and 1 U

sternal setae; propodus with 3 sternal setae. Pereopod 6 merus with 2 stout tergal spines; carpus with hooklike

protuberance (indicated by arrow in figure) and 1 sternal seta; propodus with 2 stout tergal spines, a few •

setules along sternal margin, and 3 long sternal spiniform setae at distal end; dactylus with rows of spinules U

along tergal margin, fused with terminal spine.

Pleopods 1-5 similar. Exopodite with proximal pilose seta (indicated by arrow in figure) on outer n

margin. Endopodite with 1 distal seta (indicated by arrow in figure) on inner border.

Pleotelson, 1.8x wider than tong, with rounded protuberance on posterior margin.

Uropod biramous. Exopodite indistinctly 2-segmented, 2.6x longer than broad, 1 seta at midlength, I

and 1 large and 1 small setae terminally. Endopodite indistinctly 2-segmented; first segment, about 2x

Ill

longer than broad, bearing 1 distal seta; second segment about 3x longer than broad, with 1 subterminal seta

and 4 setae and 1 feathered hair terminally, n

Remarks. Typhlotanais wiUiamsae can be distinguished from other Santa Mafia Basin and Western

Santa Barbara Channel tanaid species by the presence of a 3-segmented first antenna in the female, a broad i

crashing area of the pars molaris surrounded by a crenulated walt, a slender cheliped, and normal-sized m

pereonites 1 and 2 (combined longer than pereonite 3). Typhlotanais williamsae differs from T. crassus by

the characters listed in the Remarks section for the latter species.

Etymology. This species is named in honor of Ms. IsabeUe "Izzie" Williams, Woods Hole, MA,

who assisted in putting the finishing touches on this chapter and who critically reviewed the entire manuscript.

Without her efforts, this manuscript would not have been completed. II

Type Locality and Type Specimens. Santa Mafia Basin, California. Types deposited inthe National II

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

"Type Material." I Distribution. Santa Mafia Basin, California to San Clemente Island, California.

N

!

254 N
Figure 3.27. Typhlotanais williamsae Dojiri and Sieg, new species, female: Q, general habitus of female; A.1,

first antenna; Md, mandible; Mxp, maxilliped; Che, cheliped; P.1, pereopod 1; P.6, pereopod 6; P1,

I pleopod; Plt, pteotelson; Uro, uropod.

255
!

Typhlotanais crassus Dojiri and Sieg, new species i

Figure 3.28

I

Type Material. 1 neuter holotype (USNM) and 3 paratypes (neuters; SBMNH) from Sta. D3, i

33°52'47"N 118°35'15"W, 80 m, Santa Monica Bay, California, collected by Biology Laboratory,

Environmental Monitoring Division, City of Los Angeles, 14 July 1988. i

Material Examined. From Santa Monica Bay, California, collected by Biology Laboratory,

Environmental Monitoring Division, City of Los Angeles: 4 females and 1neuter from Sta. D3, 33°52'47"N

118°35'15"W, 79 m, 16 January 1992; 3 females and 2neuters from Sta. D3, 33°52'47"N 118°35'15"W, 80 1

m, 23 January 1993; 1 female from Sta. D3, 33°52'47"N 118°35'15"W, 77 m, 18 July 1991.

Description (neuter) (Figure 3.28). Body, 1.20-1.32 mm long, about 5x longer than broad. i

Cephalothorax about as long as broad, with anterior 2/3 tapered. Pereonites 1-6 with convex lateral margins; i

pereonite 1 about 8.3x broader than long; pereonite 4 longest of pereonites and 1.6x broader than long.

Io

Pleonites 1-5 about 5-6x broader than long. i

First antenna 3-segmented; first segment about 2.4x longer than broad; second segment shortest, i

about as long as broad; third segment almost 4x longer than broad; formula for armature: 6 setae + 5

feathered hairs, 2 setae + 1 feathered hair, and 5 setae + 1 aesthetasc. I

Mandible with truncate pars molaris; crushing area with crenulate wall and spiniform processes. I

Maxilliped with basis fused and bearing 1 distal seta near insertion of each palpus. Endites each

with 2 distal setae. Palpus 4-segmented; first segment unarmed; second segment with 1 lateral and 3 medial I

setae (2 of them spiniform); third segment with 4 medial setae; fourth segment with 1 lateral and 5 terminal II

inner setae.

Cheliped with sternal protuberance, articulating with side-piece. Merus small, with distal margin i

bearing 1 sternal seta. Carpus slender, about 2.2x longer than broad (length measured along tergal margin),

with 2 small tergal setae (1 proximal, another distal), and 2 sternal setae. Fixed finger with spiniform tip;

tergal border with 3 setae; sternal margin with 2 setae. Dactylus as long as f'Lxedfinger and lacking seta. i

Pereopod 1 with basis bearing several (about 4) setae along sternal border; merus about as long as

carpus and bearing 2 setae; carpus with 1tergal and 3 sternal setae; propodus with 3 setae. Pereopod 6 merus is

with stout, pinnate, tergal seta, 1 tergal naked seta (indicated by dotted lines in figure) and several rows of i

minute spinules; carpus with several rows of minute spinules, a hooklike protuberance, and 1 sternal seta;

u

propodus with 2 stout tergal spines, a few setules along sternal margin, and 3 distal spiniform setae; dactylus i

and terminal spine fused; dactylus with minute spinules; terminal spine serrated along tergal margin. !

Pleopods 1-5 similar. Exopodite with proximal pilose seta (indicated by arrow in figure) on outer

margin. Endopodite with distal pilose seta (indicated by arrow in figure) on inner border.

Pleotelson, 1.9x broader than long, with conical protuberance on posterior margin, l

Uropod biramous. Exopodite I-segmented, 4.5x longer than broad, with 1 seta at about midlength

and 1 small and 1 long setae terminally. Endopodite 1-segmented, about 3.8x longer than broad, bearing 5 II

distal setae and 4 feathered hairs. II

Remarks. Typhlotanais crassus differs from T. williamsae by the combination of several characters:

(1) cephalothorax with anterior 2/3 tapered (nearly parallel lateral margins in T. wiUiamsae); (2) convex i

lateral borders of the pereonites (relatively straight in T. williamsae); (3) exopodite and endopodite of l

uropod 1-segmented (indistinctly 2-segmented in T. williamsae); (4) exopodite reaching about 2/3 length of

endopodite (only slightly longer than midlength of endopodite in T. williamsae); and (5) two setae on I

sternal border of propodus of leg 2 unequal in size (these two setae equal in size in T. williamsae). These

specimens were previously identified by SCAMIT as "Leptognathia sp. H" and later as "'Typhlotanais sp. A".

I

256 i
I Figure 3.28. Typhlotanais crassus Dojiri and Sieg, new species, female: Q, general habitus of female; A. 1,first

antenna; Md, mandible; Mxp, maxilliped; Che, cheliped; P. 1, pereopod 1;P.6, pereopod 6; P1,pleopod;

i Pit, pleotelson; Uro, uropod.

i 257
!

Etymology. The specific name crassus is Latin meaning "fat" and alludes to the stout body of this I

species.

Type Locality and Type Specimens. Santa Monica Bay, California. Types deposited in the National m

Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History (SBMNH); see

"Type Material."

Distribution. Presently, this species has been reported only from Santa Monica Bay, California, 1

but may be expected to inhabit the Santa Maria Basin and Western Santa Barbara Channel. m

Family Pseudotanaidae Sieg, 1973 I

Genus Pseudotanais G.O. Sars, 1882 I

Pseudotanais makrothrix Dojiri and Sieg, new species

Figure 3.29 I

Type Material. 1 female holotype 0dSNM) and 1 paratype (SBMNH; dissected female) from Sta. •

4, 35°26.56' N 121°14.93'W, 393 m, off Pt. Estero, Santa Maria Basin, California, March 1985, originally |

identified as Tanaidacea sp. B

Material Examined. 1 female from Sta. BRC-1, 35°27.86'N 121°05.33'W, 98 m, off Pt. Estero, n

santa Maria Basin, California, March 1985, originally identified as Tanaidacea sp. B. II

Description (female) (Figure 3.29). Body about 3.6x longer than broad. Cephalothorax, 1.2x broader

than long, broadest at posterior end, and bearing a pair of eyes. Pereonites 1 and 2 combined shorter than n

pereonite 3; pereonites 4 and 5 longest of pereonites and about 2x broader than long; pereonite 6 with

curved posterior margin. Pleonite 1 longest of pleonites, 3.6x broader than long; pleonites 2-5 approximately

4-5x broader than long. I

II First antenna 3-segmented; first segment long and slender, about 5x longer than broad; second

segment shortest of 3 segments, 2x longer than broad; third segment 4.4x longer than broad; formula for I

armature: 2 setae, 2 setae + 1 feathered hair, and 5 setae + 1 aesthetasc.

u

Mandible with pars molaris whiplike; pars incisiva and lacinia mobilis denticulate.

Maxilliped with basis fused medially arid carrying I seta near insertion of each palpus. Endites m

fused and bearing a pair of short distal setae and a few setules. Palpus 4-segmented; first segment stout and II

unarmed; second segment with 1 spiniform seta and 1 long seta; third segment with 3 spiniform setae and 1

short seta; terminal segment with 1 outer seta near midlength and 5 distal spiniform setae. •

Cheliped of typical shape. Merus triangular and bearing sternal seta. Carpus about 1.5x longer than II

broad, with 1proximal tergal seta, 1distal tergal seta, and 2 sternal setae. Propodus with I seta near insertion

of dactylus and "comb" consisting of about 5 setae. Fixed finger elongate, with 3 distal tergal setae and 1 I

sternal seta. Dactylus with 1 proximal seta and fused terminal spine. |

Pereopod 2 with ischium bearing 1 feathered hair; merus with 2 distal tergal setae; carpus with 1

large spine and 1 large seta on distal tergat margin; propodus with 1 distal tergal seta. Pereopod 6 carpus

with 3 large spines and 1 small, distal, sternal seta. Propodus with 2 terminal spiniform setae, with 1 long II

terminal spine and i shorter spiniform seta.

Pleopods with exopodite and endopodite about 2.4x longer than broad; exopodite with 9 pinnate I

setae; endopodite with 7 pinnate setae. m

Pleotetson about 2.4x broader than long, with protuberant median posterior margin. I

258 I
I

Figure 3.29. Pseudotanais makrothrix Dojiri and Sieg, new species, female: Q, general habitus of female; A. 1,

first antenna; Md, mandible; Mxp, maxiUiped; Che, cheliped; P.2, pereopod 2; P.6, pereopod 6; Pl,

I pleopod; Uro, uropod.

' I 259
!

Uropods biramous. Exopodite indistinctly 2-segmented; first segment with 1 distal seta; second i

segment with 2 distal setae. Endopodite indistinctly 2-segmented; first segment with 1 seta and 1 feathered

hair; second segment with 4 large setae, 1 smaller seta, and 3 feathered hairs, i

Remarks. Pseudotanais makrothrix is characterized by possessing a pair of eyes, typical (not I

modified to bladelike elements) carpal spines on pereopods 2-6, and a whiplike pars molaris of the mandible.

Additionally, the merus of pereopod 4 has two spines of unequal size and the second segment of the •

maxillipedal palpus bears a very long seta on the inner margin. This latter character is shared with the

subantarctic species P. guilIei Shiino, 1978 which was redescribed bySieg (1986a). However, P. makrothrix

may be distinguished from this species by its slender and longer propodal spines, most noticeable on pereopod 1

6: in P. makrothrix these spines are more than half as long as the propodus, but inP. guillei they are less than II

half the propodal length. Also, in P. guillei the third spine (sternal spine) is conspicuously smaller than the

other two, while in P. makrothrix all three spines are similar in size. •

Etymology. The specific name is a combination of the Greek words makros, meaning long or large,

and trichos (=thrix), meaning hair, and alludes to the long and slender propodal spines, most noticeable on

pereopod 6. i

Type Locality and Type Specimens. Off Pt. Estero, Santa Mafia Basin, California. Type deposited

in the National Museum of Natural History (USNM) and the Santa Barbara Museum of Natural History

(SBMNH); see "Type Material." i

I

Distribution. Off Pt. Estero in the Santa Maria Basin, California.

Pseudotanais cahforniensis Dojiri and Sieg, new species I

Figure 3.30, 3.31 I

Type Material. 1 female holotype (USNM) and 31 paratypes (16 females and 15 neuters; 1female

and 1 neuter dissected; SBMNH) from cruise 2-3, Sta. R-5, 34°42.70'N 120°50.80'W, 154 m, southwest of

Purisima Pt., Santa Mafia Basin, California, MIVAIoha, January 1988, originally identified asPseudotanais I

sp. A by J. Toal.

Material Examined. Santa Maria Basin, phase I: 4 females, 1neuter, and 1 fragment from Sta. 3, m

35°27.07'N 121°10.20'W, 291 m, originally identified as Cryptocope sp. A. Santa Maria Basin, phase II: 4 I

females and 1 neuter from cruise 1-1, Sta. PJ 15 (rep. 1), 34°55.80'N 120°50.60'W, 155 rn, M/V Aloha,

October 1986, originally identified as Pseudotanais sp. A by G. Gillingham; 17 females, 4 neuters, 1fragment, •

and 1 male (poor condition) from cruise 2-3, Sta. R-8 (rep. 1), 34°55.30'N 120°45.90'W, 90 m, M/V Aloha, I

January 1988, originally identified as Pseudotanais sp. A by J. Toal; 39 females and 9 neuters from cruise

2-4, Sta. R-4 (rep. 3), 34°43.01 'N 120°47.39'W, 93 m, off Purisima Point, Santa Mafia Basin, California; 16 II

females from cruise 2-4, Sta. R-4 (rep. 1), 34°43.01'N 120°47.30%V, 92 m, off Purisima Point, Santa Mafia I

Basin, California; 32 females from cruise 2-4, Sta. R-5 (rep. 3), 34°42.69'N 120°50.83W¢, 154 m, off Purisima

Point, Santa Mafia Basin, California; 26 females from cruise 2-4, Sta. R-5 (rep. 2), 34°42.69'N 120°50.83'W, i

154 m, off Purisima Point, Santa Maria Basin, California; 1 female from cruise 1-1, Sta. PJ-4 (rep. 3); 32 |

females and 7 neuters from cruise 2-4, Sta. R-4 (rep. 2), 34°43.01'N 120°47.39'W, 92 m, off Purisima Point,

Santa Mafia Basin, California. i

Description (female) (Figure 3.30). Body, 0.90-1.10 mm long, 3.8x longer than broad. I

Cephalothorax, about as long as broad, with tapered anterior end. Pereonites 1 and 2 short, 7.2x and 5.5x

broader than long, respectively; pereonites 3 and 6 about 3x broader than long; pereonites 4 and 5 about •

1.5x broader than long. Pleonites 1-5 approximately 6-7x broader than long. I

First antenna 3-segmented; first segment long and slender, 5x longer than broad; second segment

2x longer than broad; third segment about 3x longer than broad; formula for armature: 2 setae + 2 feathered •

hairs, 1 seta + 2 feathered hairs, and 6 setae (2 have dendriform tips) + I aesthetasc.

i
i Figure 3.30. Pseudotanais californiensis Dojiri and Sieg, new species, female: 9, general habitus of female;

A.1, first antenna; Md, mandible; Mxp, maxilliped; Che, cheliped; P.1, pcreopod 1; P.2, pereopod 2;

I P.6, pereopod 6; P1,pleopod; Uro, uropod.

i 261
!

Mandible with pars molaris whiplike, m

Maxilliped with basis and endites fused, beating 1 pair of small distal setae. Palpus 4-segmented;

first segment unarmed; second segment with 1 outer distal seta and 2 inner distal setae; third segment with i

3 long setae and 1 small seta; terminal segment with 5 long inner setae and 1 smaller outer seta.

Cheliped basis with protuberance. Meres triangular and carrying 1 sternal seta. Carpus about 1.3x

longer than broad with 1 proximal tergal seta, 1 distal tergal seta, and 2 stemal setae. Chela having distinct n

gap between fixed finger and dactylus. Propodus with 1 sternal seta and 1 seta near insertion of dactylus. m

Fixed finger with 3 tergal setae. Dactylus with 1 small tergal seta.

Pereopods 2-6 with modified (bladelike) spine on carpus, i

Pleopods with exopodite about 2/3 length of endopodite and beating 3 terminal pinnate setae;

endopodite with 6 pinnate setae. •

Pleotelson 2x broader than long, with rounded posterior margin. m

Uropod with exopodite indistinctly 2-segrnented; first segment unarmed; second segment with 2

terminal setae. Endopodite indistinctly 2-segmented; first segment with feathered hair; second segment n

bearing 4 setae and 2 feathered hairs.

Description (male) (Figure 3.31). Body 4.3x longer than broad. Cephalothorax about as long as i

broad, broadest in posterior half. Pereonites 1 and 2 similar, 6x broader than long; pereonites 3 and 6 about i

3.6x broader than long; pereonites 4 and 5 about 1.8-1.9x broader than long. Pleonites 1-4 about 4x broader u

than long; pleonite 5 narrowest of all pleonites, about 3.6x broader than long. m

First antenna 7-segmented, bearing numerous aesthetascs on segments 4-6, and carrying 5 setae and i

1 aesthetasc at tip of terminal segment.

Maxilliped with basis and endites completely fused; basis partially fused to cephalothorax, i

Cheliped, with rounded protuberance, articulating with side-piece. Merus with sternal seta at about

midlength. Carpus, about 1.3x longer than broad, bearing 1 proximal and 1 distal tergal seta and 1 long

sternal seta. Propodus with 1 distal tergal seta near comblike row of spinules. Fixed finger with 1 proximal m

seta and 3 tergal setae. Dactylus slightly expanded proximally and attenuate distally.

Pereopod 1 with unarmed merus; carpus with small distal seta. Pereopod 2 with distal seta on

merus; carpus with 1 seta and 1 spine distally. Pereopod 6 ischium with conspicuous spine, i

m

Pteopod with unarmed basis; exopodite and endopodite with several pinnate setae.

Pleotelson triangular, 1.3x broader than long. •

Uropod biramous. Exopodite 2-segmented; first segment with 1 seta; second segment with 2 terminal m

setae. Endopodite 2-segmented; first segment with 1seta and 2 pinnate setules; second segment with 4 setae

and 1 pinnate setule, i

Remarks. The transformed bladelike carpal spines on pereopods 2-6 places this species in the m

subgenus Pseudotanais (Pseudotanais) sensu stricto. Additional important characters are the absence of

eyes, the structure of the pleopods, and the pointed whiplike pars molaris of the mandible. The gap between m

the fixed finger and the articulation of the dactylus in the cheliped indicate that P. californiensis is II

morphologically similar to P. jonesi Sieg, 1977 and P. abyssi Hansen, 1913. All three species also share the

dendritic (or split) setal tips on the third antennal segment. Pseudotanais caIiforniensis can be distinguished m

from these two congeners by the strong distal tergal spine on the propodus of pereopods 2 and 3.

The general outline of this male provides additional evidence that determination of the swimming

male-type specimens may be quite difficult, but not impossible under certain conditions. Firstly, only a i

detailed examination of all apendages will allow the assignment of the specimens under study to the

appropriate family. Secondly, if the male is accompanied by only one species of females of the same genus,

as in the present case of P. caIiforniensis, it is even possible to identify the specimens to species, i

262 i
I

I

I

I

Figure 3.31. Pseudotanais californiensis Dojiri and Sieg, new species, male: C_, general habitus of male; A.1,

first antenna; Mxp, maxilliped; Che, cheliped; P. 1,pereopod 1; P.2, pereopod 2; P.6, pereopod 6; P1,

I pleopod; Plt, pleotelson; Uro, uropod.

I 263
!

The present male does not differ from the general habitus of"swimming males." The second antenna, I

pereopods, pteopods, and uropods provide no special information since they are very similar within the

Anarthruridae, Pseudotanaidae, and Typhlotanaidae. But, the elongate third segment in the 7-segmented i

first antenna, as well as the totally fused maxillipedal endites are characteristic of the Pseudotanaidae. Since II

the cheliped is articulated with the cephalothorax via a side-piece and the chela shows the typical

"pseudotanaid" shape, there is no doubt that the specimen represents a species of Pseudotanais. Finally, the •

male corresponds to the accompanying Pseudotanais females by lacking eyes and having a gap between the n

fixed finger and the dactylus of the cheliped. Therefore, the specimen is identified herein as the male of the

new species P. californiensis.

i

In the case of P. californiensis, the armament of the pereopods is misleading. Pereopod 1 fits the i

general scheme by having no meral spines. However, pereopods 2-6 are misleading. The carpus of these

appendages do not bear the expected transformed spine. All pereopodal segments exclusively bear strong •

spines. Only looking at the pereopods and not knowing the female, the male would have been placed within i

the subgenusAkanthinotanais and notPseudotanais sensu slricto. This shows once more that in "swimming

males" quite often anchestral characters may "reappear" (as the re-segmentation of the uropodal endopodite •

or the 3-segmented uropodai endopodite in some anarthrurid species). i

Finally, the ischium of pereopod 6 bears a spine similar to that of the akanthophoreid male-type 2

(see above). But taking into account the details mentioned above, distinguishing the two species should not i

be difficult.

Etymology. The specific name alludes to the state where the specimens were collected.

Type Locality and Type Specimens. Off Purisima Pt., Santa Mafia Basin, California. Types 1

deposited in the National Museum of Natural History (USNM) and the Santa Barbara Museum of Natural

History (SBMNH); see "Type Material." m

Distribution. Off Pt. Estero, Purisima I>L,Pt. Sal, Santa Maria Basin, California. i

!

!

I

!

!

!

i
I

I Literature Cited

I Barnard, J.L. 1970. Benthic ecology of Bahla de San Quintin, Baja, California. Smithsonian Contributions

to Zoology 44: 1-60.

I Bate, C.S. 1856. On the British Edriophthalma. Report of the British Association for the Advancement of Science 1855:18-62.

Bate, C.S. and J.O. Westwood. 1868. A history of British sessile-eyed Crustacea, volume 2. J. van Voorst,

i I London, 536 pp.

B_ickle-Ramirez, L.F. 1965. Untersuchungen fiber die Biologie vonHeterotanais oerstedi Kr_yer (Crustacea,

i Tanaidacea). Zeitschrift fiir Morphotogie und Okologie der Tiere 55: 714-782. Claus, C. 1888. Ueber Apseudes latreillii Edw. und die Tanaiden, II. Arbeiten aus der Zoologischen

Instituten den Universit_t Wien und der Zoologischen Station in Trieste, 7: 139-220.

I Dana, J.D. 1852. Crustacea. U.S. Expl. Exped. (1838-1842). Vol. 13(2),686-1612. Philadelphia. Feller,

R.J. 1978. Predation on meiofauna established with immunological methods (abstract). American

i Zoologist 18(3): 662. Fryer, G. 1964. Studies on the functional morphology and feeding mechanism of Monodella argentarii

Stella (Crustacea: Thermosbaenacea). Transactions of the Royal Society of Edinburgh 66: 49-90.

I Gardiner, L.F. 1975. The systematics, postmarsupial development, and ecology of the deep-sea family

Neotanaldae (Crustacea: Tanaldacea). Smithsonian Contributions to Zoology 170: 1-265.

I Gerstaecker, A. 1883. Amphipoda - Flohkrebse. In: S.A. Ortmann, ed., Bronns Klassen und Ordnungen des Tierreichs. Die Klassen und Ordnungen der Arthropoden. Abteilung 2: Cmstacea Malacostraca

(1; Auflage), 5(2): 279-543.

I Hansen, H.J. 1895. Isopoden, Cumaceen und Stomatopoden der Plankton Expedition. U. Ordung:

Tanaidacea. Ergebnisse der Plankton-Expedition 2(G.C.), 49-50.

I Hessler, overview. R.R., G.D.Sarsia Wilson64:68-75. , and D. Thistle. 1979. The deep-sea isopods: a biographic and phylogenetic

I Holdich, D.M. and G.J. Bird. 1985. A preliminary report on "dikonophoran" tanaids (Crustacea). Pp. 441-

447, in: L. Laubier and C. Monniot, eds., Pleuplements profonds du Golfe de Gascogne Campagnes.

BIOGAS. - IFREMER Serv. Doc. Publ. de Brest, 630 pp.

I Johnson, S.B. and Y.G. Attramadal. 1982. A functional-morphological model of Tanais cavolinii Milne- Edwards (Crustacea, Tanaidacea) adapted to a tubicolous Me-strategy. Sarsia, 67: 29-42.

i KrOyer, Kudinova-Pasternak, H. 1842. NyeR.arter K. af slagten Tanais. Naturhistoriska Tidsskritt 6: 167-188. 1977. Tanaidacea (Crustacea, Malacostraca) from the deep-sea trenches of the

western part of the Pacific. Trudy Instituta Okeanologii 108:115-135.

I Lang, K. 1949. Contribution to the systematics and synonymies of the Tanaidacea. Arkiv f6r Zoologie

42a(18): 1-14.

I Lang, K.Bertil 1956a. HanstrGm Kalliapseu Zoologi didae, a new family of Tanaidacea. Pp. 205-225 in K.G. Wingstrand, e.d., cal Papers (in Honor of His Sixty-Fifth Birthday). Lund: Zoological Institute.

!

i 265
!

Lang, K. 1956b. Neotanaidae nov. fam., with some remarks on the phylogeny of the Tanaidacea_ Arkiv f6r i

Zoologi, series 2, 9: 469-475.

Lang, K. 1957. Tanaidacea from Canada and Alaska. Contributions du D_partment des Pecheries, Qurbec, i

52: 1-54.

Lang, K. 1960. The genus Oosaccus Richardson and the brood pouch of some tanaids. Arkiv frr Zoologi, •

series 2, 13(3): 77-79. |

Lang, K. 1967. Taxonomische und phylogenetische Untersuchungen fiber die Tanaidaceen, 3: Der Umfang

der Familien Tanaidae Sars, Land und Paratanaidae Lang nebst Bermerkungen fiber den i

taxonomischen Wert der Mandibeln und Maxillulae. Dazu eine taxonomische - monographische

Darstellung der Gattung Tanaopsis Sars. Arkiv frr Zoologi, series 2, 19: 343-368. i

Lang, K. 1968. Deep-sea Tanaidacea. Galathea Report 9: 23-209. I

Lang, K. 1970. Taxonomische und phylogenetische Untersuchungen fiber die Tanaidaceen, 4: Aufteilung

der Apseudiden in vier Familien nebst Aufstellung von zwei Gattungen und einer Art der neuen i

Familie Leiopidae. Arkiv frr Zoologi, 22: 595-626.

Lang, K. 1971. Taxonomische und phylogenetische Untersuchungen fiber die Tanaidaceen, 6. Revision i

der Gattung Paranarthrura Hansen, 19t3, und Aufstellung von zwei neuen Familien, vier neuen U

Gattungen und zwei neuen Arten. Arkiv frr Zoologi, 23:361-401.

Lang, K. 1973. Taxonomische und phylogenetische Untersuchungen fiber die Tanaidaceen, 8. Die Gattungen i

Leptochelia Dana, Paratanais Dana, Heterotanais G.O. Sars und Neotanais Richardson. Dazu

einige Bemerkungen fiber die Monokonophora und ein Nachtrag. Zootogica Scripta 2, 197-229.

p.8 i

Latreille, P.A. 1831. H_t_ropodes (Heteropoda). In: P.A. Latreille, Cours d'Entomologie, ou de l'Histoire

Naturelle des Crustac_es, des Arachnides, des Myriapodes et des Insects. (Accompagn_ d'un Atlas). •

Librairie Encycloprdique de Roret, Paris, 403 pp. |

Lauterbach, K.-E. 1970. Der Cephalothomx yon Tanais cavolinii Milne Edwards (Cmstacea- Malacostmca),

ein Beitrag zur vergleichenden Anatomie und Phylogenie der Tanaidacea. Zoologische Jahrbiicher, i

Abteilung fiir Anatomie und Ontogenie der Tiere, 87: 94-204.

Messing, C.G. 1979. Pagurapseudes (Crustacea: Tanaidacea) in Southereastern Florida: functional i

morphology, post-marsupial development, ecology, and shell use. University of Miami, Ph.D. I

thesis.

Messing, C.G. 1981. Notes on recent changes in tanaidacean terminology. Crustaceana, 41(1): 96-101. i

Milne-Edwards, H. 1840. Isopodes. Histoire naturelte des Cmstacres 3:115-283. pls. 31-33.

Montagu, G. 1808. Description of several marine animals found on the south coast of Devonshire. i

Transactions of the Linnean Society of London, 9:81-114.

Sars, G.O. 1882. Revision af Gruppen: Isopoda Chelifera med. charakteristik af mye herhen hcrende Arter i

og slaegter. Arch. Math. Naturv., 7: 1-54. B

Schram, 1974. Palaeozoic Peracarida of North America. Fieldiana Geol., 33: 95-124.

Schram, F.R., J. Sieg, and E. Malzahn. 1986. Fossil Tanaidacea. Transactions of the San Diego Society of i

Natural History 21(8): 127-144.

!
!

I Sieg, J. 1973. Zum Problem der HersteUung von Dauerpraparaten von Klein - Crustaceen, insbesondere

von Typusexemplaren. Crustaceana, 25(2): 222-224.

I Sieg, J. 1976. Zum natiirlichen System der Tanaidacea Lang (Crustacea, Tanaidae). Zeitschr. fiir Zoologic Systematik und Evolutionsforschung 14: 177-198.

I Sieg, J. Mitteilungen 1977a. Taxonomische aus dem Zoologischen Monographie Museumder Familie Pseudotanaidae (Crustacea, Tanaidacea). in Berlin, 53(1): 3-109.

Sieg, J. 1977b. Eine neue Gattung (Heterotanoides gen. n.) aus der Familie Paratanaidae Lang (Tanaidacea).

I Crustaceana, 33(2): 203-209.

Sieg, J. t980a. Sind die Dikonophora eine polyphyletische Gruppe? Zoologischer Anzeiger 205(5-6):

i 401-416. Sieg, J. 1980b. Taxonomische Monographie der Tanaidae Dana, 1849 (Crustacea: Tanaidacea),

Abhandlungen der Senckenbergischen Naturforschenden GeseUschaft, 537:1-267.

I Sieg, J. 1983a. Evolution of Tanaidacea. Pp. 229-256/n F.R Schram, ed., Crustacean Phylogeny- Crustacean

Issues 1. A.A. Balkema, Rotterdam, The Netherlands, 365 pp.

I Sieg, J. 1983b. Tanaidacea in H.E. Gruner and L.B. Holthuis, eds., Crustaceorum Catalogus. W. Junk Publishers, The Hague, 6: 1-552.

I Sieg, J. Zoologi 1984a. Neuere Erkermtnisse zum natiirtichen System der Tanaidacea. Eine phylogenetische Studie. ca (Stuttgart), 136: 1-132.

Sieg, J. 1984b. Fact documentation and literature database for the crustacean order Tanaidacea. The

I Systematics Association, special volume 26: 125-136.

Sieg, J. 1986a. Crustacea Tanaidacea of the Antarctic and the Subantarctic. 1. On material collected at

i Tierra del Fuego, Isla de los Estados, and the west coast of the Antarctic Peninsula. In L.S. Kornicker, ed., Biology of the Antarctic Seas XVIII, volume 45, Antarctic Research Series. American

Geophysical Union, Washington, D.C., 180 pp.

I Sieg, J. 1986b. Distribution of the Tanaidacea: synopsis of the known data and suggestions on possible distribution patterns. Pp. 165-194 in R.H. Gore and K.L. Heck, eds., Crustacean Biogeography -

Crustacean Issues 4. A.A. Balkema, Rotterdam, The Netherlands, 303 pp.

I Sieg, J. and M. Dojiri. 1989. Remarks on Araphura Bird & Holdich (Crustacea, Tanaldacea) and allied

genera, including descriptions of three new species. Zoologica Scripta, 18(1): 115-137.

I Sieg, J. and Tanaidacea) M. Dojiri. 1991. Two new species and a new genus of the suborder Tanaidomorpha (Crustacea: from Californian waters. Journal of Natural History, 25: 1493-1512.

i Sieg, J. and R.W. Heard. 1983. Distribution patterns of Tanaidacea in the Caribbean Sea and the Gulf of Mexico (Abstract). Association of Southern Biologists Bulletin 30: 225.

Sieg, J. and R.N. Winn. 1981. The Tanaidae (Crustacea; Tanaldacea) of California, with a key to the world

I genera. Proceedings Biological of the Society of Washington 94(2): 315-343.

Siewing, R. 1953. Morphologische Untersuchungen an Tanaidaceen und Lophogastfiden. Zeitschrift fiir

Wissenschaftliche Zoologie, 157: 333-426.

I Viviani, D. 1805. Phosphorescentia maris quatuordecim lucescentium animalculorum novis spiecibus

illustrata. Giossi, Genua, I7 pp.

!

i 267
!

Watling, L. 1981. An alternative phylogeny of peracarid crustaceans. Journal of Crustacean Biology 1(2); I

201-210.

White, A. 1847. List of the specimens of the Crustacea ha the collection of the British Museum - London. I Newman, London, 143 pp.

White, A.vi:1850.CrustaOfTrustees,the cea.ListpecimenSLondon,Of the141BritiShpp, animals in the collection of the British Museum, part I

Wolff, T. 1956. Crustacea Tanaidacea from depths exceeding 6000 meters. Galathea Reports, 2:187-241.

Wolff, T. 1977. Diversity and faunal composition of the deep-sea benthos. Nature, 267:780-785. I

I

I

I

I

I

I

I

I

I

I

I

!

I

268 I
Appendix

Lists and Maps of Stations

Table A.1. Position of soft-substrate stations taken during the Phase I Reconnaissance.

Station Latitude Longitude Depth

(m)

1 35°27.86'N 121°05.33"W 98

2 35%7.70'N 12t °06.52"W 200

3 35°27.07'N 121° I0.20'W 29 !

4 35°26.56'N 121°14.93'W 393

5 35°25.77'N 121°21.69'W 585

6 35°20.88'N 120°59.62"W 109

7 35°20.65'N 121°02.57'W 197

8 35°20.00'N 121°06.58'W 308

9 35°19.48'N 121°10.06'W 398

10 35°18.28'N 121°18.65'W 591

11 35° 17.80"N 121°22.13'W 690

12 35°15.03'N 120°57.31'W 98

13 35° 14.54'N 120°59.77'W 197

14 35°14.15'N 121°02.04'W 299

15 35° 13.98'N 121°04.54'W 393

16 35°12.23"N 121°16.29'W 591

17 35o11.61"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°05.85'3t 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°i 5.99"W 590

27 35°04.30'N 121°19.27'W 611

28 35°04.22"N 121°19.65"W 603

30 34°54.19'N 120°47.07'W 98

31 34°53.76'N 120°52.96'W 200

32 34°53.56'31 120°56.8 I'W 297

33 34°53.43"N 120°59.66'W 396

34 34°53.15"N 121°04.40'W 492

35 34°52.96'N 121°10.30'W 548

36 34°52.77'N 121°15.37"W 492

38 34°49.81'N 120°52.66'W 197

39 34°49.53"N 120°56.85 'W 294

40 34°49.24"N !21 °00.8 I'W 392

41 34°48.35'N 121°19.14'W 495

42 34°48.04"N 120°47.50'W 100

43 34°46.59"N 120°52.92"W 197

45 34°44.9 I'N 120°59.59'W 395

46 34°41.222',I 121o13.56'W 597

47 34041.99'N 121o10.8 I'W 378

48 34045.1 I'N 120°52.85"W 196

49 34°45.03"N 120056.3 I'W 290

50 34°37.80"N 121°01.66'W 591

52 34°39.56"3I 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

269
!

Table A.1 (Continued)

Station Latitude Longitude Depth B

(m)

U

58 34°34.35'N 120°45.18"W 99 n

|

59 34°33.65"N 120°47.18"W 216

60 34°33.25"N 120°48.34'W 275

61 34°33.01'N 120°48.89'W 345

62 34°30.46'N 120°52.13'W 582 •

i

63 34°26.29'N 120°58.08"W 930

64 34°33.15"N 120°40.90'W 59

65 34°31.27'N 120°43.27'W 107

66 34°30.46'N 120°44.55'W 20l •

67 34°30.29'N 120°45.50'W 282

68 34°29.24'N 120°45.99'W 390

69 34°22.88'N 120°54.20"W 927 B

70 34°29.67'N 120°43.70'W 200 •

71 34°29.04'N 120°44.01"W 306

72 34°28.41'N 120°44.76'W 401

73 34°28.2 I'N 120°36.80"W 98 I

|

74 34°26.84'N 120°38.6 I'W 201

75 34°26.08'N 120°39.65'W 293

76 34°25.59'N 120°40.98'W 387

77 34°22.62'N 120°44.02'W 578 •

l

78 34° 18.78'N 120°49.30"W 762

79 34°24.12'N 120°28.32'W 98

80 34°22.86"N 120°28.34'W 196 i

81 34°21.26"N 120°28.83'W 294 •

82 34° 18.7 I'N ! 20°29.55'W 394

83 34° 17.20'N 120°30.20'W 444

84 34° 13.54%1 120°31.19'W 394 lain

85 34°25.88'N L20°16.3 I'W 113 •

86 34°24.45'N 120°17.02'W 197

87 34°21.60"N 120°17.11'W 299

88 34°17.89"N 120°16.86'W 393 •

II 89 34°13.7TN 120°16.56"W 471

90 34°09.44'N 120°16.30'W 375

91 34° 11.73'N 120°07.43'W 540

92 34°08.70'N 120°07.50'W 444 •

l

93 34°07.63'N 120°07.5 I'W 357 i

96 34°22.91'N 120°05.42'W 296

94 34°24.54'N 120°05.47'W 96

95 34"23.70'N 120°05.47"W 198 •

97 34°22.28'N 120°05.49'W 393

98 34°12.87'N 120°05.59"W 561

99 34°11.22"N 120°05.86'W 540 n

100 34°08.67'N 120°05.50"W 443 •

101 34°07.5 I'N 120°05.65'W 357

102 34°59.7 I'N 120°48.22'W 99

103 34°59.63'N 120°53.56'W 197 •

|

104 34°59.45'N 120°56.49'W 294

105 34°59.23'N 120°59.60'W 392

106 34°58.95'N 121°04.42"W 492

107 34°58.65'N 121°15.08'W 573 •

l

108 34°58.21'N 121°17.88"W 492

Note: Sample labels from the Soft-suhstrate stations have several identification codes which include a station number, sample

replicate number, and analysis type. These are as follows: (301 to 200 = the range of station numbers; BSS = Benthic •

Sediment Single (i.e., a non-replicated station); BSR = Benthic Sediment Replicate (three replicates taken at this station); BSV =

Benthic Sediment Variance (subsamples); 01-09 = replicate numbers; TX = a taxonomy sample. Sample labels having the

designation BRA, represents a sample from rocks taken as part of the hard bottom survey, n

270 I
n

I ''2 l _ ESTEFIO I I

• s _""7 _ k _ CALIFORNIA

m ix ° \ I * 20 11 12 ( Pt.BUOHON

• l,, "-, * _

m : 160 15° " ]9-_':t,, "IS _ _ ___.--LUIS 35°10 "N

° °.. _2 RI

i . _- _ _ * (

•,,,, I

,n,c 104 103:': 10(7. I

,. 35 34 " _2 311: ° 30- /Pt.SAL

41 • * T I

• ,_ 4_ /

i ,_ _ " (PURISlMA

IOt'K_ 46 47 49 i lu¢, O"_ '_

I "", • 54 Ae3.,

,. L • _- _ . A:j_FIGUEL&O

m ..... * _oi.59 "

........."..... ,_. __..,_ \ ,_oo.o_T.

m 63 '..e 72 71-......... 08,.5

i " ....... 0e74 •.... 94

_.T: ¢_6 75 '.. 0_ .... .ms ............... Qg_

., ......... .so ......... 200 * 96 .....

--,. 08'7 e97

I ,. *77 *sl

i o78 o82 • 88

/ 083 059

m .,

: 0_4 • 98

91 •

099

-- :)4° lO'N ,"

'" _ ego 92 • ol.O0

i . _ 93 • • I01

/

/ = s_,_St_,_dr=Mooit,,rh_g(L_,a.e_

m Figure A.1. Map showing location of soft-substrate stations from the Phase I Re.connaJssance and Phase 1I

Monitoring Programs.

|

I

m 271
!

Table A.2. Location of soft-substrate stations taken during the Phase II Monitoring Program. I

Station Latitude Longitude Depth i

(m)

i

R-) 35°05.83'N 120049.16'W 91 •

R-2 35°05.50'N 120°53.40"W 161

R-3 35°05.30'N 12l°00.90'W 409

R-4 34°43.0 t'N 120°47.39'W 92 i

R-5 34°42.69"N 120°50.83"W 154 •

R-6 34°4 ) .40'N 120°57.90'W 410

R-7 34°52.90'N 121°10.30'W 565

R-8 34°55.30"N 120°45.87"W 90 •

R-9 34°53.68'N 120059.12'W 410 |

PJ-t 34°55.79"N 120°49.91'W 145

PJ-2 34°55.32'N 120°49.59'W 142

PJ-3 34°56.26'N 120°49.58'W 138 •

l

PJ-4 34°56.26'N 120°50.24'W 150

PJ-5 34°55.32'N 120°50.2A'W 152

PJ-6 34°54.7 I'N 120°49.9 )'W 148 i

PJ-7 34°55.79'N 120°48.60'W 123 •

PJ-8 34°56.87'N ) 20°49.9 I'W 142

PJ-9 34°55.79"N ) 20°51.23"W 169

PJ- 10 34°53.63'N 120°49.9 I'W 147 i

PJ- 11 34°57.95"N 120°49.9 I'W 136 •

i PJ-12 34°55.58"N 120°49.91'W 145

PJ- 13 34°56.01"N 120°49.9 I'W 144

PJ- 14 34°55.79'N 120°49.26'W 134 •

|

P J- 15 34°55.79"N 120°50.57"W 155

PJ- 16 34°55.03'N 120°48.99'W 130

PJ-17 34°56.56'N 120°48.98"W 126

PJ- 18 34°56.56'N 120°50.84"W 158 •

PJ- 19 34°55.03"N 120o50.84'W 167 l

PJ-20 34°50.38'5I 120°49.9 I'W 148

PJ-21 35°01.23'N 120°51.15'W 143

PJ-22 34°55.25'N 120°49.93'W 143 •

PJ-23 34°56.33"N 120°49.90'W 143

!

Table A.3. Sampling dates of MMS Phase l] Monitoring Program. I

Cruise Date i

1-I October 1986

1-2 January 1987

1-3 May 1987 •

2-1 July 1987

2-3 October 1987 |

2-4 January 1988

2-5 May 1988 i

3-1 October 1988

3-4 May 1989 |

!

272 i
Table A.4. MMS Phase I - Locations of hard-substrate transects.

Station Beginning End Depth

Latitude Longitude Latitude Longitude (m)

1 A/B 34°24.454'N 120°01.876"W 34°24.464'N 120°00.878'W 69-73.5

1 C/D 34°24.076"N 120°00.443'W 34°24.184"N 120°01.480'W 73.5-78

2 A/B 34° 11.377'N 120°29.318'W 34° 11.289'N 120°28.774'W 1!0-126

2 C/D 34° 10.984'N 120°28.094'W 34° 10.780'N 120°27.554'W 120-123

4 A/B 34°27.539'N 120°40.364"W 34°28.162'N 120°40.189'W 168-237

6 A/B 34°30.246'N 120°35.555"W 54-63

6 C/D 34°30".421'N 120°3"4.3 I5"W 54-63

13 A/B 34°42".570'N 120°4"7.899'W 34°42.107'N 120°48.253"W 92-100

13 C/D 34°42.556'N 120°48.147'W 34°42.974'N 120°47.424'W 88.5-100.5

14 A/B 34°43.589'N 120°49.093'W 34°42.826'N 120°48.370"W 96-105

14 C/D 34°43.244"N 120°49.406'W 34°42.893'N 120°48.822'W 105-117

16 A/B 34°46.544"N 120°50.197'W 34°45.912'N 120°49.726"W" 91.5-123

17 A/B 34°49.382"N 120°50.768'W 34°49.600'N 120°50.688"W 160.5 - 168

19 A/B 34°47.833'N 120°51.425"W 34°47.097'N 120°50.793"W 148.5-177

20 A/B 34°46.470'N 120°50.289'W 34°46.140"N ! 20°49.885'W 90-130.5

21 A/B 34°47.335"N 120°45.903'W 34°47.548'N 120046.123'W 75-90

22 A/B 34°50.365'N 120°48.221'W 34°50.990"N I20°48.365'W 114-115.5

23 A/B 34°49.868"N 120°47.393'W 34°50.003'N 120°47.480"W 93-102

25 A/B 35°05.662"N 120°47.562'W 35°06.036'N 120°47.652'W 64.5-72

26 C/D 35°11.586'N 120°55.556'W 35°11.555'N 120°55.233'W 108-111

27 A/B 35°20.906'N 120°59.657'W 35°21.035'N 120°59.603'W 96-126

28 A/B 35°21.539'N 120°59.641"W 35021.867'N 120°59.299'W 96-105

29 A/B 35°27.864'N 121°05.331'W 35°27.805'N 121°05.277'W 102-106.5

Table A.5. MMS Phase II - Locations of hard-subslrate photosurvey stations.

Station Latitude Longitude Depth

(m)

PH-E 34°30.26'N 120°42.76'W 119

PH-F 34030.8 I'N 120°42.36'W 105

PH-I 34°29.96'N 120°41.68"W 107

PH-J 34°29.82'N 120041.82'W 117

PH-K 34°29.37'N 120°42.26"W 160

PH-N 34°29.21"N 120°42.05"W 166

PH-R 34°29.11"N 120°42.67'W 213

PH-U 34031.48"N 120043.5 I'W 113

PH-W 34°31.52'N 120°45.86"W 195

273
I

"_ ESTERO I I I

_. ,. _> _o CALIFORNIA _ I

' • 26 Ft. SAN L.UIS

:': 23

!17 D 23 '%_

! ee22 I

-"_-, "_ / I i

I

PH4J '"" :_" I

• _ -'___---_1

_w • _ • "__ " '"...'..2............ 2oo ............................. I

PH-R Q_ PH-N /

34o28,N I

'

I

120 °44'W

i

Figure A.2. Map showing location of hard-substrate stations from the Phase I Reconnaissance and Phase 11

Monitoring Programs. Phase II stations are indicated in the inset. •

!

I

I

274 |
I

I

I

I

I

i

!

i

I

I

The Department of the interior Mission

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 encouragingstewardship and citizen participation in their care.

I The Department also has a major responsibility for American Indian reservation communities

and for people who live in islandterritories under U.S. administration.

The Minerals Management Service Mission

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 Federat and Indian

i __,_/ lands, Moreover, and distribute those revenues. in working to meet its responsibilities, the Offshore Minerals Management Program

administers the OCS competitive feasing program and oversees the safe and environmentally

i sound exploration and production of our Nation's offshore natural gas, oil and other mineral

resources. The MMS Royalty Management Program meets its responsibilities by ensuring the

efficient, timely and accurate collection and disbursement of revenue from mineral leasing and

production due to Indian tribes and allottees, States and the U.S. Treasury.

i The responMMSsivestrives to the topublic's fulfill itsconcerns responsibi andlitinterests iesthroughby the maintgeneral aining guiding a dialogueprinwith ciplesallof:potentially (1) being

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

i development and environmental protection.

I
'l

I

I

I

I

,i

i

I

,I

I

I

I

I

l

I

i

l

i

i