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I OCS Study MMS 96-0031

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I TAXONOMIC ATLAS OF THE BENTHIC FAUNA OF

| THE SANTA MARIA BASIN AND

WESTERN SANTA BARBARA CHANNEL

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I FINAL REPORT

Volume 10 of 14

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The Arthropoda m The Pycnogonida

I The Crustacea Part 1 m The Decapoda

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I U.S. Department of the Interior

MM_ Minerals Pacific OCSManagement Region Service

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I OCS Study

MMS 96-0031

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| TAXONOMIC ATLAS OF THE BENTmC FAUNA OF

THE SANTA MARIA BASIN AND

WESTERN SANTA BARBARA CHANNEL

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I FINAL REPORT Volume 10 of 14

I

The Arthropoda -- The Pycnogonida

I The Crustacea Part I --The Decapoda

I Edited by:

i James A. Blake

Paul H. Scott

Submitted by:

I Science Applications International Corporation

10260 Campus Point Drive

I San Diego, California 92121 For:

I U.S. Min Department of the Interior erals Management Service

Pacific OCS Region

i 770 CamariUo, Paseo California Camarillo 93010

Under Contract No. 14-35-0001-30484

I

Minerals Management Service

I PacificOCS Region July 1996

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DISCLAIMER I

TSheisrvicrepor e, U.S. t hasDepart been rmevent iewedof theby theIntePrior acifiand c Ouapproved ter Continenfortalpubl SheiclfaRtion. egion, ThMeinopinions, erals Mangfindings, ement I

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

nnot ececonsitiute ssarily reflect endorsement the view oorf threcommendation e Minerals Managfor emeuse. nt SeThis rvice.report Mentihas on onot f tradebeennames editeddoesfor I

conformity with Minerals Management Service editorial standards.

I

TAXONOMIC DISCLAIMER

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

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

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i PROJECT ORGANIZATION

I PROGRAMMANAGER

I DR. ANDREWL. L_SSNER Science Applications International Corporation

i San1026Diego, 0 Campus California Point Drive 92121

I DEPUTY PROGRAM MANAGER

I DR.JAMESA. BLAKE

i ENSR Consulting 89 WaterandStreet Engineering

Woods Hole, Massachusetts 02543

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I PROJECT TECHNICAL OFFICER

MR. FRANKJ. MANAGO

I U.S. Department of the Interior

Minerals Management Service, Pacific OCS Region

I 770 Paseo Camarillo

Camarillo, California 93010

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i CONTRIBUTINGAUTHORS

MR. DONALD B. CADmN

I DR.DR.TODJOEL D L.W.Z_ MARTIN

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ACCESS NUMBER: 30484.10

TECHNICAL SUMMARY I

STUDYTITLE: TaxonomicAtlasof the SantaMaria Basinand Western SantaBarbaraChannel. •

REPORTTITLE: TaxonomicAtlasof the SantaMaria BasinandWesternSanta BarbaraChannel. Volume

10.The Arthropoda--The Pycnogonida-- The CrustaceaPart 1.

CONTRACTNUMBER: MMS ContractNo. 14-35-0001-30484. i

SPONSORINGOCS REGION: Pacific. n

APPLICABLEPLANNING AREA: Southern California.

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

COMPLETIONDATEOF REPORT: May 1996. i

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

CUMULATIVEPROJECT COST: $724,384. n

PROJECTMANAGER: Andrew L. Lissner.

AFFILIATION: ScienceApplications International Corporation. n

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

i

PRINCIPALINVESTIGATORS" JamesA. Blake,Andrew L. Lissner. i

KEY WORDS: Marine invertebrates,Arthropoda,Crustacea, Pychnogonida,Decapoda,crabs, shrimp, •

taxonomy,California,SantaMaria Basin,Santa BarbaraChannel,continentalshelf.

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BACKGROUND:The TaxonomicAtlasofthe Santa MariaBasinandwesternSanta BarbaraChannelisan

extensionofthe benthicreconnaissance(PhaseI)and monitoringprograms(Phase II)that wereconducted n

by the MMS since 1983. The organisms that were collectedas part ofthose programsprovidethe material

on whichthe Atlas isdeveloped. In orderto fully documentthe faunacollected by thoseprograms, a series

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

hardandsoftsubstrateenvironments.A team of 40 expertson thefaunahas been assembledto carryout =

this work andtheir contributions are distributedamongthe14volumes.

OBJECTIVES: The objectivesof Volume10 areto continuecoverageof the Crustaceathat was initiatedin B

Volumes11and12. Thisvolumetreatsthepycnogonidarthropodsandbeginstreatmentofthecrustaceans

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withthedecapods.

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DESCRIPTION: Volume 10 includesa treatment of the sea spiders(Pycnogonida)and the shrimpsand B

crabs (Decapoda). The 2 chaptersare organizedinto sectionsthat includethe morphology,taxonomic

history,biology,keysofspecies,and descriptionsofgeneraand species. Eachspeciesisfullyillustratedwith •

relevantcharacteristicslabeledand identified. =

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ACCESS NUMBER: 30484.10

I SIGNIFICANT CONCLUSIONS: Volume10 representsthefirstof three arthropodvolumesand includes

treatment of the Pycnogonida(sea spiders)and the decapod crustaceans(shrimpsand crabs). The

i Pycnogonidsinclude 6 families,8 genera,and 11species. All are previouslyknown. A key has been preparedto these species andothers that are likely to occur in the study area. The author hasprepared an

extensive review of pycnogonid morphology that will assist readers in using the key and interpretingthe

descriptions and illustrations. The chapter onthe decapods includes treatment of 14crab and 11shrimp

B species distributedamong 14families and20genera. One species of commensalcrab, Pinnixiascamit,was

III new to science and describedin an earlier paper. A key is developedto the species occurringin the MMS

collections, but is not inclusivetothose other specieslikely to occur in the area. Each species isdescribed

and illustrated. Bothchaptersincludeanextensivereviewofthe literatureand acomprehensivebibliography.

STUDY RESULTS: Thefauna ofthe Santa MariaBasinand Western Santa Barbara Channel is evaluated

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

,11 sediments using box cores andfrom rocks using manipulatorarms of submersibles and remotelyoperated

am vehicles. The collectionsare organizedinto setsof vouchersfrom Phase I and II that have been made

availableto theteam ofinvestigators,Additionalmaterialfromthe bulkcollectionsnowarchivedwiththeLos

I AnTaxonomic gelesMuAtlas seum(14of Navolumes) turalHismay toryiexceed salsobei1,000 ngexspecies. amined. Thetotalnumberofspeciestreatedintheentire

i The pycnogonids or sea spiders, are familiar, yet enigmatic invertebrates found in epifaunal and cryptic habitats. The present study represents the first comprehensive compilation of the offshore pycnogonid

faunaof the central Californiaregion. The authorhas compileda key to allof the species expected to occur

ii in the area thus ensuring that readers will be able to account for rare species. A total of 11species were

,11 encounteredinthe MMS collections. All were previouslyknown to science. Many ofthe commoncrabs and aB shrimpsencountered in MMS studies have been well known since the monographicworks of Schmidt and

Rathbun. Althoughthe 25species encounteredinthe MMS studies were not a high number for the number

I of in species anearlierexpected paper byinthetheauthors region,ofatthis leastchapter one new. The speci descriptions es,Pinnixiaandscamit illustrations was encountered providedasandpart described of both

chapters will greatly assist users in the identificationof the local arthropods.

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

SantaMariaBasinandWesternSanta BarbaraChannel.Volume10.TheArthropoda-- The Pycnongonida,

TheCrustaceaPart1--Decapoda. A finalreportpreparedbyScienceApplicationsInternationalCorporation,

I fOCS ortheStudy U.S. MMS Departmentof theInterior,MineralsManagementService, PacificOCS Region,Camarillo,CA. 96-0031. Contract No. 14-35-0001-30484.

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I "P.rsaffiliationmaybedifferenthanthatlistedfor ProjectManager(s).

5O272 ° 101

REPORTDOCUMENTATION,. nEPORNTO. MMS 96-0031 =" 3._,_mt's _.,m. No. PAGE

4. 11tie end Subtit_ 11.Report I)ete

TaxonomicAtlasoftheSanta MariaBasinandWesternSantaBarbaraChannel.Volume July 1996

10.The Arthropoda-- The Pychnogonida,The Crustacea Part 1. 6.

7, Author(s) 8. Performinl_Orllanizati_n Rept, No.

JamesA. Blakeand Paul H. Scott (editors)

9. Perfonl_nl[ OrlPlnization Neme lendAddress 10. Pfojectfrask/1Work Unit No.

ScienceApplications InternationalCorporation z:.coertrl_ or Gm_z(No. G)

10260Campus Point Drive

SanDiego, California92121 _c_ 14-35-001-30484

(G)

112.SPonsoringOfilanlZetton Name and Address |3. Type of Report & Period Covered

Final 10of 14volumes,

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

Camarillo,California 93010 z4.

]S. Supp4ementaryNotes

Contributingauthors: Mr. Donald B. Cadien, Dr. Joel W. Martin, Dr. Todd I.Zimmerman.

16. Albstmct(Llm_t: 200 words)

Volume10 representsthe firstof threearthropodvolumesand includestreatment of the Pycnogonida(seaspiders)

and the decapodcrustaceans (shrimpsandcrabs). The Pycnogonids include 6families, 8 genera,and 11species. All

are previouslyknown. A key has beenpreparedto these species and others that are likelyto occur inthe study area.

The authorhas preparedan extensive review of pycnogonid morphology that will assist readers in using the key and

interpretingthe descriptions and illustrations. The chapter on the decapods includes treatment of 14 crab and 11

shrimpspecies distributedamong 14families and 20genera. One species of commensal crab, Pinnixiascamit,was

newto scienceand describedin anearlier paper. A key is developedto the species occurringinthe MMScollections,

but is not inclusiveto those other species likely to occur in the area. Each species is describedand illustrated. Both

chapters include anextensive review ofthe literatureand a comprehensive bibliography.

17. Document Asmlysls a. O_.rlptom

MarineinvertebratesA, rthropoda,Crustacea, Pychnogonida,Decapoda,taxonomy,California,Santa MariaBasin,

Santa BarbaraChannel, continental shelf.

b. IdentifiemfOpen-F.ndedTerms

TaxonomicAtlas, Keys to species, faunal guide, marine ecology, crustaceans, marine benthos.

r. COSATi Field/Group

|L Avalleblllty Statement 19. Security Class (This Report) 21. No, o! PIIses

Unlimited Unclassified 126

29. Security Class (This PaGe) 22. Price

(See ANSl-Z39.18) See fnstructtons on Reverse OPTIONALFORM 27") {4-_*_)

(Formerly NTIS-35)

I Oepartmenta( Cornmtrce
I

Table of Contents [

1

1. StmPm'L_ I_CNOC,_NmA (by Donald B. Cadien)

Introduction ..................................................................................................................................... 1 i

Morphplogy ..................................................................................................................................... 2 [

Ecology and Behavior ..................................................................................................................... 9

Collection and Preservation .......................................................................................................... 12 !

Laboratory Methods ...................................................................................................................... 13 I

Glossary ......................................................................................................................................... t4

Key to the Pychnogonids of California ......................................................................................... 16 •

Description of Species ................................................................................................................... 20 !

Family Ammotheidae ....................................................................................................... 20

Achelia echinata Hodge 1864 ................................................................ 20 Ill

Eurycyde spinosa Hilton 1916a ............................................................. 22 l

Prototrygaeus jordani Child 1990a ....................................................... 24

Family Rhynchothoracidae .............................................................................................. 26 I

Rhynchothorax philopsammum Hedgpeth 1951 .................................... 26

Family Callipallenidae ..................................................................................................... 28

CalIipallene pacifica (Hedgpeth 1939) .................................................. 28 I

Family Nymphonidae ....................................................................................................... 29 1

Nymphon pixellae Scott 1912 ................................................................ 29

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Nymphon heterodenticulamm Hedgpeth 1941a ..................................... 31 i

Family Phoxichilidiidae ................................................................................................... 33 •

Anoplodactylus erectus Cole 1904 ......................................................... 33

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Anoplodactylus oculospinus Hilton 1942d ............................................ 35

Anoplodactylus pacificus Hilton 1942d ................................................. 37 II

Family Pycnogonidae ....................................................................................................... 39

Pycnogonum rickettsi Schmitt 1934 ...................................................... 39 lilt

Literature Cited ............................................................................................................................. 41 i

SUBPHYLUM CRUSTACEA I

2. Tim DECAPODA(by Joel W. Martin and Todd L. Zimmermaa)

Introduction ................................................................................................................................... 49 •

General Biology and Ecology ....................................................................................................... 50 l

Terminology and Glossary ............................................................................................................ 50

Glossary ......................................................................................................................................... 53 i

Institutional Abbreviations ............................................................................................................ 56

Methods of Preservation ................................................................................................................ 57

List of Families, Genera, and Species in the MMS Collections ................................................... 57 _l[

Key to Families and Species ......................................................................................................... 59 l

Descriptions of Species ................................................................................................................. 62

Suborder Dendrobranchiata Bate, 1888 ........................................................................................ 62 •

Infraorder Penaeidea de Haan, 1849 ............................................................................................. 62 n

Family Sicyoniidae Ortman, 1898 ................................................................................... 62

Sicyonia ingentis (Burkenroad, 1938) .................................................... 62 i

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I Suborder Pleocyemata Burkenroad, 1963 ..................................................................................... 64

Infraorder Caridea Dana, 1852 ...................................................................................................... 64

I Family Crangonidae Neocrangon Haworth,zacae 1825(Chace ................................................................................ 1937) ........................................................... 64

Neocrangon resima (Rathbun, 1902) .................................................... 66

l Neocrangon communis (Rathbun, 1899) ............................................... 68

Ii Crangon alaskensis Lockington, 1877 ................................................... 70

Metacrangon spinosissima (Rathbun, 1902) ......................................... 73

i Family AlpheidaeAlpheus Rafinesque, bellimanus 1815 Lockington, ................................................................................ 1877 ................................................... 7575

Family Hippolytidae Dana, 1852 ..................................................................................... 77

I Eualus lineatus Wicksten and Butler, 1983 ........................................... 77 Spirontocaris sica Rathbun, 1902 .......................................................... 80

Spirontocaris holmesi Holthuis, 1947 .................................................... 82

i Family Palaemonidae Rafinesque, 1815 .......................................................................... 84 Pseudocoutierea eIegans Holthuis, 1951 ............................................... 84

Family Axiidae Huxley, 1879 .......................................................................................... 86

Acanthaxius spinulicaudus (Rathbun, 1902) ......................................... 86

I CaIocarides quinqueseriatus (Rathbun, 1902) ...................................... 88

Calocarides sp........................................................................................ 89

Family Diogenidae Ortmann, 1892 .................................................................................. 91

I Paguristes ulreyi Schmitt, 1921 ............................................................. 91

Family Paguridae Latreille, 1803 ..................................................................................... 93

Parapagurodes laurentae McLaughlin and Haig, 1973 ........................ 93

I Galatheidae 1819 95 Family Samouelle,

Munida quadrispina Benedict, 1902 ...................................................... 95

Pleuroncodes planipes Stimpson, 1860 ................................................. 96

i Family Lithodidae Samouelle, 1819 98 Paralomis multispina (Benedict, 1894) ................................................. 98

Family Homolidae De Haan, 1839 ................................................................................. 100

i Paromolafaxoni Schmitt, 1921 100 Family Xanthidae MacLeay, 1838 ................................................................................. 103

Lophopanopeus beUus diegensis (Rathbun, 1900) .............................. 103

i Family Parthenopidae Alcock, 1895 105 Heterocrypta occidentalis (Dana, 1854) .............................................. 105

Family Majidae Samouelle, 1819 .................................................................................. 107

I Erileptus spinosus Rathbun, 1893 ........................................................ 107 Family Pinnotheridae De Haan, 1833 ............................................................................ 110

Pinnixa occidentalis Rathbun, 1893 .................................................... 110

i Acknowledgments Pinnixa scamit Martin and Zmarzly, 1994 .......................................... 112 ....................................................................................................... ............... 114

Literature Cited ........................................................................................................................... 114

I APPENDIX..................................................................................................................................... 121

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List of Figures !

Figure 1.1. Pycnogonid external morphology; A. body, B. second and third coxae, and femur of leg n

(after Child, 1979) .............................................................................................................. 3

Figure 1.2. Appendages of the cephalon; A.-D. chelae, E. and F. palpi, G. and H. ovigers ................ 5 I

Figure 1.3. Location and extent of gonads in; A. male (after E1-Hawawi and King, 1978), and B. female

Pycnogonum littorale (after Jarvis and King, 1972) .......................................................... 8 1

Figure 1.4. Protonymphon larva ofAchelia echinata (after King, 1973) ............................................. 9 U

Figure 1.5. Achelia echinata; A. body, dorsal, B. terminal claw, C. female oviger, D. juvenile chela, •

and E. adult chela (all after Utinomi, 1954) ..................................................................... 21 il

Figure 1.6. Eurycyde spinosa; A. body, dorsal (after Hedgpeth, 1941b), B. chelifore, C. proboscis, D.

palp, E. oviger, and F. terminal claw ............................................................................... 23 HN

Figure 1.7. Prototrygaeus jordani; A. body, dorsal, 13.terminal claw, C. palp, and D. oviger (after

Child, 1990) ...................................................................................................................... 25 1

Figure 1.8. Rhynchothoraxphilopsammum; A. body, dorsal, B. palp, C. oviger, D. terminal claw (after

Hedgpeth, 1951) ............................................................................................................... 27

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Figure 1.9. CallipalIene pacifica; A. body, dorsal, B. chelae and proboscis, C. oviger, D. terminal claw H

(after Hedgpeth, 1939) ..................................................................................................... 29

Figure 1.10. Nymphon pixellae; A. body, dorsal, B. oviger, C. palp, D. chela, E. terminal claw (after n

Hilton 1942a) .................................................................................................................... 31 .I

Figure 1.11. Nymphon heterodenticulatum; A. body, dorsal, B. chela, C. oviger, D. terminal claw (after Ill

Hedgpeth 1941a) .............................................................................................................. 32 Ii

Figure 1.12. Anoplodactylus erectus; A. body, lateral, B. chela, C. oviger, D. terminal claw, E. leg (after

Cole, 1904) ....................................................................................................................... 35 i

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Figure 1.13. Anoplodactylus oculospinus; A. body, dorsal, B. chela, C. terminal claw, D. leg (after Child,

1975) ................................................................................................................................. 36 I

Figure 1.14. AnopIodactylus pacificus; A. body, dorsal, B. chela, C. oviger, D. terminal claw (after

Child, 1975) ...................................................................................................................... 38

m

Figure 1.15. Pycnogonum rickettsi; A. habitus, dorsal, B. body, lateral, C. oviger, D. leg (after Schmitt, n

1934) ................................................................................................................................. 40

Figure 2.1. Schematic drawing of shrimp in lateral view: ai, appendix interna; as, antennal spine; asc, H

antennal scale; b, basis; bs branchiostegal spine; cp, carpus; cx, coxa; d, dactyl; end, endopod;

ep, epipod; ex, exopod; hs, hepatic spine; i, ischium; m, merus; p, propodus; pg, postorbital

groove; ps, pterygostomian spine; ss supraorbital spine; st, stylocerite (from Williams 1984, I

modified from Holthuis 1955; Schmitt 1921) .................................................................. 51

Figure 2.2. Schematic drawing of brachyuran crab in: A--dorsal view; areas of carapace indicated; legs n

of right side only shown; b, basis; cp, carpus; d, dactytus; i,ischium; m, merus; p, propodus. l

B--ventral view; areas of carapace indicated; legs of right side only shown; b, basis; cp,

carpus; cx, coxa; d, dactyl; end, endognath; ex, exognath; i, ischium; m, merus; p, propodus I

(from Williams, 1984) ...................................................................................................... 52 •

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I Figure 2.3. Sicyonia ingentis (Burkenroad, 1938): A, lateral view; B, dorsal view of petasma; C,

thelycum; scale for B = lmm, scale for C = 2mm (after Perez-Farfante, 1988) ............. 63

I 2.4. (Chace, 1937): A, Female lateral view; B, female

Figure Neocrangon zacae holotype, holotype,

dorsal view of carapace; C, paratype, antennal scale; D, paratype, chela of first pereiopod

(from Chace, 1937) .......................................................................................................... 65

I Figure 2.5. Neocrangon resima (Rathbun, 1902): A, lateral view of carapace; B, antennal scale; C,

anterodorsal view of carapace; D, chela (A from Word and Charwat, 1976; B-D from

I Rathbun, 1904) ................................................................................................................. 67 Figure 2.6. Neocrangon communis (Rathbun 1899): A, dorsal and lateral views (from Butler 1980); B,

antennal scale; C, anterolateral view of carapace; D, anterodorsal view of carapace; E,

I chela (b-e from Rathbun, 1904); F, anterodorsal view; G, lateral view (F and G from Schmitt, 1921) ................................................................................................................................. 69

i Figure 2.7. Crangon view of carapace; alaskensisC, Locldngton, dorsal viewI877: of carapace; A, lateral D, viewantermal of ovigerous female; B, anterolateral scale; E, third maxilliped; F,

chela. Original illustration from specimen taken from Los Angeles Harbor ................... 72

I Figure 2.8. Metacrangon spinosissima (Rathbun, 1902): A, dorsal and lateral views (from Butler, 1980); B, dorsal view of carapace (from Rathbun 1904); C, lateral view (from Rathbun, 1904); D,

dorsal view of carapace; E, anterolateral view (D and E from Word and Charwat, 1976) ... 74

Figure 2.9. Alpheus bellimanus Lockington, 1877: male, carapace length 5.2 ram; A, anterolateral view;

B, anterodorsal view; C, carina below fit-st antennular segment; D, right third maxilliped;

i E, outer face of major first pereiopod; F, inner face of same; G, outer face of minor first pereiopod; H, inner face of same; I, left second pereiopod; J, right third pereiopod; K,

abdomen; L, telson and uropods; M, outer face of minor chela from first pereiopod of

i ovigerous female (carapace length 6.5mm); N, right second pleopod (female, carapace length 4.6 mm)(from Kim and Abele, 1988) .............................................................................. 76

Figure 2.10. Eualus lineatus Wicksten and Butler, 1983: A, dorsal and lateral views; B, anterolateral

I view of female (carapace length 3.1 mm); C, left third pereiopod of same; D, dactyl of left third pereiopod of same; E, appendix masculina of male (carapace length 4.1 mm) (from

Wicksten and Butler, 1983) .............................................................................................. 79

I Figure 2.11. Spirontocaris sica Rathbun, 1902: A, dorsal and lateral views; B, anterolateral view (A,

from Butler, 1980; B, from Word and Charwat, 1976) .................................................... 81

I Figure 2.12. Spirontocaris from Butler, 1980; holmesi B, Holthuis, from Word1947: and A, Charwat, dorsal and 1976)lateral .................................................... views; B, anterolaterat view (A, 83

Figure 2.13. Pseudocoutierea elegans Holthuis, 1951: A, anterolateral view; B, third pereiopod (both

I from Holthuis, 1951) ........................................................................................................ 85

Figure 2.14. Acanthaxius spinulicauda (Rathbun, 1902): A, male, lateral view; B, carapace dorsal view;

I C, 1982, female, see text; fightB,Echeliped; from Rathbun, D, male, 1904) fight cheliped; ......................................................................... E, tail fan. (A,C,D modified from Hart, 87

Figure 2.15. Calocarides quinqueseriatus (Rathbun, 1902): A, male, lateral view; B, male, fight cheliped;

I C, male, dorsal view; D, tailfan Hart, 89 carapace, (from 1982)

Figure 2.16. Calocarides sp. A.: A, lateral view; B, dorsal view of rostrum and anterior carapace; C,

I lateral and leftviewuropodof rostrum ................................................................................................................. and anterior carapace; D, right cheliped; E, left cheliped; F, telson90

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Figure 2.17. Paguristes ulreyi Schmitt, 1921: A, male, dorsal view; B, telson (from Hart, 1982) ..... 92 i

Figure 2.18. Parapagurodes laurentae McLaughlin and Haig, 1973: male paratype - A, shield; B, mesial In

view of right cheliped: C, dorsal view of right cheliped: D, mesial view of left cheliped i

(modified from McLaughlin and Haig, 1973) ................................................................. 94

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Figure 2.19. Munida quadrispina Benedict, 1902: male, dorsal view (from Hart, 1982) .................... 96 I

n

Figure 2.20. Pleuroncodesplanipes Stimpson, 1860: male, dorsal view (from Boyd, 1967) ............. 97

Figure 2.21. Paralomis multispina (Benedict, 1894): male, dorsal view (from Hart, 1982) ............... 99 i

II

Figure 2.22. Paramola faxoni (Schmitt, 1921): Holotype female (carapace width 39.60 ram) USNM

53331, dorsal view (from Kuck and Martin, 1994) ....................................................... 102 II

Figure 2.23. Lophopanopeus bellus diegensis Rathbun, 1900: A, anterodorsal margin of carapace; B, m

male, dorsal view; C, male, lateral view of fight cheliped; D, male holotype, dorsal view of

right carpus; E, male holotype, lateral view of second ambulatory leg; F, male holotype, I

lateral view of fourth ambulatory leg (A, from Allen, 1980; B,C, from Hart, 1982; D-F, l

from Menzies, 1948) ...................................................................................................... 104

Figure 2.24. Heterocrypta occidentalis (Dana, 1854): A, dorsal view; B, male, f'n'st pleopod; C, male, I

second pleopod (A, from Allen, 1980; B-C, from Garth, 1958) .................................... 106

Figure 2.25. Erileptus spinosus Rathbun, 1893: A, male; B, female (A and B after Rathbun, 1904, as i

Erileptus spinosus and Anasimus rostratus respectively) .............................................. 109 IN

Figure 2.26. Pinnixa occidentalis Rathbun, 1893: A, anterodorsal view of male; B, anterodorsal view of

carapace (juvenile male); C, anterior face of female cheliped; D, anterior face of male

cheliped; E, abdominal face of fight gonopod (A-E from Zmarzly, 1992) ................... 111

n

Figure 2.27. Pinnixa scamit Martin and Zmarzly, 1994: USNM 267500, and USNM 267501 (C only), III

western Santa Barbara Channel, California. A, female holotype, dorsal view of carapace II

and right side appendages, pereiopods 1-3 drawn in situ, pereiopods 4 and 5 removed and

figured separately (to enssure correct proportions): B, same specimen, frontal view showing •

sculpturing of carapace and subhepatic tooth (arrow): C, dorsal view of immature paratype; !

note that subhepatic tooth (arrow) and acute serrations along anterolateral margin are already

present at this size: D, third maxilliped of holotype, left side, outer view; E, abdomen of fl

holotype, ventral view illustrated in situ (first two segments not visible, and segments not n

visible, and segment 3 shortened due to curvature of abdomen). Scale bar = 2.0 mm for AC, E; 1.0 mm for D (from Martin and Zmarzty, 1994) .................................................. 113 i

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I Acknowledgments

i This study was funded by the U.S. Department of the Interior, Minerals Management Service,

Washington, D.C., under Contract No. 14-35-0001-30484. Ms. Marie Murphy of the Santa Barbara Museum

I of Natural History provided production assistance for this volume.

I List of Acronyms

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ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania.

I BLM Bureau of Land Management.

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

transect A/B.

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

transect C/D.

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

California, USA.

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

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

I MMS Minerals Management Service.

NHML The Natural History Museum, London, United Kingdom.

I SCAMIT Southern California Association of Invertebrate Taxonomists.

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

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

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

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

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1. SUBPHYLUM PYCNOGONIDA

| by

I Donald B. Cadien _

i Introduction

The pycnogonids (or sea-spiders) are a small group of cheliceriforms with about 1,100 species

I world-wide. They are exclusively marine, although some species are tolerant of reduced salinities. Pycnogonids can be found from the intertidal zone to depths of nearly 7,400m. They have a long evolutionary

history, but the fragility of the pycnogonid body (or failure to recognize members of the group) has left this

I history poorly documented in the fossil record. They are only distantly related to the true spiders (Arachnida), having diverged from a common ancestor prior to the Carboniferous (Manton, 1978). One group of

pycnogonids, the Pantopoda, is extant, and has Devonian fossil representatives similar to the living genus

I Pig extinct. rogromitus (Arnand and Bamber, 1987). Two other groups, the Paleoisopoda and Paleopantopoda, are

Of the approximately 44 species of pycnogonids which are known to occur in or adjacent to the

I study area, only a few are routinely taken on soft substrata. The vast majority of pycnogonids are found on hard substrates, or on the epibiota which occupies them. At least one genus, Rhynchothorax, has interstitial

members (Child, 1988).

I are separate pycnogonids, although hermaphroditism reported one species.

Sexes in has been in

Sexual dimorphism is usually manifested in presence or absence of ovigers; in differences in oviger size,

segmentation, or ornamentation; and in number, shape, and position of the gonopores. Gynandromorphy is

I also in the literature (Child, 1978; Child and Nakamura, 1982; and Makioka, Both

reported Miyazaki 1993).

sexual mosaics and bilateral gynandromorphs have been found.

i The higher classification of animals of the arthropod grade is still unsettled. Definitive determination of the position of the Arthropoda may always remain a matter of conjecture. Although there is little controversy

about the relatedness of the various taxa with jointed appendages encased in a chitinous exoskeleton

(arthropods), there are two clear lines of thought as to their status. One holds that the concept of"arthropod"

I embraces a series of extant and extinct phyla and, if used at all, must be accorded the level of Superphylum.

Willmer (1990), for instance, considers that "arthropodization" has occurred in a number of related, but

independent evolutionary lines. The other maintains that, while there is considerable diversity among

I "arthropods", they a common ancestor belong a phylum taxon, adopt view

share and within level Here I the

that embryological and morphological evidence available (summarized in Schram, 1986) supports phylum

status for each of the three major extant divisions within the arthropod grade. Of these the Phylum

I Cheliceriformes has two the Chelicerata and the For within

subphyla Pycnogonida. organizational purposes

the Atlas, these animals have been included with the arthropods.

I

I t Marine Biology Laboratory, Los Angeles County Sanitation District, 24501 S. Figueroa St., Carson, CA 90745 USA
!

It must be emphasized, however, that the controversy over a monophyletic or polyphyletic Arthropoda I

still rages. Brusca and Brusca (1990) provide cogent summary of the issue (and support monophyly), based

on much the same evidence used by Schram (1986), who supports polyphyly. Even more recent phyletic

reanalyses (Eemisse et al., 1992; and Backeljau et al., 1993) of a common character set assembled by II

Schram (1991), while yielding congruent consensus trees, were interpreted as evidence of arthropod polyphyly

and monophyly, respectively. I

il Knowledge of the natural history, ecology, functional morphology, and physiology of pycnogonids

is relatively recent. Until the 1960' s, most reports of pycnogonids were restricted to either anatomy, taxonomy,

or biogeography. Two recent comprehensive reviews of the Pycnogonida are available (King,1973; Arnaud I

and Bamber, 1987) and should be consulted for additional information. M

Although opinions differ, an arrangement of the pycnogonids into nine families is generally used.

Fry (1978) attempted a radical reorganization of the group based on application of numerical taxonomic I

methods, but his system has received little acceptance. There is only one extant order, the Pantopoda, i

containing the families Endeidae, Pycnogonidae, Phoxichilidiidae, Colossendeidae, Nymphonidae,

Callipallenidae, Ammotheidae, Austrodecidae, and Rhynchothoracidae.

W

Morphology I

The first impression of most people upon viewing a pycnogonld is of appendages, but no body. So I

decentralized is the appearance of sea spiders that the Reverend T. R. R. Stebbing used the term "nobodies" I

as a popular name for the group (Stebbing, 1902). Ignoring appendages for the moment, the basic pycnogonid

plan consists of a body divided into four sections - the proboscis, the cephalon, the thorax (or trunk), and the !1

abdomen (Figurel. 1A). The proboscis is a tube or cone-like anterior projection on the front of the animal. It I

is attached to the cephalon, or "head", which is the first section of the body proper. The cephalon is the

attachment site of the chelifores, palpi, and ovigers, and also bears a dorsal ocular tubercle. The f'trstpair of m

walking legs are attached near the posterior margin of the cephalon. The next three (or atypically four or I

five) body segments form the thorax or trunk. Each bears one pair of walking legs, but no other appendages.

The final body segment is the abdomen, which is attached terminally on the last thoracic segment. This

segment is often flexed dorsally, especially if it is elongate. I

Proboscis I The proboscis is perhaps the most unique feature of the group, being present in one form or another

in every pycnogonid. It is the feeding organ of sea-spiders, and its form reflects the type of prey each animal

consumes. It is essentially a muscular tube projecting anteriorly or ventrally from the cephalon (Figure

1.1A). The proboscis bears no appendages, but is terminally equipped with small lappet-like lips. Proboscis II

shape is generally species specific (although either shape or relative size of the proboscis changes with

growth in some species), and particular proboscis arrangements may be characteristic at the family level.

The proboscis is attached to the underside or anterior face of the cephalon by an arthrodial membrane of II

varying width. This is also termed the soft collar, and its width is generally an indication of the degree of

mobility of the proboscis; species with narrow soft collars having nearly immobile proboscides.

!1 Proboscis shape can be complex. The organ is never just a simple tube, being either tapered, bent, or

with some complex of bends, swellings, and tapers over it' s length. Proboscis length relative to the remainder

of the body also varies widely within the group, from a mere rounded button on the cephalon to a long stylus I

nearly the length of the body. Fry and Hedgpeth (1969) introduced a system of expressing proboscis shape

which, while generally accepted, is often not included with descriptions.

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1 FiN 1.1, PycnogonidChild, 1979).externalmorphology; A. body, B. second and thirdcoxae, and femur of leg (after

i

Cephalon

I palpi, theTheovigers, proboscis and istheattafirst chedsettooftheambulatory cephalon, legs. as areDepending the first foonur sets species, of append age, orages; sex the of anchelifores, animal, one th

or more pairs of appendages may be missing. These appendages are distributed around the attachment of the

i proboscis. surface. TheThepalpi chelifores originatearelateral dorsal,tooverhanging the proboscistheinsertion, proboscisand(Figure extend 1.2A) anteriorly or extending along itsalong sides.itsOvigers upper

are ventrally attached to the cephalon anterior to the attachment of the legs. The legs attach laterally on the

i cephalon, posterior to the insertion of the proboscis. The cephalon usually bears an ocular tubercle dorsally,

I
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on which the eyes are situated. The position of this tubercle varies from the anterior margin of the cephalon I

to near the posterior end of the segment. It is anocular in some deep-sea and interstitial forms, but usually

bears two pairs of eyes distally. These pairs are usually equal in size, but in some forms the anterior pair is •

enlarged. The ocular tubercle is usually of moderate size (three to six times higher than broad), but ranges

from a barely perceptible bump on the cephalon to a tall turret raised high above the segment surface.

Cephalon shape varies from compact to attenuated. The insertion of the proboscis is close to that of

the walking legs in the former, while in the latter the proboscis, chelae, and palpi are situated at the distal

end of an extended neck. A further variation is seen in the local species Prototrygaeusjordani, in which the

ocular lobe is extended forward and overhangs the insertions of the proboscis, palpi, and chelifores. N

I

The chelifores are composed of a basal scape of one or two segments and a terminal chela which

bears one moveable and one fixed f'mger (Figure 1.2B and D). Functional chelae are always longer than the

proboscis, and are assumed to be involved in transferring food to the mouth. Chelifores are present in all i

pycnogonid larvae. In the Ammotheidae, chelae are fully developed in juveniles, but become reduced or

I

vestigial in the adult (Figure 1.2C). In the Austrodecidae, Colossendeidae (with the exception of Decolopoda),

Pycnogonidae, Rhynchothoracidae, and Endeidae chelifores are present in juveniles and absent in adults, i

Members of the Callipallenidae, Nymphonidae, and Phoxichilidiidae have well-developed chelae both as IB

juveniles and adults.

i

The palpi are composed of between one (Pallenopsis) and twenty (Nymphonella) segments, although g

most palps have between five and ten segments (Figure 1.2E and F). Their function is assumed to be sensory,

m

and they bear sensory hairs, especially on their terminal segments. InAchelia echinata they appear to assist

in orientation of the proboscis to the ectoproct prey (Wyer and King, 1973). As with chelifores, palpi are not i

found in all families. Palpi are present in Ammotheidae, Nymphonidae, Rhynchothoracidae, and m

Colossendeidae; either present or absent in Callipallenidae; and absent in the remaining families. In the i

Ammotheidae the number of palp segments varies considerably, but is always at least four. Colossendeids

have eight or nine, nymphonids five, and callipallenids four or fewer palp segments. In

Ovigers (Figure 1.2G and H) are specialized cleaning appendages which have become secondarily In

modified in the males of some species into egg carriers. In the Pycnogonidae, Endeidae, and Phoxichilidiidae i

they are found only on males, while they occur on both sexes in the remaining five families. They may bear

a brush-like elaboration of the last few segments termed a strigillus, which is of diagnostic value, as are both

the number of oviger articles and their proportions. •m

Thol'8.x I The thorax typically consists of three segments and their associated walking legs. The segments are

joined along their midline and separate towards the side of the body into lateral processes. These are closely

approximated in some species, forming a disc with barely perceptible gaps between adjacent processes, i

More often, however, they are broadly separated so that the lateral processes appear to be the bases of the i

legs rather than extensions of the body. In some forms the cuticle of adjacent segments is fused in the adult,

obscuring their articulation, i

In a number of species from Antarctic seas (and a few from the Caribbean), there are four or five

thoracic segments, each beating a pair of walking legs. When first found, such forms were considered

aberrant, and some individuals with extra legs have been reported in species normally having three thoracic •

segments (Arita, 1936). Additional observation has shown that most specimens with extra legs are not II

aberrant, although the co-occurrence of very similar three segmented species nearby remains suggestive.

Ten or twelve-legged species have not been reported from the northeastern Pacific. i

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i Figure 1.2. Appendages of the cephalon; A.-D. chelae, E. and F. palpi, G. and H. ovigers.

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Thoracic legs are invariably nine-segmented. This is also true of the cephalic legs with the exception

I ofmultisegmented the genus Nympho (Ohshima, nella, 1927). whereThethe segments last two are leg (sequentially segments of from the first the lateral walkingprocess) legs arethreethemselves coxae, a

femur, two tibiae, a tarsus, a propodus, and a terminal claw (Figure 1.1A). In species with short legs (about

i equal to thorax width) none of these leg segments are elongate. In long-legged species, the added length results from elongation of the femur and/or tibiae, with the other segments remaining relatively short.

Segment widths do not normally differ greatly in any given leg, with the widest segment typically being the

i femur.

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Abdomen i

The abdomen is terminally attached to the posterior thorax. Its articulation may be readily apparent,

or obscure so that the abdomen appears fused to the last thoracic segment. The abdomen itself terminates in 1

the anus. There are no abdominal appendages. i

The Integument I

The pycnogonid external surface is composed of a chitinous cuticle overlying a layer of epithelium.

Unlike the exoskeleton of crustaceans, the cuticle of pycnogonids is never stiffened by calcium deposits I

(Arnaud and Bamber, 1987). The cuticle is perforated by pores providing exits to cutaneous glands for !I

mucous production and excretion. It is composed of three layers: the epi-, ecto-, and endocuticles. At points

of flexion, such as joint articulations, the cuticle is thinned by loss of the ectocuticular layer (King, 1973). i

All organisms with exoskeletal structures must either modify or shed these in order to accommodate

soft tissue growth. In pycnogonids, the cuticle is shed in sections rather than as a single piece as in crustaceans.

During molting, the cuticle of each leg separates from the body and is shed as a sleeve, while that coveting i

the body splits laterally, and is shed as dorsal and ventral halves (Lotz and Biickmann, 1968). 1

Sensory Organs and Nervous System i

Although it may be inferred with some confidence that at least some of the hairs and setae of the

palpi are related to environmental sensing, such function has only been demonstrated for the eyes. These are 1

simple, with a cupped retina, a single lens, and a layer of photoreceptive cells. They are light sensitive, but I

not image-forming. There appear to be no other light sensitive areas on the body, as loss of the ocular

tubercle results in unresponsiveness to light (Cole, t901). A number of deep-water species have the eyes 1

reduced or lacking, as do some true infaunat forms such as Rhynchothorax philopsammum (Krapp, 1973). I

Chemoreception has been demonstrated for pycnogonids, but has not yet been associated with

particular structures. Ablation of chelifores, patpi and ovigers did not remove chemosensitivity. Innervated i

epidermal setae were suspected of being the site of chemosensory localization (Stock, 1978). Such structures il

occur on both the proboscis and on walking legs. Bifid setae, hollow setae with terminal pores, and setae set

into cuticular cups are seen on various pycnogonids. All are basally innervated, and all certainly serve either I

a mechanosensory or chemosensory purpose. Either chemoreception or thermoreception has also been I

suspected in specialized structures (Sokolow's organ - Hanstrrm, 1965)) on the ocular tubercle adjacent to

the eyes (Richter, 1982). •

The central nervous system consists of a bilobed cephalic "brain" of paired supraesophageal ganglia, II

circumesophageal commissures, and subesophageal ganglia. The optic nerve arises dorsally from the

supraesophageal ganglion, while the chelifore nerve arises anteriorly. The stomodeal nerve, dorsal and I

ventral proboscis nerves, and the palp nerve arise anteriorly on the subesophageal ganglion, while the oviger

nerve arises ventrally. A ventral chain of ganglia extends from the rear of the cephalon, through the trunk,

and into the abdomen. Except for the last three, all of which supply the abdomen, each ventral ganglion •

innervates a leg pair. II

The Digestive System I

The mouth, at the tip of the proboscis, forms the entry point of the digestive system. This empties

into the esophagus (foregut), then into the midgut (in the posterior portion of the cephalon, and throughout IBm

the trunk), and into the hindgut in the abdomen. Each division of the alimentary canal is separated by a l

tripartite valve. These one-way valves are held closed by presence of food on their posterior sides. Once

digestion and/or evacuation has reduced the volume of the material occupying the digestive lumen, muscular I
I

I activity in the proboscis can force more food through the valve. The system terminates in a slit-like anus at

the end of the abdomen. The foregut is lined with a continuation of the external cuticle, which is shed

I during mass, andmolting. adequate The digestive distributedsurface nature isofprovided the pycnobygonidlateral bodymidgut precludes caecaedevel extending opment of intoantheenlarged legs. Inmidgut some

Nymphonidae and Phoxichilidiidae the midgut is also anteriorly elaborated into the chelifore scape.

I to the esophagus. Ingested food The isresulting maceratfood ed bypulpmuscular is strained actionthrough withinathe grouppharynx, of setaewhich projecting extendsintofromthe the proboscis mouth

lumen at the beginning of the esophagus. These form a sieve (the "oyster basket") serving to retain larger

I particles within the pharynx for further maceration. Digestion is performed by epithelial cells of the midgut lining. Schlottke (1933) proposed a model

of pycnogonid intracellular digestion in which these cells become dislodged during digestion and wander

I freely through the lumen, later becoming secondarily reattacbed to the gut wall. Intracellular digestion proceeds within these cells, which retain unusable digestive products within themselves. They then detach

from the gut wall, and axe, along with their unusable inclusions, voided through the anus. Observations on

I digestion in several Antarctic species (Richards and Fry, 1978) suggest that this model is misleading, and that digestion normally proceeds within attached digestive cells, followed by release of digestive byproducts

into the gut lumen. Enzymes released into the lumen concurrently may also provide some extracellular

I digestion.At least in the antarctic species investigated digestion was a very slow process, with two months of

non-feeding to empty the guts. It is likely that higher ambient temperatures would raise the rate of digestion

I along with other aspects of metabolic activity. Such low rates of digestion also provided high resistance to starvation, with survival for over 18 months without apparent damage (Richards and Fry, 1978).

I The Circulatory System

The pycnogonid heart is an elongate sinus extending along the underside of the dorsal body wall

I from the dorsal the rearsideofofthethecephalon gut), andthrough its upper the thorax. is the inside Its lowerof the boundary body wall is the(Tjonneland horizontal septum et al., (1985). attachedTheto

horizontal septum divides the entire body into an upper and lower half and separates the haemocoel into

I uispper slit effere near thent and lateral lowerproacffesses, ernt cconsequently ompartments. separation The septumofdoe thestwo not reach flowstheis only ends of partial. the append It is sufficient, ages, and

however, to guarantee movement of hemolymph from the heart throughout the body. It appears that the

I heart of gutitself peristalsis provides and little muscular propulsion movement to circulate within the the legs. hemolymph, Up to threewhichpulsesis accomplished of hemolymphbymovement a combination have

been observed per second (Loman, 1907). The ventral return flow to the heart is via paired ostia opposite the

_ bases of the second and third legs, and an unpaired opening at the base of the abdomen (Cole, 1910).

| Pycnogonid respiratory pigment is apparently hemocyanin, which is present in the plasma but not

localized in blood cells. This may reflect the lack of respiratory organs in pycnogonids. It is assumed, partly

I because volume ratio, of thethat abseoxnycegenofisspecialized normally obgas tainexchange ed over thestruc entitreures, bodand y surface partlybybecdiffu ausesionof from their thhighe surrounding surface to

medium. Blood cells similar in appearance to those normally associated with oxygen transport are present

I (Dawson, 1934; Redmond and Swanson, 1968), but their function is unclear.

The Excretory System

I Specialized organs for excretion are lacking in pycnogonids. Removal of metabolic byproducts is

thus either across the general body surface, or into the gut for subsequent elimination through the anus.

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The Reproductive System I

Sexes are separate in all known pycnogonids with the exception of Ascorhynchus corderoi, a true

hermaphrodite species from Brazil (Marcus, 1952). Fertilization is external, and fertilized eggs are usually i

borne on the ovigers of the male.

The ovaries of the female lay along and above the gut, with lateral diverticulae extending into the i

legs in most species (Figure 1.3B). The testes of the male (Figure 1.3A) are similarly situated, and usually 1

extend into the legs. Ova are usually stored in the femurs of all legs between their production in the ovary

i

and their fertilization and release. In some species it is the second coxae that serve as the storage area for the

eggs. In either case, the segment becomes expanded to accommodate storage of the mature eggs prior to 1

their release. i

Eggs are released just prior to fertilization through a female genital pore on the second coxa of each am

walking leg. Typically these pores are antemventral, but are posterodorsal in Pycnogonum Iittorale. In the I

genus Rhynchothorax, they are found ventrally on the second coxae and only on the last pair of legs. The

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opening of the pore is closed by a lid or operculum, but not in Achelia echinata, where the pore is raised on

tubercle, rather than flush with the segment surface, i

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Early stages in development of pycnogonids from vitellogenesis of the egg to embryology is

extensively reviewed by King (1973). Postembryonic development is anamorphic, with the developing egg el

yielding a protonymphon larva (Figure 1.4). Subsequent growth is through addition of segments posteriorly. |

Very juvenile pycnogonids are two-legged (the first leg pair on the cephalon). Addition of the thoracic

segments yields four, six, and fmally eight legged individuals. More than one molt may be required for I

complete development of each leg pair, so that the last pair may consist of incomplete legs composed of just |

a few segments.

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Figure 1.3. Location and extent of gonads in; A. male (after El-Hawawi and King, 1978), and B. female I

Pycnogonum littorale (after Jarvis and King, 1972).

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Figure 1.4. Protonymphonlarva ofAchefia echinata (after King, 1973). !

i Ecology and Behavior

Reproductive Behavior

I The reproductive behavior of relatively few species of pycnogonids has been directly observed, and

the existing observations indicate that the process is far from uniform. Some variation is dictated by the

positions of the genital openings in variousspecies. Males and females typically engage in pseudocopulation,

I with their gonopores opposed. In Anoplodactylus lentus, where both the male and female gonopores are on

the ventral surfaces of the leg, pairing is ventrum to ventrum. In Pycnogonum littorale the male gonopores

open on the ventral side of the leg, while the female gonopores are dorsal. Consequently, this species pairs

I ventrum to dorsum, with the gonopores appressed. As eggs are extruded by the female they are fertilized and gathered into a ball by the ovigers of the male (King and E1-Hawawi, 1978). The ball is carried under

the male until hatching. The form of the egg mass differs in other members of the genus, being a flattened

I mass held laterally by the ovigers in P. panamum.

Nakamura and Sekiguchi (1980) made detailed observations of PropaUene longiceps reproductive

behavior in the laboratory. They found that oviposition was stimulated by male mate holding. The egg then

I migrated from the ovary to the female gonopore, was extruded, and was caught by the female with her

ovigers. It was then transferred to the male's ovigers where it was added to others to fashion "bracelets" of

eggs which were carried around the ovigers until hatching. The pseudoeopulatory position in this species

I was male ventrum to female dorsum, with both facing in the same direction. Pairing lasted several hours, with the male maintaining his position by grasping the base of the female's chelifores with his chelae. A

similar period of pairing and the same ventrum to dorsum position was reported for ParapalIene avida

I (Hooper, 1980).

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The duration of pairing was much shorter (minutes) in AnoplodactyIus lentus (Cole, 1901), and I

particularly extended (up to five weeks) in Pycnogonum littoraIe (Jarvis and King, 1972).

Seasonality of reproduction is characteristic of littoral species such asNymphon gracile while offshore i

species tend to have maturing eggs throughout the year. Species with both littoral and sublittoral populations,

such as Achelia echinata, have pulsed seasonal reproduction in the former and fairly continual reproduction

in the latter (King, 1973). I

Locomotion II

Pycnogonids are capable both of walking and swimming. The former was discussed at length by I

Schram and Hedgpeth (1978), who separated walking motion into two general categories related to leg

length: the crawling motion of short-legged species, and the walking of long-legged species. In

Swimming is used by pycnogonids to raise themselves off the bottom both for onshore/offshore II

tidal driven migration (Morgan, 1977) and current driven relocation by individuals in deeper water (Grassle

et al., 1975). Such activity may be relatively frequent as reports of positive phototropism and catches of II

pycnogonids in the plankton with night lights are numerous (see reports listed by Arnaud and Bamber,

1987).

The functional morphology of the musculature and leg articulations of pycnogonids with regard to i

both walking and swimming was discussed by Manton (1978). g

Feeding Behavior I

Most pycnogonids are carnivores feeding on sessile marine invertebrates, especially ectoprocts and

cnidarians. Algae and/or detritus apparently forms the majority of the diet in other species. Some Antarctic •

species have been found to be necrophagous and can be attracted to baited traps (Arnaud, 1972). !1

Careful observation or experimentation is necessary to separate recurrent association from predator/

prey relationship. Consistent association ofAchelia longipes with Flustellidra hispida suggested feeding by i

the pycnogonid on the ectoproct until Wyer and King (1973) observed the pycnogonid ignoring ectoproct

zooids, and consuming red algal sporelings growing epiphytically on the colony. Consumption by Achelia

echinata of both red and brown algae, but not bacteria or other epibionts on the algae, was demonstrated •

using radioactive tracers (Bamber and Davis, 1982). Where AcheIia echinata was associated with Bugula II

turbinata, the food source was organic detritus accumulated near the colony base, not the ectoproct (Ryland,

1976). I

In other cases, observation has demonstrated predator/prey relations in commonly associated

organisms. Members of the genus Pycnogonum have, for instance, been shown to prey on anemones with

which they are usually found (Gruet, 1972; Wyer and King, 1974). Preference experiments show P. littorale i

will also feed avidly on anemones not normally accessible to it (Bamber, 1985). g

The actual attack on the prey usually involves the proboscis and some combination of the chelifores I

and palpi if they are present. Ectoproct feeders may range over the face of an encrusting colony, avoiding J

defensive avicularia and orienting themselves for a forced entry past the operculum and into the zoarial

aperture, where the zooid is then consumed. The shape of the proboscis is usually modified in ectoproct

feeders with distinct preferences for particular prey species. In species of Austrodecus with particularly I

attenuate flexible proboscides, the apparent prey has the zoarial aperture restricted by internal structures

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(Fry, 1965). Movements of the lips and pumping by the muscular pharynx are involved in ingestion of prey.

Prey may be partially digested prior to ingestion by enzymes pushed forward from the midgut and released I

through the proboscis.

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I Cnidarian feeders tend to feed suctorially, but may also tear apart hydroid polyps with their chelae

and then carry the fragments to the mouth. Pycnogonum feeds suctorially on anemones, and Nymphon

I rubrum cheliforepumps (King, body 1973).fluids Pigrogromitus from the hydroid timsanusLaomedea suctorial feeding after first on the tearing anemone a holeBartholomea in the hydroid annulata with isa

tolerated by the anemone if only a single pycnogonid is involved. Attack by multiple individuals, however,

I led to death of the anemone within one month (Mercier and Hamel, 1994) Feeding habits of most California species can only be inferred from investigations on related species

from other areas. Bain (1991), however, provides some direct observations on the feeding of Anoplodactylus

i I californicus in captivity. Both living and dead animals were consumed. The soft-bodied opisthobranch

_ gastropod Berthella engeli was immediately attacked when the pycnogonids were first introduced into an

aquarium. First attack was with the proboscis unaided by the chelae, but chelae were seen carrying fragments

I of chelae the mollusk were usedpreyto the rapidly following snap swimming day. Capturebrineandshri consumption mp from theofwater brineandshripass mp was themalsoto theobserved. mouth. The The

pycnogonids would also accept pieces of mussel flesh offered to them. Pycnogonids have been observed

I feeding was termed on mussels a parasiticin situinfestation as well (Benson at the time, and but Chivers, can also1960), be interpreted causing extensive as predation. damage to the clams. This

At least some pycnogonids may be somewhat opportunistic, feeding on organisms not usually

I considered prey. Piel (1991) observed Anoplodactylus carvalhoi attack the aeolid nudibranch Dondice occidentalis, provoking it into defensive shedding of the finger-like processes (cerata) on its back. Ceratal

autotomy is a defensive strategy of many aeolids, but in this case it failed. The pycnogonids harvested cemta

I from the mollusk, one in each chela, then proceeded to suck out the ceratal tissue with the proboscis. This same species also consumed small polychaetes taken from the same hydroid mass.

I Symbioses

Both parasitic and commensal relationships between pycnogonids and other organisms have been

i reporte for asteroids, d. Commensal ophiuroids, associ andation echinoids. is usuallyStock betwee (1981) n pyfound cnogoninumerous ds andecPycnosomia hinoderms, and asterophila have beenclinging reportedto

the oral surface of the starfish Calliaster corynetes in the Philippines, andDecachela dogieli has been found

i attached to the ambulacral groove of Pteraster tesselatus in the Santa Barbara Channel (D. B. Cadien, unpublished). In both of these cases, and in the observation ofAnoplodactylus ophiurophilus on three ophiuroid

species by Sloan (1979), the pycnogonids may have been feeding on the mucus produced by the echinoderm.

i Associations between pycnogonids and echinoids have been reported for Pycnosomia strongylocentroti by

Losina-Losinsky (1933), and for Anoplodactylus insigniformis and Ascorhynchus castelloides by Stock

(1975a), in all cases as ectocommensals clinging to the urchin test or spines.

i Both Ectoparasitic relations with pycnogonids are reported for polychaetes, cnidarians, and mollusks. reports of ectoparasitism on polychaetes involve juvenile pycnogonids: Hannonia sp on Audouinia

australis (Stock, 1959), and Ammothella spinifera on Sabella melanosfigma (Salazar-Vallejo and Stock,

i 1987). Ectoparasitism on cnidarians also typically involves juvenile not adult pycnogonids. Russel (1990) reports juveniles of Tanystylum duospinum as ectoparasites of Eucopella everta, and Ammothea hilgendorfi

juveniles endoparasitic in the same hydroid in the Central California intertidal.

I cnidariansBoth(Hedgpeth adults and,1962; youngMauchline, of several1984), speciesandofcnidarian Pallenopsistissues are have associated been found with midw in theatergutpeoflagicthe

pycnogonid by Child and Harbison (1986). In California waters Anoropallene palpida have been taken

I from shallowthe they bell ooccasionally f a large undescribed contactedmedusa the bottom, (Martinthis and may Kuck,reflect 1991).onlySinceantheaccidental medusae phoresy were in water and not so

ectoparasitism.

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I 11
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Juvenile pycnogonids have been reported as either endo- or ectoparasites of a variety of bivalves, I

and both benthic and pelagic mollusks (summarized in Arnaud and Bamber, 1987). The only reports of

parasitism by adults are those of Benson and Chivers (1960 - here considered predation) and that of Arnaud I

(1978) who found an adult female Ascorhynchus endoparasiticus nearly filling the mantle cavity of the II

shelled opisthobranch Scaphander punctostriatus.

Predators l

Pycnogonids have been viewed as being incidentally consumed along with substrate (such as algae), II

or with other target invertebrates (Fry, 1980; Arnaud and Bamber, 1987). At least in one case, however, •

they are apparently actively sought by fish predators. Up to 30% (Zander and Heymer, 1992) and 40%

Ill

(Krapp and Nieder, 1993) of examined guts of several Mediterranean blennioid fish species contained II

pycnogonids. Blennioid fishes have also been reported to take pycnogonids in the northeast Atlantic (Gibson, II

1972), and in the Gulf of California (Kotrschal and Thomson, 1986). Further investigations may find similar

patterns of predation by some groups of specialized vertebrates or invertebrates. Hi

With their thick cuticle pycnogonids are probably not normal prey of most small invertebrates. I

Larger invertebrates, which are either strong enough to dismember pycnogonids or consume them whole,

require protection beyond that offered by the cuticle alone. Ecdysteroids used normally in control of growth, I

molting, and gametogenesis by Pycnogonum littorale are also released as a feeding deterrent in response to |

molestation by potential predators (Tomaschko, 1994b). They effectively deter predation by Carcinus maenas,

leading within seconds of release to the suspension of feeding by the crab (Tomaschko, 1994a). Chemical I

defenses have yet to be demonstrated for other pycnogonid species. I

Collection and Preservation I

Few pycnogonids are taken on soft substrates compared to the number that frequent rocks and their •

associated epibiota. In the deep-sea soft substrate forms are more common, as the lightweight pycnogonid II

body is well suited to the muds and clays of deep basins. As these forms tend towards large size, they can be

individually harvested from grab samples or trawls. In shallower areas soft bottom pycnogonids are usually

smaller, although some nymphonids reach relatively large sizes. Such forms are still not likely to be noted I

among the screenings of benthic grab samples and will probably only be found during processing of preserved

samples. I Samples collected remotely from offshore rock substrates, either by directly in scrapings, or indirectly

on collected rocks, are likely to yield at least some pycnogonids. Again, these cryptic organisms will probably

only be detected during laboratory processing of field collections. 1

Although the strong thigmotaxis of pycnogonids, combined with their gangly body form, makes

relaxation in isolation desirable prior to fixation, it is usually not possible. Relaxation of bulk samples with mlh

propylene phenoxytol or magnesium chloride prior to fixation may help prevent damage to pycnogonids 1

they contain. Most arthropods have less damage and leg loss in samples where relaxing agents have been HI

added prior to preservation. Collected pycnogonids do not require different fixation from other benthic

forms; fixation in 10%buffered seawater formalin being recommended. After a 24 to72-hour fixation period, 1

samples should be washed in freshwater until the formalin is removed, and preserved in either 70% isopropyl J

or ethyl alcohol.

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I Laboratory Methods

I Particular must be exercised in the removal of from debris and other organisms in pycnogonids care

bulk samples. Their eight legs are usually spread in such a way that as soon as one leg is freed from

entanglement, two more become caught. Application of force to extricate tangled specimens will usually

I result in lost legs, or loss of terminal claws. It is beneficial to place a pycnogonid, once f'mally extricated

from all entanglements, in a separate vial. Placing it in a vial with other arthropod specimens will only result

in re-entanglement, especially if more than one pycnogonid is involved. When live, pycnogonids are strongly

I thigmotactic, and will gather together in a ball of embracing arms. Somehow this tendency seems to persist

even in preserved specimens.

Once a single animal has been separated it should be examined with a dissecting microscope.

I Availability magnifications up to generally advisable, propodal spination

of 100x is because details of

usually require close examination. Adequate lighting is a necessity, with both fiber optic illumination, and

dark field substage lighting advantageous. Where possible, the animals should be examined entire, with

I disarticulation of appendages undertaken only if absolutely It be to gently brush

necessary. may necessary

debris from the specimen with an extra free sable or camel hair artists brush during the examination.

i A few very small species, such as Rhynchothorax philopsammum can be usefully examined in whole mount under a compound microscope. The animal should be protected from crushing by the coverstip

either by use of a depression slide, or through use of wedges (coverslip fragments, bits of clay or sand, etc.)

i to provide additional space between slide and coverstip. Fine detail of legs of larger species (such as femoral cement gland shape, or propodal armament) can also be examined in this manner. Specimens should be

temporarily mounted in glycerol for examination, then removed and returned to preservative for maintenance.

Rather than preparation of permanent slides for small specimens or body parts, these should be kept in small

I vials associated with the main sample. Rubber or silicone stoppered glass insect genitalia vials provide safe

storage, and are recommended for retention of small disarticulated parts.

Since all characters of taxonomic importance in pycnogonids are external, cleating and staining are

I not necessary for species determination.

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Glossary I

I

Abdomen. Last body segment. Articulated or Femoral cement gland(s). Glands located in the •

fused to the posterior end of the last thoracic femur of the males of many species which

Jl$

segment. Often bent dorsally if elongate, secrete a sticky mucus used to form egg m

and usually continuing any dorsal balls carried by the ovigers, or to stick eggs 1

ornamentation of the thorax. Bears the anus to other substrates. Usually a single exit w

at its terminus, pore which may be raised on a tubercle, or

prolonged into a tube. In Colossendeidae I

Auxiliary claw(s). Secondary claws (one or two) usually numerous simple pores flush with

which may be present behind the primary the segment surface.

claw; they may be larger, but are usually •

smaller than the primary claw. Femur(ae). Fourth segments of the walking legs. II

Attached proximally to the ends of the third

Cephalic hood. An anterior prolongation of the

ocular tubercle over the attachment points of cases,C°xae malsand distallYbear onet°heomr orefirst tibia.cemensotmein I

the proboscis, chelae, and palpi in some glands in the femur. Used by females of

species, many species for storage of eggs prior to

Chela(ae). Paired pincer-like structures flanking their release and inflated as a result.

the proboscis and borne on a one or two- Genital pore(s) See Gonopores.

segmented scape. A chela has a fixed basal •

segment and a moveable terminal segment Gonopores. Openings of the male or female

(finger). In some families they are reproductive system, usually located on the

degenerate in the adult; reduced to knobs at second coxae of one or more pairs of legs. •

the end of the scape. Heel. Tarsus of the walking legs, often bearing !i

Chelifore(s). The first pair of cephalic large spines.

Ii

appendages, which bear the chelae at their

Lateral processes. Lateral extensions of the body I

ends. Formed of a basal scape of one or two

(either cephalon or thorax) to which the first

segments, and the chelae (unless lost or

reduced). Attached on the dorsal side of the coxae of the walking legs are attached.

proboscis just behind its insertion on the Ocular tubercle. Dorsal hump or tubercle on the II

cephalon, cephalon which bears the eyes. Typically

lit

Coxa(e). The first three segments of the walking near the middle of the cephalon, I

legs (first, second, and third coxae). The exceptionally near its anterior or posterior

first coxa is attached to the lateral process of edges. May be extremely tall to nearly absent. 1

the body, and the third is attached to the I

proximal end of the femur. Usually short Oviger(s). Third cephalic appendages (although

relative to the femur and tibiae. In the one or both of the first pairs may be absent)

female the gonopore is typically located which originate ventrally just in front of the i

anteroventrally on the second coxae of the first leg pair. May be used for egg carrying

walking legs. in male, and as grooming appendages in B

Crufiger(s). See Lateral processes, both sexes. 1

Denticulate spines. Spines which bear lateral Palp(s). Second cephalic appendages (first pair

denticles. Often located on the strigillus, or may be absent) which originate lateral to the I

on the chelae, but occasionally on the proboscis near its base.

walking legs. I
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I Primary claw(s). Moveable sickle shaped Tarsus. The seventh segment of the walking legs.

structure(s) which form the last segment of Attached proximally to the second tibia and

I the auxiliary wlkingclaws legs.near Usually the attachment with one oroftwo the distally segmenttowiththe apropodus. short anterior Usuallyand along small

primary to the pmpodus. Exceptionally posterior margin; seldom ornamented with

I reduced or absent, tubercles or spines. Proboscis (ides). Muscular bulbous, tubular, or Terminal claw(s). See Primary claw(s).

cone-shaped organ bearing the mouth at its

I end and attached to the cephalon along the Tibia(et)ibi . aThe ) segments fifth (firofst tithebiawalking ) and sixlegs. th (second Often midline of the body. Usually points

anteriorly, but is bent down or even under elongate in long-legged species, and often

I the body in some species, bearing tibia is atsetae, tachedtubercles proximallyor spines. to the femur, The firstthe

Propodus. The eighth (penultimate) segment of second tibia distally to the tarsus.

i the walking leg. Attached proximally to the Tubercle(s). a rounded to sharply pointed

tarsus, and distally to the primary claw.

outgrowth of the cuticle, sometimes bearing Usually curved and bearing spines and/or a

blade of fused spines on its concave surface, setae or spines at its apex. Tubercles can

I occur on all portions of the body and Strigillus(ae). The modified four terminal walking legs, with the exception of the

segments of the ovigers. Curved inward into proboscis.

I a near circle, and bearing modified spines on the concave side. Used in grooming of

legs.

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I 15
I

Key to the Pychnogonids of California I

The present key is based on those previously presented by Child, 1979; Hedgpeth, 1941b; and Hedgpeth, I

1975. Species reported or known to occur from intertidal to upper slope depths in California are included.

Those marked with an asterisk (*) were not taken in the present collection, and are not described in the text. 1

They are included to increase the utility of the key, which otherwise would include only one-quarter of the 1

species one might encounter in California waters. Species from basin and lower slope depths are not treated

i

here (see Child, 1994, for information on this deep-water fauna). II

HI

1A. Chelifores and palps present; ovigers in both sexes ........................................................................ 2

lB. Chelifores and/or palps absent or greatly reduced; ovigers in both sexes or male only ................. 4 I

2A. Chelifores and/or chelae vestigial in adult; palps 4 to 10-segmented - Family Ammotheidae ...... 8 I

2B. Chelifores and chelae well developed in adult; palps 4 to 5-segmented -Families Nymphonidae and I

Callipallenidae (in part) .................................................................................................................. 3

I

3A. Palps 5-segmented, ovigers 10-segmented - Family Nymphonidae ............................................. 29

3B. Palps 4-segmented, ovigers 9-segmented .................................................... AnoropaIlene palpida* I

4A. Palps present - Families Colossendeidae and Rhynchothoracidae ................................................. 5 1

4B. Palps absent ..................................................................................................................................... 6 !

5A. Proboscis longer than body, fusiform; ocular tubercle tall, eyes well developed; auxiliary claws 1

lacking on legs ................................................... Hedgpethia californica* (Family Colossendeidae) I

5B. Proboscis shorter than body, widest near base and tapering distally; ocular tubercle a low anoculate

boss; short auxiliary claws present on legs ..................................... Rhynchothorax philopsammum I

6A. Chelifores absent - Family Pycnogonidae ..................................................................................... 31 I

II 6B. Chelifores present - Families CaUipallenidae (in part) and Phoxichilidiidae ................................. 7

7A. Ovigers in both sexes, 1O-segmented - Family Callipallenidae (in part) ...................................... 32 I

7B. Ovigers in male only, 5 to 6-segmented - Family Phoxichilidiidae .............................................. 35

1

8A. Body circular or discoidal, lateral processes touching or only narrowly separated distally; chelifore I

scape 1-segmented; palps 4 to 8-segmented ................................................................................... 9

8B. Body more slender, lateral processes separated for at least half their length or widely separated; I1

chelifore scape 1 or 2-segmented; palps 9 to 10-segmented ......................................................... 10

Illl[

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I 9A. Palps 8-segmented; chelae vestigial; proboscis pyriform; 1st and 2nd coxae with tall dorsolateral

tubercles .......................................................................................................................... Achelia* 14

I 9B. Palps cylinder4 toor7tubular; -segmented; 1st coxae chelaeonlyusual withly absent, low rounded scape tubercles reduced t.............................. o short knob; proboTanystylum scis a tapering * 20

I 10A. Proboscis 2-segmented, a proximal cylinder articulated with a distal pyriform segment ................

............................................................................................................................... Eurycyde spinosa

I 10B. Proboscis l-segmented, pyriform, ovoid, or cylindrical ............................................................... 11

11A. Palps 9 or 10-segmented; chelifores inserted at anterior margin of cephalon .............................. 12

I 11B. Palps 8-segmented; chelifores and palps inserted underneath anterior margin cephalon ............

of

........................................................................................................................ Prototrygaeus jordani

I 12A. Proboscis ovoid or modified cylindrical; palps 9-segmented ....................................................... 13

12B. Proboscis pyriform; palps 9 or 10-segmented ......................................................... AmmothelIa* 24 !

13A. Proboscis ovoid; trunk with spiny median tubercles; legs with tall setose tubercles; chelifores 3-

I segmented; abdomen long, bent posteriorly ........................................... Nymphopsis spinosissima*

13B. Proboscis cylindrical, ends constricted; lacking median tubercles; chelifores blunt, 1-segmented;

abdomen short, blunt ................................................................................... Ammothea hilgendorfi*

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14A. Chelae lacking in adult .................................................................................................................. 15

I 14B. Chelate in adult ....................................................................................................... Achelia chelata*

i5A. Trunk and legs conspicuously spinose, with small spinose tubercles on 1st coxae ...................... 16

I 15B. Trunk and legs at most slightly spinose; 1st coxae lack spinose tubercles, but may bear finger-like

spurs ............................................................................................................................................... 17

I . . ,

16A. Terminal segments of palps with ventral lobes ................................................... Acheha spmoseta

16B. Terminal palp segments subcylindrical, lacking ventral lobes .............................. Achelia echinata !

17A. First coxae with fingerlike dorsal processes; propodus with basal spines; palps 8-segmented .... 18

I 17B. First coxae lack fmgerlike processes; no large basal spines on propodus; palps 7-segmented .... 19

i 18A. Processes on 1st coxae three-quarters coxa length; proboscis narrowly elliptical ...........................

'_ ............................................................................................................................ Achelia gracilipes*

18B. Processes on 1st coxae less than half coxa length; proboscis broadly elliptical AcheIia alaskensis*

I 19A. Chelifore with conspicuous dorsal tubercles ........................................................ Achelia latifrons*

i 19B. Chelifore smooth, without dorsal tubercles .................................................... Achelia simplissima*
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20A. Proboscis tapered to a point; body not particularly compact ........................................................ 21

20B. Proboscis rounded; body very compact ................................................... Tanystylum occidentalis* I

mill

21A. Without spines on body or lateral processes; proboscis bluntly pointed .......... Tanystylum nudum* l

2lB. Lateral processes with spines or tubercles; proboscis acutely pointed ......................................... 22

22A. Fourth palp segment with medial proximal tubercle .............................. Tanystylum californicum* I

22B. Fourth palp segment lacking proximal tubercle ............................................................................ 23

I1

23A. Lateral processes beating sharp spine-like tubercles; auxiliary claws nearly half length of primary I

claw .......................................................................................................... Tanystylum intermedium* m

23B. Lateral processes without spine-like tubercles; auxiliary claws only one-fourth length of primary I

claw ............................................................................................................ Tanystylum duospinum*

24A. Legs with both primary and auxiliary claws well developed ........................................................ 25 I

24B. Primary claw absent or vestigial ........................................................... Ammothella biunguiculata*

I

25A. Second segment of cheliforc scape clavate ................................................................................... 26

25B. Second segment of chelifore scape linear ..................................................................................... 28 I

26A. Ocular tubercule height several times diameter; lateral processes separated by one-fourth their width •

or more .......................................................................................................................................... 27 il

26B. Ocular tubercle about as wide as tall; lateral processes touching or nearly touching .......................

.................................................................................................................. Ammothella tuberculata* I

I

27A. Single tubercles mid-dorsally ........................................................................ Ammothella menziesi*

I

27B. Paired hollow tubes mid-dorsally .................................................................. Ammothella spinifera* •

28A. Lateral processes each with single smooth projection; abdomen lacking spines or tubercles .......... II

........................................................................................................................... Ammothella setosa* II

28B. Lateral processes each with multiple spined tubercles; abdomen with spined tubercles ................. m

AmmothelIa kilIix* •

I!

29A. Ocular tubercle located about midlength on cephalon, preceeding insertion of ovigers, and well 1

before first leg pair ............................................................................................ Nymphon stipulum*

29B. Ocular tubercle located above or slightly behind insertion of ovigers, and next to first leg pair. 30

I

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I 30A. Tarsus slightly longer than propod; terminal oviger spine nearly equal to length of terminal segment

............................................................................................................ Nymphon heterodenticulaturn

I 30B. Tarsus more than twice propod length; terminal oviger spine half length of terminal segment or less

.............................................................................................................................. Nymphon pixelIae

I 3IA. Body unicolor; dorsal tubercles short, about as tall as basal diameter ......... Pycnogonum stearnsi*

3lB. Body reticulate; dorsal tubercles tall, height greater than basal diameter ..... Pycnogonum rickettsi

I 32A. Chelae with serrate or denticulate fingers; auxiliary claws present on legs ............ Callipallene 33

32B. Chelae with smooth f'mgers; auxiliary claws lacking ............................................... Decachela* 34 !

33A. Segmentation of thorax incomplete ...................................................... CallipaUene californiensis*

I 33B. Segmentation of thorax complete CaUipaUene pacifica

I 34A. 34B. Thorax Thorax and and legs legs not heavily conspicuously spinose ........................................................................ spinose ....................................................... Decachela Decachela discata dogieli**

I 35A. Cephalon projecting over proboscis forming long neck (except in A. viridintestinalis); oviger 6-

segmented, with terminal segment very small ........................................... Anoplodactylus ........ 36

I 35B. Cephalon or completely not elongated fused to 5thanteri segment orly int........................................................... o neck; oviger 6-segmented, with terminal Phoxichilidium segment* partia ...... lly41

I 36A. Cephalon only slightly overhanging proboscis ............................ Anoplodactylus viridintestinalis*

36B. Cephalon extended forward over proboscis forming long neck ................................................... 37

I 37A. Lateral processes separated at base ............................................................................................... 38

37B. Lateral processes contiguous at base, body very compact ................. Anoplodactylus californicus*

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38A. Lateral processes and first coxae each with a pair of tubercules ......... Anoplodactylus oculospinus

I 38B. Lateral processes and fast coxae each with one tubercle or none ................................................ 39

39A. Tibiae and femur of legs with prominent nodulation ............................... Anoplodactylus nodosus*

I 39B. Tibiae and femur of legs not conspicuously nodose ..................................................................... 40

I 40A. Lateral processes with a single dorsal tubercle ............................................ Anoplodactylus erectus

40B. Lateral processes lacking dorsal tubercles ................................................ Anoplodactylus pacificus

I 41A. With small knobs (vestigial palps) just posterior to chelifores .................. Phoxichilidium parvum*

4lB. Without vestigial palp knobs posterior to chelifores ............................. Phoxichilidumfemoratum*

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I

Description of Species I

Family Ammotheidae I

Achelia echinata Hodge 1864 i

Figure 1.5

Achelia echinata Hodge, 1864:115; pl. XII, figs. 7-10.--Hilton, 1943a:94-95.--Utinomi, 1954:11-14, figs. I

4-5;1959:201-202, fig. 1--King, 1974:28, fig. 11.--Lissner et al., 1986:D-26.

Ammothea echinata Bouvier, 1923:55-57, fig. 55. i

Material Examined. California, off Morro Bay, Sta. 6 (1).

Description. Thorax compact, circular; lateral processes short, subequal in length, little separated, 1

bearing 2 dorsal tubercles near ends. Thoracic segmentation well defined to nearly absent. Abdomen about Ii

equal in length to cephalon, angled upward, slightly larger in diameter terminally.

Proboscis longer than cephalon, about equal to thorax in length; conical, widest at about two-thirds 1

of length. m

Neck nonexistent. Ocular tubercle low, height about twice basal diameter; 2 pairs of eyes borne

near top. A pair of laterally directed tubercles at base of ocular tubercle, larger and more attenuate in male. !

Chelate only in juvenile, although scape retained in adult with chela reduced to small terminal

knob. Chelifore extending one-half to two-thirds length of proboscis. Scape bearing variable number of am

spines and tubercles dorsally. Palpi 8-segmented, with f'mal 4 segments setose; segment 2 longest, segment i

4 somewhat shorter, with remaining segments much shorter and subequal.

Ovigers 9-segmented in both male and female, but slightly smaller in female; both sexes with well- •

developed strigiUus bearing digitate spines. Few setae except on strigillus. Segments 1 and 4 longest; remainder U

somewhat shorter and subequal.

Legs not particularly long (about 3 times width of thorax), length about 10 times diameter, spiny, •

spine pattern variable. Coxa 1 about 2 times length of other coxae; femur and tibia 2 subequal, about twice

length of coxa 2. Tibia 1 intermediate in length between coxa and femur. Tarsus about one-fifth tibia 2

length; propod about two-thirds length. Terminal claw about one-half propod length; auxiliaries between 1

one-half and two-thirds length of primary, longer in male. |

Coxa 1 bearing pair of tubercles in male, only one in female; male genital pore situated at end of a

large genital spur on second coxae of third and fourth legs. Female pore not elevated; second coxa broadened 1

distally. Femoral cement gland of male not prominent. Tarsus armed with 3 or 4 spines ventrally. Heel of II

propodus weak, scarcely differentiated from rest of segment, bearing 3 to 4 large spines; about 10 sole

spines; no lamina.

Biology. Achelia echinata is a widely distributed species associated with sponges, hydroids,

ectoprocts, and algae (Peatde and Hoare 1981). The species feeds on hydroids (Wyer, 1972), ectoprncts

(Wyer and King, 1973), algae (Bamber and Davis, 1985), and detritus (Ryland, 1976). The unspecialized 1

diet and association with fouling organisms, may account for its wide distribution in the northem hemisphere. i

Remarks. The examined specimen is not typical for the species. The cephalic tubercles are more

slender and acute than usual, the chelifore scapes bear dorsal tubercles, and the lateral margins of the coxae i

are very setose. The combination of these characters is reminiscent ofAchelia salebrosa (Losina-Losinsky,

m

t961). Widely-distributed species should be more polymorphic than those with narrowly constrained niches !

20 I
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I

I ._.._il c

I, _ \_ ! A

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I Figure 1.5. Achelia echinata; A. body, dorsal, B. terminal claw, C. female oviger, D.juvenile chela, and E. adult chela (all after Utirtomi, 1954).

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and distributions. Such variation has been documented for A. dohrni by Clark (1971), and the A. echinata I

illustrations of Utinomi (1954) show an animal differing in some respects (especially proboscis shape) from

North Atlantic material (King, 1974). Alternatively, an unresolved species complex may exist. In England, I

A. echinata has been reliably differentiated from several similar species. North Pacific material has yet to be

critically examined to determine the limits of variation in this and related species. Additional specimens

from the northeast Pacific with the same or similar variations from typical A. echinata should prompt re- •

evaluation of this specimen, which may represent a new species. m

Type Locality and Type Specimens. Not designated by Hodge (1864).

Distribution. North Atlantic; Mediterranean; Alaska, Hawaii; California, San Francisco to San 1

Diego; intertidal- 109m. I1

Eurycyde spinosa Hilton 1916a I

Figure 1.6

I

Eurycyde spinosa Hilton, 1916:19-24, figs. 1-6;1939:72, fig. 11;1943a:98.--Hedgpeth, 1941b:254, pl. 9.--

Lissner et al., 1986:D-26. 1

Material Examined. California: off Morro Bay, Sta. 27 A/B (1); Santa Mafia Basin, off Purisima 1

Pt. Sta. 20 A/B (3).

Description. Thorax compact, nearly circular; segmentation complete, distinct; lateral processes of •

moderate length, separated by less than their diameter, divergent, and broadening distally. Abdomen longer II

than cephalon, slightly longer than lateral processes of fourth legs; borne vertically over last thoracic segment.

Swollen at about 80% of length, with abdomen diameter increased bynearly half. A corona of 4 long spines •

(length 4 times abdomen diameter) directed anterolaterally on the swelling. Beyond abdomen tapering II

rapidly to rounded tip.

Proboscis with 2 joints, with bulbous terminal segment connected to narrow cylindrical pedicel, as i

typical of genus. Pedicel as long as cephalon; terminal proboscis segment haft again as long, its greatest II

diameter about one-third segment length from pedicel, tapering evenly from that point to bluntly rounded

tip. Second proboscis segment bent downward and to rear under body. i

Neck very short and broad, half length of rest of cephalon. Ocular tubercle at base of neck, just

behind insertion of palps; height two and one-half times basal diameter, with 2 equal pairs of eyes just

below midpoint. Ocular tubercle topped by 3 dorsally directed spines shorter than those on abdomen. 1

I

Chelifore scape 2-segmented, with chelae nonfunctional in adult; fingers lost and palm reduced to

stub bearing 2 terminal spines. Scape basal segment smooth, without tubercles or setae; segment 2 with

series of sharply pointed tubercles, 3 on segment dorsal midline, 1 ventrolateral, and 1 at distal edge of •

segment at base of chela stub. Palp 9-segmented, slender, about double length of pedicel, but shorter than u

combined proboscis length. Basal segment, and segments 3, 5, 6, and 8 subequal. Segment 2 about 5 times,

segment 4 three times their length. Segments 7 and 9 slightly shorter than segment 8. Last 5 segments setose •

dorsally. III

Oviger 10-segmented, about as long as thorax. Segments 1-3 short and subequal; segments 4 and 5 u

subequal and equal to combined length of first 3 segments. Segments 6 through 10 successively shorter; i

segment 10 bearing pair of large simple spines at its end. Segments 7-10 beating a double row of denticulate

wll

spines, i

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I Figure 1.6. E.Eurycydespin°Sa and F.;oterminalA" viger,bodY,claw.dorsal (after Hedgpeth, 1941b), B. chelifore, C. proboscis, D. palp,

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Legs slender, length 25 times diameter. Coxa 2 twice length of coxae 1 and 3, and half femur length. I

Femur four-fifths length of tibia 1, and three-quarters length of tibia 2; tarsus equal to coxa 1 length, propod

3 times tarsus length. Propod lacking heel and lamina, with about 9 sole spines, and 3 longer dorsal spines. In

Terminal claw strongly tapering, about half propod length; no auxiliary claws II

Biology. The broad bathymetric occurrence of this species may indicate a lack of habitat specificity.

The present specimens were taken from small rocks that were retrieved with a submarine. The species has l

also been collected among Zostera roots (Hilton, 1939). No direct observations of feeding in Eurycyde II

spinosa have been reported. Absence of functional chelae in the adult, the terminally narrowed bulbous

proboscis born on a basal pedicel, andthe well developed palps suggest this is an ectoproct feeder. Collection •

among eel-grass roots suggests that detritus may also serve as a food source.

Type Locality and Type Specimens. Types were collected intertidally at Laguna Beach, California,

and deposited at Pomona College, California. I

Ig

Distribution. California: Morro Bay to Laguna Beach; intertidal-117m.

Prototrygaeusjordani Child 1990a I

Figure 1.7

I

PrototrygaeusjordaniChild, 1990a:157-160, fig. 1. •

Ammotheidae Genus Asp. A Lissner et al., 1986:D-26. 1

B

Material Examined, California: eastern Santa Barbara Channel, between Santa Cruz and Anacapa

Islands, BLM Sta. 24848 (1).

Description. Thorax slender, longer than wide; lateral processes slender, each as long as thorax, II

approximated basally, but diverging over length, with large tapering tubercle distally; thoracic segmentation

incomplete. Abdomen about as long as cephalon, held nearly vertically, slightly broadened distally, with a 1

few distal setae. U

Proboscis nearly twice length of cephalon; inserted ventrally with insertion overhung by anterior

edge of cephalon; sharply angled down, cylindrical, blunt, slightly narrowed distally. Neck very short, IJ

broad, not differentiated from rest of cephalon. Ocular tubercle shorter than abdomen, about twice basal

diameter in height, with 2 pairs of eyes near top, anterior pair slightly distal to posterior pair.

Scape of chelifore 2-segmented, inserted beneath overhanging front of cephalon, and extending to 1

end of proboscis. Chela nonfunctional in adult, with fingers lost and palm reduced to nub at end of scape. U

Basal scape article about half length of clavate second article. Second article with 2 distal setae; chela with

1; chelifore otherwise bare of setae. 1

I

Palp with 7 articles, fourth longest, article 2 slightly shorter, and remaining 5 one-quarter the length

of article 2. Only last 4 segments setose. Palps, like proboscis and chelifores, inserted underneath overhanging I

anterior cephalon margin. 1

Ovigers lO-segmented inboth sexes, with that of male considerably larger. Male oviger segments 4

and5 each shorter than segment preceding them, segments 2 and 3 subequal and longer than others. Basal I

segment about half length of segment 4. Segments 6-10 decreasing in length; segment 10 very short and il

bearing 2 curved tridentate spines, central tooth longest. Segment 9 lacks spines or setae, segments 6-8 bear

setae longer than segment diameter. Female oviger with fourth and fifth segments subequal and longest, m

articles 1-3 subequal and shorter than 4 and 5; strigillus with few setae, articles 8 and 9 each with pair of •m

non-digitate spines, article 10 with 1 pair.

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Figure 1.7. Prototrygaeusjordani; A. body, dorsal, B. terminal claw, C. palp, and D. oviger (after Child,

i 1990).

Legs of moderate length (about 3 times thorax width), appearing long because of elongate lateral

I processes. First coxa about two-thirds length of coxae 2 and 3. Female genital pores ventrodistal on coxa 2

of all legs. Femur and tibiae subequal, each about twice length of last 2 coxae. Tarsus short, less than onequarter length of propodus. Terminal claw about half length of propod, which is two-thirds length of long

i segments. Articles increasingly setose distally; first coxa bearing only 1 long seta. Femoral cement gland

distal, with short tapering duct. Terminal claw with 3 basal denticles on concave face, distally increasing in

size. Auxiliary claws nearly length of primary; slender. Propod heel poorly marked, with 3 large spines; sole

I with 12-13 anteriorly curved spines.

i 25
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Biology. The species was found to co-occur with Anoplodactylus erectus in Monterey Bay, and U

may have similar habitat requirements. The species has only been taken with remote gear (epibenthic sled,

grab sampler), and has not been observed in aquaria. Hydroids are numerous on the soft bottom from which •

the types were collected, and may represent its food.

Remarks. The single specimen examined came from a sample collected off the north side of

Anacapa Island during the BLM study of 1976-79. It was not taken either further to the west in the Santa n

Barbara Channel, or in the Santa Mafia Basin, although many samples were collected from it's reported II

bathymetric range. The species has been seen further south in the Southern California Bight, but no definite

•southern range limit has been established. This is the second species in the genus, which was established by me

Stock (1975a) for the similar Caribbean Prototrygaeus ammothelloides.

Type Locality and Type Specimens. Holotype male (USNM 234509); paratypes 1 female, 1

juvenile (USNM 234510), on soft sediments with hydroids, mollusks, and wood debris, Monterey Bay, n

California. I

Distribution. California: Monterey Bay to the eastern Santa Barbara Channel; 50-70m. I

Family Rhynchothoracidae I

Rhynchotlmrax philopsammum l-Iedgpeth 1951

Figure 1.8 U

Rhynchothorax philopsammum Hedgpeth, 1951:111 - 115, pl. 3;1975: 418; 421; 424; pl. 101, fig. 15.--

Krapp, 1973:123.---Clark, 1976:295.---Child, 1979:72.--Lissner etal., 1986:D-26. n

U

Material Examined. California: Santa Maria Basin, off Purisima Pt., Sta. 16 A/B (6).

Description. Thorax compact, elongate, segmentation well defined, with short lateral processes n

separated by less than their diameter and diverging distally. Integument with a granular texture caused by J

presence of numerous tiny pearl-like rounded processes. Abdomen horizontal, bluntly tubular, short, only

about half cephalon length. Cephalon and thorax subequal in length, m

Proboscis about as long as cephalon, widest just above base, and tapering to broad conical tip. Neck

broad, short, with pair of processes on anterior lateral corners beating palps. Ocular tubercle and eyes

absent, but in large specimens a low anoculate boss located dorsally near anterior margin of cephalon n

between 2 stubby anteriorly directed tubercles. Two smaller acute tubercles located behind and slightly I

lateral to boss.

Chelifores absent in adult, but may be present in juveniles. Palpi 4-segmented; segments 1 and 3 I

longest, subequal; extending to end of proboscis. Attached basally to lateral projection from cephalon with

i

dorsal tubercle at midlength; segment 3 with setose tubercle dorsally near end. Terminal segment distally

rounded, expanded dorsally, ending in setal brush. I

Ovigers similar in both sexes, with 10 segments and terminal claw; lacking setae. Segments 4-6

somewhat longer than others; first six segments smooth, segments 7, 8, and 9 with 2 distally directed spines; •

segment 10 terminating in large claw, fitting into pocket at end of segment. |

Leg length about 3 times width of thorax, with sparse short setae, and single long spine near ends of

femur and tibiae. Coxae 1 and 3 subequal in length, and half as long as coxa 2. Femur and tibiae subequal •

and about half again as long as coxa 2. Tarsus short, about one-fifth the length of propodus or coxa 2. II

Propodus only slightly curved, covered with short setae, with about 7 sole spines. Heel not defined, without

enlarged spines at proximal end of propod. Terminal claw robust, terminally hooked; auxiliaries about half n

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I Figure 1.8. Rhynchotho Hedgpeth, raxphilopsammum; A. body, dorsal, B. palp, C. oviger, D. terminal claw (after 1951).

I

length of primary and less curved. Lateral process of first and second legs with pair of tubercles postemdistally;

I legs occur.3 andFemale 4 withwithan anterior genital pore tubercle projecting on lateralposterodorsally process; in largeas males a spur aonsmaller coxa posterior 2 of fourthtubercle leg; male may with also

corresponding spur posteriorly on coxa 2 of third leg.

I sands, andBioislogconsidered y. This small to be species one of thewasthreeoriginally true interdescribed stitial formsfromin the undergenus 5-6(Clark, inches 1976). of coarse The intertidal attenuate

shape, lack of eyes, and small size are all specializations for this habitat. It is assumed that the species was

I living reported, in sand and pockets the first ontimetheitrocks has beercovered n reportedoffoffshore. Purisima InPt.addition This is only to thethetypethirdcollection, time the species specimens has been were

taken in the Gulf of Tehuantepec, southern Mexico (Child, 1979). These were shore and shallow water

I collections lots of this from speciesintertidal from southern rocks andCalifornia coral rubblein (Ftheraser, collections 1943). Child of the(1979) National mentioned Museumfurther of Natural unaccessioned History,

but provided no further details.

I Remarks. Several points in the original description were misleading or inaccurate. Child (1979) reexamined the type, and clarified the structure of the palp. Hedgpeth originally described it as five-segmented.

Child found the basal "segment" not to be separated from the cephalon by a suture line, concluding it was

i actually Hedgpetha described process toitwhich as 9-segmented the palp wasin theattached. accompanying Simlarly,text. whileAs illustrating pointed out aby10-segmented Child (op. cit.), oviger, the

supposed chelifores mentioned by Hedgpeth are really tubercles at the anterior rim of the cephalon. In a

I particularly It is unlikely large these female were overlooked from the present by Child, collections and must abesecon assumed d set toofbesmall of variable cephalicoccurrence, tubercles was or restricted located.

to the largest individuals.

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Type Locality and Type Specimens. Holotype female (USNM 91245), 2paratype females (USNM), I

1 paratype female (Pacific Marine Station, Dillon Beach); Tomales Bluff, opposite Dillon Beach, Tomales

Bay, CaliDifornia. I

stribution. Tomales Bay, California to Tangola Tangola Bay, Mexico; intertidal-123m.

Family Callipallenidae I

CaUipallene pacifica (Hedgpeth 1939) 1

II

Figure 1.9

PalIene pacifica Hedgpeth, 1939:460-461, figs. f-l.--Hilton, 1939:72, fig. 4. I

Callipallenepacifica Hedgpeth, 1941b:257, pL 1l.----_lton, 1942c:38.

I

Material Examined. California: Santa Maria Basin, off Purisima Pt., Sta. R5 (1). !

Description. Thorax slender, elongate; segmentation complete, distinct; lateral processes of moderate

length, with first and fourth pairs shorter than others; separated by about half their diameter. Abdomen 1

erect, shorter than lateral processes of fourth leg, but equal in height to ocular tubercle; slightly bilobed

posteriorly.

Proboscis short, bluntly rounded, as wide as long. Neck long, nearly 4 times length of cephalon l

posterior to it; narrowed at midlength then broadened distally for insertion of chelae; equal in width to

lateral processes of first leg. Ocular tubercle at base of neck, just above insertion of oviger; only slightly

taller than basal diameter; bearing 2 equal pairs of large eyes; bluntly pointed dorsally. 1

II

Chelifore scape 1-segmented, reaching slightly beyond end of proboscis; chelae robust, longer than

scape; fingers longer than setose palm. Palpi lacking. Oviger twice thorax length, 10-segmented. Segments It

1, 2 and 10 subequal; segments 3 and 4 slightly longer; segments 5, 8, and 9 half again as long as basal •

segment; segment 7 twice and segment 6 more than twice length of basal segment. Last 4 oviger segments

m

armed ventrally with leaf shaped spines, forming a strigillus.

Legs slender, length 23 times diameter, with a few scattered short setae. Coxa 2 twice length of I

coxa 1 and 3, and half femur length. Tibia 1 slightly shorter than femur, second tibia 25% longer. Tarsus

Ill

extremely short, only one-eighth length of coxa 1, propodus subequal to coxa 1 length. Propodal heel i

undeveloped, marked only by presence of 5 large spines; sole lacking lamina, and bearing 15-16 small 1

spines. Terminal claw two-thirds propod length; auxiliary claws three-quarters length of primary.

Biology. The single specimen examined was taken on soft sediments. The type specimens may • 

have come from offshore rocks. Additional specimens were reported by Hilton (1939) and Hewatt (1946), 1

from collections in shallow water at Santa Cruz Island. The area sampled has both kelp beds and a mixed

rock and soft-sediment bottom, so it is not possible to determine the exact substrate from which CallipalIene •

pacifica was taken. |

Remarks. The specimen taken in the Santa Maria Basin was quite near the type locality of this

species, but from somewhat deeper water. The other member of CallipalIene in California waters, C. •

californiensis, can easily be separated from C. pacifica by a much shorter neck, more compact thorax, and II

lack of complete dorsal thorax segmentation.

Type Locality and Type Specimens. Holotype female (No. 3211), paratype female (No. 3212), •

Stanford University Natural History Museum. Taken off Pt. San Luis, California; 112m.

Distribution. Puget Sound, Washington to Santa Cruz Island, northern Channel Islands, California;

4-154m. I
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I Figure 1.9. Callipaltenepacifica; A. body, dorsal, B. chelae andproboscis, C. oviger, D. terminal claw (after Hedgpeth, 1939).

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Family Nymphonidae

I Nymphon pixellae Scott 1912

i Figure 1.10

Nymphon pixellae Scott, 1912:206-209, pl. VII.----_lton, 1942a:4; 1942b:279-280, pl. 35.---Hedgpeth,

1941b:258.--Stock, 1955:216-217, fig. lb-d.---Child, 1979:40;1990b:42.

I Nymphon solitarium Exline, 1936:414-416, fig. 33a-d.

Nymphon variatum Hilton 1942a:4.--Hedgpeth, 1949:27 I, fig. 34b.

I Material Examined. California: Santa Maria Basin, off Pt. Sal; Sta. PJ-6 (1), Sta. PJ-10 (1).

Description. Thorax slender, elongate; segmentation complete, distinct; lateral processes moderately

I long, their diameter; separated byall apairs bout subequal their diameinterlength. exceptAbdomen for first and short, secosomewhat nd pair, which longerarethanseparlastatedthoracic by nearsegment, ly twice

but shorter than lateral processes of fourth legs; not fused to last thoracic segment, and inclined at about 30°.

I Proboscis shorter than neck, about one-third combined length of cephaion and thorax; cylindrical, with slight constriction at midlength, bluntly rounded.

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Neck slender and very long, 3 times length of remainder of cephalon; broadened distally I

accommodating insertion of chelifores and palpi. Ocular tubercle at base of neck, over and slightly behind

insertion of oviger bases, immediately anterior to lateral processes of first leg pair. Ocular tubercle low, •

height slightly less than twice basal diameter, with 2 equal pairs of large eyes occupying central half. Above I

eyes, tubercle attenuates rapidly to sharp point.

Chelifore scape long, reaching end of proboscis, broadening slightly distally. Chelae about three- II

quarters length of scape, overhanging end of proboscis; palm and fingers subequal in length, both f'mgers

attenuate, with acutely pointed crossing tips. Fixed finger bearing 24-30 distally pointing spines, moveable

finger with 40-45 similar spines. Palpi 5-segmented, slender, about twice proboscis length. Basal segment I

very short; segments 2 and 3 subequal, and about one-third longer than segments 4 and 5. II

Oviger 10-segmented, longer than combined length of cephalon and thorax if extended. Segments

1-3 short, together only one-third length of segment 4; segment 5 subequal to 4; segment 6 one-third length 1

of 5. Segments 7-10 comprising a strigillus, armed ventrally with rows of denticulate spines. Spines evenly

denticulate, tapered distally from origin of denticulation. Segment 10 bearing long terminal spine about half

length of segment. I

Legs slender, length 17 times diameter, bearing numerous short setae. Coxae together less than

10% of leg length; coxa 2 twice length of other coxae, and one-third femur length. Tibia I 25% longer than III

femur; tibia 2 nearly double femur length, narrower than segments proximal to it. Tarsus one-sixth length of 1

tibia 2, and two and one-half times propod length. Both tarsus and propod slightly narrower than tibia 2. II

Propod lacking heel and lamina, with short ventral setae only. Terminal claw one-third propodus length; am

auxiliary claws half length of primary. Iw

Biology. This species bears sufficiently long legs that it is frequently caught in trawl nets fished

over sandy mud bottoms offshore. The specimens examined were taken with a benthic grab sampler, and I1

are small for the species. Large specimens can have a leg spread exceeding 5 cm. In southern California •

Nymphon pixellae has been taken several times in association with the sand star Luidiafoliolata. Although

llll

these specimens were recovered from trawl catches, they were repeatedly found in the same situation, on

the dorsal surface of the star, and not elsewhere in the trawl collection. It seems incredible that so delicate an 1

II

organism could withstand the rigors of such handling without dislodgement, but it has happened on several

occasions. The pycnogonids remain in association with the star when In'st placed in an aquarium, but soon Ill

disappear (R. Velarde, pers. com. 1993). When live these animals have red-banded legs. II

U

Remarks. This is the common shallow shelf representative of the genus Nymphon in the northeast

Pacific. Nearly the same depth range is occupied by N. heterodenticulatum, but it is rarely reported. The two •

species are similar in many respects, but differ in the number of spines on the chelae, the relative length of |

the tarsi and propodi, the relative length of the oviger terminal spine, and the length of the neck. The

differences in shape of the oviger denticulate spines illustrated by Hilton (1942b) and by Hedgpeth (1941a) II

are of no diagnostic value as the illustrated range in spine morphology can be found on a single oviger. •

Unfortunately N. pixellae is a large species, and N. heterodenticulatum is a small delicate form, so the

Ill

specter of ontogenic change cannot be easily dismissed. Until full growth series of both species can be

assembled, the possibility that N. heterodenticulatum is a junior synonym of N. pixellae based on juvenile 1

females must be considered. A third species, Nymphon stipuIum was recently described from upper slope

depths in the northern Channel Islands (Child, 1990b). This species differs from both N. pixellae and N.

heterodenticulatum by the position of its ocular tubercle in advance of the oviger insertion, by proboscis •

shape, and by having tarsi shorter than propodi. Child (1994) added yet another species to the northeast R

Pacific fauna, Nymphon aculeatum. This deep-water species (2450-4100m) is blind, lacks an ocular tubercle,

has chelae longer than the chelifore scape, and also differs in leg and oviger spination from it's shallower •

water congeners.

1

Type Locality and Type Specimens. Type in British Museum of Natural History.

Distribution. Gulf of Alaska to San Diego; 59-196m. I

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I Figure 1.10. Nymphonpixellae; A. body, dorsal, B. oviger, C. palp, D. chela, E. terminal claw (afterHilton 1942a).

I Nymphon heterodenticulatum Hedgpeth 1941a

i Figure 1.11

Nymphon heterodenticulatum Hedgpeth, 1941a:447-449, pl. 1.----Child, 1990b:42.

I Material Examined. California: Santa Maria Basin, off Pt. Sal; Sta. PJ-1 (2), Sta. PJ-10 (2).

Description. Thorax slender, elongate; segmentation complete, and distinct; lateral processes

I mshort, oderately equalloinng,length separated to lastbythoracic more thansegment; their width, taperingslightly slightlydivergent; to bluntlyall pairs rounded subequal tip; lacking in length. setation, Abdomen and

inclined upwards at about 45°.

I Proboscis equal in length to neck and thorax, cylindrical, constricted slightly at two-thirds length. Neck long, slender, broadened distally for attachment of proboscis, chelae and palpi; more than

twice length of rest of cephalon, and equal to proboscis and thorax in length. Ocular tubercle at base of neck

I above insertion of ovigers, and just anterior to lateral processes of first legs. Two equal pairs of eyes located

near base of ocular tubercle, which is twice basal diameter in height, terminating in sharp anteriorly bent

point.

i Chelifore of which is chelae. Chelae about two-thirds scape nearly length proboscis, overhung by

scape length, slender, with fmger length subequal to palm. Chelae slender, with sharply pointed tips, with

17 small spines lining fixed finger, and 19 on moveable finger. Palpi 5-segmented, slender, nearly twice

length of proboscis. First short, 2 and 3 subequal and about 50% longer than

segment very segments segments
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Figure 1.U. Nymphon heterodenticulanon; A. body, dorsal, B. chela, C. oviger, D. terminal claw (after

Hedgpeth 1941a). i

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I 4 or 5. Oviger 10-segmented, male and female ovigers very similar; fourth and fifth segments longest and

subequal; segments 1-3 together half length of segment 4, and subequal in length to segment 6. Strigillus

I segments terminal spine 7-10 nearly of decreasing equal tolength; length.each bearing row of denticulate spines ventrally; segment 10 bearing

Legs long, slender; sparsely setose. Coxa 2 twice length of coxa 3, which is nearly twice length of

I coxa I; femur over half length of tibia 2, and about three-quarters length of tibia 1; tarsus long, equal in length to second coxa, and about one-third longer than propodus. Tarsus and propodus armed ventrally with

row of small spines, and sparsely setose dorsally. No trace of heel on propodus; lamina also absent; terminal

I claw slender, about half propod length; auxiliary claws slender, about half length of primary.

- Biology. Nymphonids are typically found on soft bottoms rather than on hard substrates.

Remarks. The examined specimens deviate in some respects from Nymphon heterodenticulatum

I described The of the Santa Maria Basin is the

as by Hedgpeth. tarsus specimens subequal to propodus

length. While the proportions of the chela fingers and palm axe the same in the present specimens as described

for the type, the entire chela of the Santa Maria Basin specimens is less slender than described by Hedgpeth.

I Spination of the chela fingers also differs, with spines larger and less numerous than described by Hedgpeth.

In other characters the present specimens agree with Hedgpeth's species, and they are treated here as variants.

i Type Locality and Type Specimens. Holotype female (No. 3215), and paratype female (No. 3216) in Stanford University Natural History Museum; paratype female in collection of J. W. Hedgpeth.

Distribution. California: off Pt. Sal to Catalina Island; 91-160m.

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Family Phoxichilidiidae

I Anoplodactylus erectus Cole 1904

i Figure 1.12

Anoplodactylus erectus Cole, 1904: 289-291; pl. X/V, fig. 12; pl. XXVI, figs. 1-9.--Stock, 1975b:74-76,

figs. 22, 23.

I Material Examined. California: Santa Maria Basin; off Pt. Sal, Sta. R8 (3); off Purisima Pt., Sta.

42 (1); off Pt. Arguello, Sta. 65 (1).

I Description. slender, elongate; segmentation complete distinct; lateral processes

Thorax and of

moderate length, separated by about their diameter, diverging slightly; those of first legs longest, with

length decreasing slightly in each remaining pair. Abdomen tapering from about midlength to blunt tip;

I about half length, than lateral of fourth inclined at 60_, and cephalon longer processes legs; upwards nearly

beating few very short setae.

Proboscis about twice neck length, cylindrical, blunt, increasing slightly in diameter distally; angled

I downward from cephalon at about 15°.

Neck as long as rest of cephalon, slender, bearing ocular tubercle dorsally at end. Ocular tubercle

I height slightlyabout lowertwice and larger basal than diameter; posterior 2 pairs pair.of eyes situated under its bluntly pointed apex; anterior pair

Chelifore scape inseaed at base of ocular tubercle; scape cylindrical, of 1 segment, longer than

I neck, palm rectangular, but shorter shorter than cephalon; than or equal lackingto fixed tubercles, finger and length; withfingers few very unequal, short moveable scattered finger setae. longer, Chelae more with

strongly curved distally, and bearing a group of setae dorsally. Neither finger with spines or teeth. Chelae

i attached to scape at nearly right angle, projecting ventrally. Palpi lacking.

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Oviger in male only; 6 segmented, slender, as long as combined cephalon and thorax. Basal segment I

with length and diameter subequal; segment 2 only two-thirds diameter and twice as long as segment 1.

Segment 3 bent at nearly right angle to segment 2, projecting along underside of body to rear. Segments 3- i

6 all of smaller diameter than segments 1-2; segments 3, 5 and 6 setose; segment 4 lacking setae. Segment il

3 about half oviger length. Segments 4-6 forming strigillus-like loop, with segments 5 and 6 bent back along

segment Legs 4. | long, about twice combined length of cephalon and thorax; slender, length about 16 times

diameter. Lateral processes with single conical tubercle distally. First and third coxae subequal; coxa 2

twice their length. Femur about 50% longer than tibia 1, 25% longer than tibia 2. and twice the length of !

coxa 2; tarsus short, about 10% length of tibia 2; pmpodus curved, about half length of femur. Terminal II

claw as long as sole, nearly equal to propodus length; auxiliary claws very short, about one-eighth length of

primary. I Coxae lacking dorsal tubercles; male genital aperture at end of ventral genital spurs distally on

second coxae; female coxa 2 swollen distally, with genital aperture flush with segment surface. Femur

beating tubercle dorsally at distal end in male, but not in female. Femoral cement gland beyond segment

midpoint, aperture at end of tapering duct projecting above femur by about one-third diameter. No other U

tubercles. Legs sparsely scattered with short setae. Propodus with heel well defined, but short; bearing 2

spines and 3 setae. Sole beating a lamina for distal one-third, with between 7-11 distally curved spines

proximally. Propodus with row of setae dorsally. roll

Biology. The life history of Anoplodactylus erectus has been detailed by Hilton (1916), who found !

them associated with the hydroid Tubularia crocea. Larvae ofA. erectus, once hatched from the egg mass, I[

pierce the body wall of the hydroid and enter the digestive tract. Eventually these endoparasites eat their u

way out, becoming adult predators on the hydroid. The shape of the terminal segments of the oviger are held ill

recurved into a loop, as are those of species whose spinose oviger ends form a true strigillus. Male _l

phoxichilidiids have, however, not been observed to use the oviger in grooming, and ovigers are lacking in UP

the female.

a.

Remarks, This is by far the most commonly encountered of the three species of Anoplodactylus i

taken in the Santa Maria Basin samples. To some extent this is due to its occurrence in the intertidal zone,

U

and in association with hydroids in the fouling community. It can be distinguished from the co-occurring

Anoplodactylus species by having one tubercle on the lateral processes; Anoplodactylus oculospinus has li

two, and A. pacificus none. It differs from A. nodosus, which was not represented in these collections, in m

lacking the tuberculations of the tibiae which characterize that species. m

Type Locality and Type Specimens. Holotype male, allotype female, (University of California i

No. 19,514); San Diego, California.

Distribution. British Columbia, Canada to Punta Arenas, Chile; intertidal-107m. In

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I Figure 1.12. Anoptodactylus Cole, 1904). erectus; A. body,lateral, B. chela, C. oviger, D. terminal claw, E. leg (after

,_i Anoplodactylus oculospinus Hilton 1942d

i Figure 1.13

Anoplodactylus oculospinus Hilton 1942d:72-73.--Child, 1975:196-198, fig. 3.

i Anoplodactylus nr. pacificus (in part) Lissner et al., 1986:D-26

Material Examined: California: Santa Maria Basin, offPurisima Pt., Sta. 16 A/B (2); Sta. 20 A/B (1).

i Description. Thorax slender, elongate; segmentation complete, distinct; lateral processes of moderate length, separated by about their diameter, diverging slightly; those of ftrst leg pair slightly longer than

others. Abdomen tapering slightly to blunt tip, about one-third cephalon length, equal in length to lateral

I processes of fourth legs, inclined upwards at about 30°, with single dorsal seta.

i 35
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Proboscis about twice length of neck, cylindrical, blunt, tapering slowly from base to about three- I

quarters of length, then enlarging again to nearly basal diameter at tip.

Neck somewhat shorter than rest of cephalon, relatively robust, and shouldered into cephalon for I

nearly half of length. Ocular tubercle at anterior cephalic margin, capped by anteriorly directed spine, with U

2 pairs of eyes at about midheight. Anterior pair of eyes somewhat below posterior pair, flanked by small

laterally directed tubercles. Ocular tubercle short, with height and diameter about equal. I

IB

Chelifore scape of single segment, about 3 times chela length, overhanging proboscis by more than

chela length; scapes approximated basally, bearing low seta-tipped tubercles both dorsally and laterally.

Chelae with palm rectangular, longer than fingers, with neither dorsal or ventral setae. Both moveable and I

fixed fingers strongly curved, attenuate, ending in sharp crossing tips. Fixed finger with several dorsal II

setae, but no ventral setae; moveable f'mger with 2 dorsal setae, and no ventral setae. Neither finger with

spines or teeth. Palpi lacking. Adult oviger unknown; juvenile male with 5-segmented oviger, im

Each lateral process with 2 seta tipped tubercles, one anterodistal, and one posterodistal. Legs

intermediate, neither slender or robust, length about 9 times diameter. First and third coxae subequal, second R

about 50% longer; femur and tibiae subequal, about 3 times length of coxa 2; tarsus very short, barely one- •

sixth length of propod; propodus subequal to coxa 2.Terminal claw as long as sole, fully four-fLfths propodus

length; auxiliary claws strong, but very short, only 10% of primary length. El

First coxae with spine tipped tubercles as on lateral processes; second and third coxae without I

tubercles or setae; femur slightly inflated, bearing scattered short setae; both tibiae with row of short lateral

setae, tibia 2 also bearing row of short setae ventrally; both with long dorsodistal seta. Propod with 4 dorsal ttl

setae, and with a distinct heel beating 2 spines and 2 large setae; sole with 10-11 anteriorly curved spines _

and 7-8 setae; lamina short, about one-eighth length of propodus.

I

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B !

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l_N 1.13. Anoplodactytus oculospinus; A. body, dorsal, B. chela, C. terminal claw, D. leg (after Child,

1975). I
!

i Biology. The type was taken from rock substrate by dredging; the specimens examined here were

associated with medium sized rocks (over 1000 cm 2 surface area) that were collected with manipulating

I arms on Remarks. a submarine.The present collection appears to include the only specimens of Anoplodactylus

oculospinus reported since the type female was described (Hilton 1942d). As no adult male is reported here,

i The detailsjuvenile of the male oviger,examined the femoral was cement not sufficiently gland andmature duct, sexually and the male to predict genitalwhat aperture these remai secondary n unknown. sexual

characteristics would be like in the adult. Of the three species of Anoplodactylus in the present collection, A.

I opaired culosptubercles inus is theoneasies the lateral t to recognize process. andThecoxa combination one quicklyof long sepacrhel atesa pitalm, fromlathe teralothers. cephaloLike n tubercles, A. pacificus an,d

more A. oculospinus probably await discovery in existing collections mixed with A. erectus.

I 91m; USNMType 81494. Locality and Type Specimens. Holotype female; offMoss Landing, Monterey Bay, California;

Distribution. Central Califomia, from Monterey Bay to off Purisima Pt.; 91-123m. I

Anoplodactylus pacificus Hilton 1942d

i Figure 1.14

Anoplodactflus paeificua Hilton 1942d:73.--Stock, 1955:242-243.---Child, 1975:198-201, fig. 4. m

I Anoplodactylus nr. pacificus (in part) Lissner et aI., 1986:D-26.

Material Examined. California: Santa Maria Basin off Purisima Ft., Sta. 14 C/D (1). Western

I: Santa Barbara Channel, Sta. S1 C/D (1).

Description. Thorax slender, oval; lateral processes long, separated by more than their diameter;

I tubercles pair a bit shorter. absent, bAbdomen ut with 2 small slender, spines aboutdistal halfly.cephalon Lateral pro length, cessesandof

longer first legthanpairlateral longest; processes each successive of fourth

legs, tapering to a relatively acute terminus bearing several distal setae, and held vertically.

slightly Proboscis swollen atnearly bend. twice length of neck, cylindrical, blunt, bent upwards at about one-third of length,

Neck equal in length to rest of cephalon, slender, with line continued anteriorly by tapering and

_1_ terminally acute ocular tubercle. Two pairs of large eyes borne near base of ocular tubercle, the latter about

m twice basal diameter in height. Chelifore scapes touching basally, long, 1-segmented, with row of dorsal

setae, and distal tubercle. Scape nearly 3 times chela length; with distal one-third overhanging proboscis.

I Chelae with palm rectangular, subequal to fingers, bearing several dorsal setae. Both moveable and

fixed fingers strongly curved, attenuate, ending in sharp crossing tips. Fixed finger bearing seta dorsally

near base; moveable finger with 3 setae ventrally, and 4 longer setae dorsally, without either spines or teeth.

i Palpi lacking.

Oviger 6-segmented, thin, basal segment short, relatively wide; segment 2 with length 9 times

width; segment 3 twice length of segment 2; segments 4 and 5 subequal, half length of segment 2; terminal

I segment a button, one quarter length segment setae on segments 2-4, segment

minute of 5. Scattered 5

strongly setose on concave surface, segment 6 with 2 small setae.

i Legs slender, length about 13 times diameter; coxa 1 and 3 subequal, coxa 2 2-3 times length of 1

and 3; femur, tibia 1, and tibia 2 subequal; tarsus very short, propodus about one-third length of 3 long

segments; terminal claw about two-thirds propod length; auxiliary claws minute, little more than setae.

i Coxa 2 bearing distal tubercle and both dorsal and ventral setae. Femur and tibiae also with short setae

| 37
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dorsal and ventral. Femoral cement gland at about midsegment, produced into raised, slightly flared duct. i

Heel poorly differentiated, bearing 2 curved spines and 2 setae. Propodus with about 15 sole spines bent

anteriorlyB at their ends; lamina less than one-third propod length. I

iology. The type was taken in a trawl, presumably over soft bottom. Both specimens examined

here were associated with small rocks collected with a submarine.

Remarks. The original description was cryptic and probably prevented recognition ofAnoplodactylus I

pacificus until the redescription and figuring of the type by Child (1975). Only Stock (1955) managed to

identify his specimens from the original description. This off-shore species is perhaps more common than

the number of specimens reported would indicate. It is found with at least two similar congeners, A. erectus I

and A. oculospinus, both represented in the present collections. Anoplodactylus pacificus can be readily

distinguished from both by the absence of tubercles distally on the lateral processes.

Type Locality and Type Specimens. Holotype male; off Avalon, Santa Catalina Island, southern i

California; 86m; USNM 77078.

Distribution. Santa Maria Basin, central California to Southern California Bight: 74-105m. I

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FiN 1,14. Anoplodactylus pacificus; A. body, dorsal, B. chela, C. oviger, D. terminal claw (after Child,

1975). i

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. Family Pycnogonidae

Pycnogonum rickettsi Sehmitt 1934

I Figure 1.15

Pycnogonum rickettsi Schmitt, 1934:62-65, fig. 1.---Hilton, 1943b: 19.--Hedgpeth, 1975:417, 424; pl. 99,

I• fig. 3.--Hedgpeth and Haderlie, 1980:638, fig. 27.2.

Material Examined. Central California off Morro Bay, Sta. 27 A/B (3); off Oso Flaeo, Sta. 25 A/

I B (1); offPt. Arguello, Sta. 6 A/B (1). Western Santa Barbara Channel off Tajiguas, Sta. 1 C/D (3). Description. Thorax compact, elongate; segmentation complete and well-defmed. Lateral processes

short, subequaI, separated by less than half their diameter, diverging slightly. Integument reticulate. Cephalon

I and first two thoracic segments bearing tall, conical middorsal tubercles; tubercles shortest on cephalon, and increasing in height posteriorly. All 3 middorsal tubercles taller than ocular tubercle. Abdomen short,

tubular, horizontal, between and not exceeding length first coxae of fourth leg.

I more length cephalon, blunt, at midlength; proboscis

Proboscis than twice of fusiform, widest

integument reticulate.

i No neck apparent; ocular tubercle near anterior cephalic margin, short, with height and basal diameter subequal, with 2 equal pairs of eyes. Chelifores absent inboth adult and juvenile, present only in protonymphon

larva. Palpi lacking. Ovigers only in male, 9-segmented with a terminal claw; basal 3 segments with width

i exceeding more length; segments 4 and 5 longer, with length twice width; strigillus segments (6-9) progressively slender. Segment 3 with 1 dorsal and 1 ventral seta, segments 5 and 6 with 1 ventral seta, segment 7

with 1dorsal and 3 ventral setae, segments 8 and 9 with field of very short ventral setae. Segment 9 tapering

to basal diameter of terminal claw.

i Legs robust, length about 8 times diameter. Coxae subequal on first 3 leg pairs; coxa 1 both longer

and broader than coxae 2 and 3 on fourth pair of legs. Femur and tibia 1 subequal and about twice coxa

length, tibia 2 about two-thirds femur length on legs 1-3. On leg 4 femur and both tibiae subequal, about

50% longer than coxae 2 and 3. Tarsus short, one-third to one-quarter length of propodus, beating field of

short setae ventrally. Propod lacking heel and enlarged heel spines; sole with numerous very small setae,

i but lacking spines and lamina. Terminal claw about half propod length; auxiliary claws lacking. Lateral processes beating low conical tubercle distally, largest on fourth legs and decreasing in size on more anterior

leg pairs. Coxa 1with distal tubercle like that of cruriger; femur and tibia 1 each bearing a pair of dorsodistal

i tubercles,Biol tibiaogy.2 unType armed. specimens were taken from soft substrate offshore, in association with the large

plumose anemoneMetridium. The present specimens were taken on rocks collected by submarine. Although

I n(pone rimarily of theMetridium collected)rocks were sabund upported ant inMetridiu the sampled m, or other habitatlikenearby ly ane(pers. moneobs.). prey for Association Pycnogonumwith, Metridium anemones

senile, Anthopleura xanthogrammica, Aglaophenia spp, and Clavelina huntsmani are reported in the Pacific

i NorthwesRet marks. by Hedgpeth The (1987). report that Pycnogonum rickettsi ranges to the Galapagos Islands (Hedgpeth, 1951),

seems to represent a misreading of Hilton's (1943b) statement that several specimens had been taken from

I Santa channelCruzislands Island(Hewatt, by Hewatt 1946), andandWilli notamsto one in 1of939. theTGalapagos his report regroup. ferredChild to oneandof Hedgpeth the southe(1971), rn California whle

not specifically addressing the prior record, did not mention P. rickettsi as a member of the Galapagos fauna

i in their review.

i 39
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I

i

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I

a

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Figure 1.15. Pycnogonum1934). rickettsi; A. habitus, dorsal, B. body, lateral, C. oviger, D. leg (after Schmitt, I

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40 |
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I This species is clearly distinguished fromP, stearnsi, it's only congener in northeast Pacific temperate

and boreal waters, by its much higher middorsal tubercles, and by the reticulation of the integument. The

I two any single speciesanemone do co-occur, (Hedgpeth and, while and reported Haderlie,from1980). the same prey species, have not been reported together on

Type Locality and Type Specimens. Holotype male; off Pacific Grove, Monterey Bay, central

I California; 120m; USNM. Distribution. Puget Sound to the Southern California Bight; intertidaI-117m.

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m

Losina-Losinsky, L. K. 1961. On the fauna of Pantopoda from the Far-Eastern seas of the USSR. Explorations B

of the Far Eastern Seas of the USSR 7:5-I 17.

Lotz, Von G. and D. Biickmarm. 1968. Die Hiiutung und die Exuvie yon Pycnogonum litorale (Str6m) i

(Pantopoda). Zoologisches Jahrbiicher, Abteilung fiir Anatornie und Ontogenie der Tiere 85:529-536. i
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I Manton, S. M. 1978. Habits, functional morphology and the evolution of pycnogonids. Zoological Journal

of the Linnean Society 63(1):1-21.

I Marcus, E. duBois-Reymond. 1952. A Anais da Academia Brasileira de Ciencias

hermaphrodite pantopod.

24(1): 23-20.

i Martin, J. ScypW.hozoa) and H.inKthucke Sou . 199the1.rnFauCalnaliforass niaociBight, ates ofwiathn unnodetesscrionbedunususpecalieosccuofrrenChrys cesaora of other (Cnidawarria, m

water species in the area. Bulletin of the Southern California Academy of Sciences 90(3):89-101.

I Mauchline, J. 1984. Pycnogonids caught in bathypelagic sarnples from the Rockall Trough, northeastern Atlantic Ocean. Journal of Natural History 18:315-322.

I Mercier,Bartholomea A. and J. -F. annulata Hamel. .1994 Canadian . Deleteri Journal ous effec of Zoology ts of a pycnog 72(7):onid1362-1364. on the sea anemone

Miyazaki, K. and T. Makioka. 1993. A case of intersexuality in the sea spider, Cilunculus armatus

I (Pycnogonida; Ammotheidae). Zoological Science 10:127-132.

Morgan, E. 1977. The swimming ofNymphon gracile (Pycnogonida): the energetics of swimming at constant

I depth. Journal of Experimental Biology 71:205-211. Nakamura, A, K. and K. Sekiguchi. 1980. Mating behavior and oviposition in the pycnogonid Propallene

longiceps. Marine Ecology - Progress Series 2 (2): 163-168.

I Ohshima, A. 1927. NymphoneUa tapetis, n. g., a pycnogon parasitic in a bivalve. Annotationes Zoologicae

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I Peattie, M. E. and R. Hoare. I981. The sublittoral ecology of the Menai Strait. II. The Halichondria

sponge

panicea (Pallas) and its associated fauna. Estuarine, Coastal and Shelf Science 13(4):621-635.

I Piel, W. H. 1991. Pycnogonid predation on nudibranchs and ceratal autotomy. Veliger 34(4):366-367. Redmond, J. R. and C. D. Swanson. 1968. Preliminary studies of the physiology of the Pycnogonida. Antarctic

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I Richards, P. R. and W. G. Fry. 1978. Digestion in pycnogonids: a study of some polar forms. Zoological

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I Richter, S. 1982.Zur Ultrastruktur der seitlichen Sirmesorgane am Augenhiigel vonAnoplodactyluspygmaeus

• (Pycnogonida). Helgol_nder Meeresuntersuchungen 35:465-478.

i Russel, DAmmothea . J. 1990. Hhilgendorfi ost utiliza)tioparasitic n duringononthe togehydroid ny by twoEucopella pycnogonid evertaspecies (Coelenterata: (TanystylumCarnpanuladidae). duospinum and

Bijdragen tot de Dierkunde 60 (3/4):215-224.

I Rytand, J. S. 1976. Physiology and ecology of marine bryozoans. Advances in Marine Biology 14:285-443.

Salazar-Vallejo, S. I. and J. H. Stock. 1987. Apparent parasitism of SabelIa meIanostigma (Polychaeta) by

i AmmotheUa spinifera (Pycnogonida) from the Gulf of California. Revistas de Biologia Tropical 35 (2): 269-275

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I Forschung, Leipzig 32(4):633-658.

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I Journal of the Washington Academy of Sciences 24(1):61-70.

I 45
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Schram, F. R. 1986. Crustacea. Oxford University Press. New York. 606 pp. I

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u

Schram, F. R. and J. W. Hedgpeth. 1978. Locomotory mechanisms in Antarctic pycnogonids. Zoological _1_

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in research. Academic Press, London.

I Wyer, D. and P. E. King. 1974. Feeding in British littoral pycnogonids. Estuarine and Coastal Marine

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Anzeiger (Leipzig) 228:220-228.

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iiiiIIII _ i
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SUBPHYLUM CRUSTACEA

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2. THE DECAPODA

I by

I Joel W. Martin 1 and Todd L. Zimmerman:

i Introduction

Decapod crustaceans are among the most common and easily recognized invertebrate groups,

i representing a large number of species found in many habitats. Because of their popularity, and because some

groups are of considerable economic importance, there are numerous guides to the identification and natural

history of decapods. However, most of these guides are restricted either to specific areas outside southern

I California (e.g., Austin Williams' 1984 guide to the decapods of the east coast of the United States) or, if

pertaining to southern California, are restricted to one or two families or genera (e.g. Zmarzly's 1992 review

of the pinnotherid crab genus Pinnixa). It is surprising that for the California coast, an area that has been

I subjected to much study, no updated and thorough treatment of the Decapoda exists subsequent to Waldo Schmitt's (1921 ) classic (but now outdated, and long out of print) volume The Marine Decapod Crustacea of

California. Several guides to, or treatments of, the west coast decapods have appeared since Schmitt's

publication (e.g. Kozloff, 1987; Allen, 1980; Jensen, 1995, etc.), but these tend to include only those species

I

restricted to more shallow waters (and therefore more likely to be seen and collected by naturalists), and so

they are of limited use in identification of the deeper fauna.

talk

I The most surprising result from the present study of the MMS collections is the relatively small

lp

number of species encountered, most of which are relatively well known, compared to those anticipated (for

expected source pool see Wicksten, 1989). For many of the taxa encountered, the descriptions and biological

I information have been summarized from older literature, such as Garth's (1958) monograph on Pacific "Oxyryncha".

One of the most valuable assets in identification and taxonomy of California invertebrates are the

l members of the Southern California Association of Marine

Invertebrate Taxonomists (SCAMIT).These

biologists have been responsible for publishing numerous new records and taxonomic changes to the knowledge

of southern California species. Primarily because of the active input of information from that group, the

I list should not be considered a definitive checklist of the fauna of the Santa Maria Basin. Despite the

present

size of most decapods and the interest they generate, it is clear thatthere still is much to learn about them, even

from such populated regions as southern California.

I The current report is a list of species identifications and a corresponding compilation of information

that stems primarily from preexisting literature. For that reason and by way of this disclaimer, this report is

i not to be considered to have met the criteria for publication as outlined in Article 8 of the International Code of Zoological Nomenclature, third edition (1985).

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1Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007

I 2Department of Biology, University of California, Los Angeles, California 90024-1606

I 49
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General Biology and Ecology I

l

Decapods occupy such a wide range of habitats and ecological "niches" that no general statements of _l

their biology and ecology can be made without many caveats inserted. The group includes scavengers, strict

w

herbivores, predators (both sit-and-wait predators as well as hunters), filter feeders, symbionts, and probably

any other feeding mode that could be imagined. The majority are benthic, although many species, particularly B

among the shrimp, are pelagic, at least during part of the life cycle; there are of course freshwater, terrestrial

and semiterrestdal species as well, but these need not be mentioned for purposes of this report. Most of the /

species mentioned in this report are benthic, macrophagous animals, as might be guessed from where and how I!

they were collected (see Volume 1 of this series). In general the group might best be described as opportunistic l

scavengers, meaning that, from what little we know, they will eat just about anything they happen upon. Some

of the species in this report are almost certainly more specialized in their feeding habits than the above I

statement would seem to indicate. Commensal pinnotherid crabs, as one example, are often highly dependent II

upon their "host" as a source of nutrition, yet for some no host is known. Although many species of decapods

will eat shrimp andjust about any other proffered food item in captivity, it is not always appropriate to extend I

observations made in the laboratory to what might be happening in the field. U

Terminology and Glossary I

Ii

For decapod crustaceans, the glossary of Williams (1984), although written to assist in the identification l[

of decapods from the east coast of the United States, serves equally well for identifying characters used in the

taxonomy of west coast decapods. We have borrowed verbatim the glossary found in his introductory section "ll

(Williams, 1984: 8-17), and his schematic labeled diagrams of a shrimp and crab are reproduced here as |

Figures 1 and 2. We have added to Williams' glossary the relatively few terms used in this report that are not

found in his glossary. In the following section, any word and its definition taken directly from Williams (1984) iI

is followed by the notation [ABW]. Other terms anddefinitions have been taken from various sources, among .1

which are McLaughlin (1980) [McL] and Moore and McCormick (1969, in Glaessner, 1969) [M&M], which

are also good starting points for learning the terminology of other crustacean taxa. Some of the more descriptive m

terms are included here as well, and definitions for some of these have been taken from the American Heritage 1

Dictionary of the English Language [AHD]. When no concise definition could be located, we have used our

_J

own; these are designated as [M&Z]. Figures referred to in the definitions are those of Williams (1984) and

not of the current report. Iii

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I

i prbit eyesfalk ro abdominal segments

corneo ..,p.g.

I a.s. _- _.s.

carapace 5

an_ennule _.s.

_s. pleura

I antenna

d.

i third maxit|ipe c)

subcheta- m. .x.

plaopods I che|Q- c_ P

i subdivided c(irpus e_d_

uropods i

teLson

1

Figure 2.1, Schematic drawing of shrimp in lateral view: ai, appendix intema; as, antennaI spine; asc, antennal

I scale; b, basis; bs branchiostegal spine; cp, carpus; cx, coxa; d, dactyl; end, endopod; ep, epipod; ex, exopod; hs, hepatic spine; i, ischium; m, merus; p, propodus; pg, postorbital groove; ps,

pterygostomian spine; ss supraorbital spine; st, stylocerite (from Williams 1984, modified from

i Holthuis 1955; Schmitt 1921). ,

I

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I Glossary

I Abdomen. Trunk somites (tagma) between thorax Antennule. One of a pair of appendages of the

and telson; somites with or without limbs; first cephalic somite; syn., 1st antenna.

I syn., pleon. [McL] IMcL] Acicle. Antennal scale reduced to a spine. [ABW] Areolation. Demarcated and usually elevated

regions of the brachyuran carapace (see M.

I Afferent passes channels. Openings through which water J. Rathbun, 1930, page 6). [M&Z] to gills. In brachyuran crabs, usually

opening behind pterygostomian regions and Basis 0aasipodite). Second article (from body) of

I inOxystomata front of chelipeds in whichexthey ceptopen in certain at leg endorofmaxilliped. limb. [ABW]Sixth segment from distal

anterolateral angles of palate or endostome.

[ABW] Basieerite. Spine on dorsal side of basis of

.I antenna; sometimes more lateral than dorsal.

I Apodeme. Any cuticular ingrowth of body wall. [ABW]

[ABW] Branehioeardiae groove. Groove separating

I Appendix interna. Small separate branch on branchial and cardiac regions. [ABW]

mesial side of pleopodal endopodite, usually

tipped with hooks which interlock with Branehiostegal spine. Spine on anterior edge of

I carapace, or near it, immediately below opposite member in swimming. [ABW] branchiostegal groove (Fig. 2). [ABW]

Appendix maseulina. Accessory male organ Branchiostegite. Part of carapace not coalesced

I located mesially on second pair of pleopods between endopodite and appendix intema, ventrally with thoracic somites, but overhanging on each side as coveting for gill

[ABW] chamber. [ABW]

t Arthrobranehiae. Gills attached to articular

membrane between coxa of appendage and Bueeal cavity. Cavity on ventral surface of body

body wall. [ABW] in which mouthparts are situated; bounded

I A anteriorly by epistome, laterally by free ntepenultimate. Second from the last; third from edges of carapace. Within this "frame" lie the

the end in a series. [AHD] mouthparts, which in most Brachyura are

I Antenna(e covered by operculiform third maxillipeds. ). One of a pair of appendages of the [ABW]

second cephalic somite; syn., 2nd antenna.

[McL] Capitate. Enlarged or globular at end. [AHD]

I Antennal scale. See stylocerite. Carina. Keel-Iike ridge or prominence. [ABW]

Carpus (earpopodite). Third article from distal

I end of leg. [ABW]

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I Figure 2,2. Schematic drawingof brachyurancrab in: A--dorsal view; areasof carapaceindicated;legs of right

It side onlyshown;b,basis; cp, carpus;d, dactylus;i,ischium;m, meres; p, propodus. B---ventralview;

areas of carapaceindicated;legs of leftside onlyshown;b, basis; cp, carpus;cx, coxa;d, dactyins;end,

i endognath;ex, exognath;i, isehium; nLmerus; p, propodus(fromWilliams, 1984).
!

Chela. Arrangement of distal 2 articles of Epipodite. Outgrowth of first 7 thoracic coxae. I

crustacean limb in which terminal article is [ABW]

opposed adaptatioton farticle or grasppreceeding ing. In trueit inchela, an Epistome. Antennal sternum mainly represented I

by epistome, a plate of varying shape lying

terminal 2 articles shaped as fingers, one betweeen labrum and bases of antennae. In

cl article osing(dactyl) against usually other. Incloses subcheagainst la, terminal distal shrimps, comparatively narrow and I

surface of penultimate article (propodus). separated on each side from lateral portions

of carapace by exhalant branchial channels.

[ABW] In Astacidea and Anomura, broad and in I Chelipeds. Pair or pairs of thoracic legs behind contact with carapace on each side; in

maxillipeds; bearing chelae, or pincer-like Brachyura, becoming firmly united with

claws, and often stouter, sometimes much carapace. The epistome forms the anterior I

stouter, than other legs. [ABW] part of the buccal frame. [ABW] 

Coxa (coxopodite). First or proximal article of leg Exognath. Lateral or secondary branch of I

or maxilliped. [ABW] maxilliped. [ABW]

Dactyl (dactylns or dactylopodite). Terminal or Exopod. see Exopodite.

distal article of leg; movable finger of I

cheliped. [ABW] Exopodite. Lateral ramus of bimmous appendage. [hBW]

EfferentpassesChannelgills.ChannelSopeningsthr°ughat S'from sidesWhiChof water Fingers (digits). Narrow scissorlike blades of I

endostome, except in Section Oxystomata in pincer end of cheliped, movable finger being

dactyl, and fixed finger the terminal part of

which[ABW]they open at middle of endostome, propodus. [ABW] I

Endognath. Inner or principal branch of a Flagellum. Multiarticulate distal portion of

maxilliped. [ABW] antennule, antenna, or exopod. [McL.] ii

Endopodite. Mesial ramus of biramous Front. Frontal portion of carapace; that portion of

g

appendage. [ABW] crab carapace lying between orbits. [ABW]

Endostome. Part of epistome forming palate in Frontal teeth. True frontal teeth; those teeth I

brachyurans and usually separated from originating on front but exclusive of inner

epistome proper by transverse ridge. [ABW] orbital teeth. [ABW] I

Epibranchial (epibranchial region). Part of Gastric region. Large median area, in crab

porcellanid (crab) carapace situated behind carapace, bounded behind by cervical suture,

orbit and above metabranchial region, laterally by hepatic regions, andanteriorly by l

Region situated between cervical groove and frontoorbital regions. Divisible into following

linea anomurica. Often a strong spine on subregions or lobes: epigastric, protogastric,

region, referredto as epibranchial spine, mesogastric, metagastric, and urogastric. I

[ABW] [_W]

Epigasgastric tric lobes. Anterior lobes or subregions of Genital region. See urogastric lobe. [ABW] i regions. [ABW] Gonopore. Outlet for genital products; syn.,

Ephnere. Lateral part of wall of body somites sexual pore. [McL.]

situated between tergum and insertion of Hand (chela). Propodus and dactyl of cheliped. I'

appendages. [ABW] [ABW]

I
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I Hepatic region. Small (paired) subtriangular, Metabranchial (metabranchial region). That

anterolateral region, wedged between region of porcellanid (crab) carapace situated

I branchial margin ofand carapace gastrico regions, and either below linea anomurica and, therefore, not

r margin of orbit in completely united with main part of

Brachyura. [ABW] carapace. [ABW]

I Hepatic spine. Spine on hepatic region in shrimps Metagastrie lobe. Posterolateral lobe or subregion (Fig. 2). [ABW] of gastric region; often not defined. [ABW]

i Holotyofpe. The single specimen used as the basis Molar process. Grinding portion of gnathal lobe of the original published description of a mandible; syn., pars molaris. [McL.]

species and later designated as the type

specimen. [AHD] Obsolescent. Vestigial. [AHD]

I Incisor process. Biting portion of gnathal lobe of Ocellus. Little eye, distinct from main organ of

mandible; syn., pars incisiva. [McL.] vision. [ABW-]

i Ischium (ischiopodite). Fifth article from distal

Ocular peduncle. Eyestalk. [ABW]

end of leg; usually first large article of Orbit. Cavity in carapace containing eye. [ABW]

i Keel, see maxilliped. Carina. [ABW] Orbital f'tssure, see orbital hiatus.

Orbital hiatus. Gap in orbital margin of carapace

i Lacinia(e). Inner distal spiny lobe of second at its lower (inner) angle. [M&M]

segment of maxillule. [McL.] Orbital region. Narrow space bordering upper

Linen anomuriea, L, homoliea, L. thalassinica, margin of orbit; not always distinguishable.

I Longitudinal groove or uncalcified line on [ABW] carapace which may form a hinge; such lines

found in various groups, from which come Palate. Roof of buccal cavity in crabs. [ABW]

I the names. [ABW] Palm. Proximal part of propodus of chela. [ABW]

Mandible. One of the third pair of cephalic Palp. Usually one ramus (endopod), sometimes

i M appendages, used to masticate food. [McL.] both, and basis, reduced distally to 1 to 3 andibular palp. Distal articulated part of segments, associated with mouthparts.

mandible used in feeding or cleaning. [McL.] [McL.]

I Manus. Broad proximal part of propodal cheliped;

Peduncle. A stalklike structure. [AHD]

syn., palm. IMcL.] Pereiopod (also spelled peraeopod or pereopod).

I Maxillipeds. Paired appendages modified for Thoracic appendage used in locomotion; feeding on 1st, up to 3rd, thoracic somites, syn., ambulatory leg, walking leg. [McL.]

usually fused to cephalon. [McL.] Petasma, Endopodite of first pleopods in male

I Maxillule. One of apair of the 4th cephalic

Penaeidae; a complicated membranous plate

appendages, usually serving as a mouthpart; beating coupling hooks mesially which

syn., 1st maxilla. [MeL.] interlock with those on member of opposite

I Meru side; may terminate distally in various s (meropodite). Fourth article from distal complex-shaped lobes. [ABW]

end of leg; sometimes called arm of cheliped.

i Phyllobranehia(e). Gill with leaflike filaments. [AWl [McL.]

Mesogastrie lobe. Median lobe or subregion of

gastric region; pentagonal in form, and with

I long, narrow, prolongation. [ABW]

anterior
!

Pleurobranehia(e). Gills attached to lateral wall Pubescence. A covering of down or short hairs. 1

of body dorsal to articulation of appendage. [AHD]

[ABW]

Rugose. Having a rough or ridged surface. [AHD] i

i

Pleuron. See ep_ere. Scaphoeedte. Antennal scale (Fig. 2). [ABV_

w

Podobranehiae. Gills attached to coxa of Ill

appendage. [ABW] Styloeerite. Spine or rounded lobe on lateral I

aspect of basal article of antennule. [ABW]

Postorbital groove. Groove on carapace behind

orbit and more or less parcel to margin of Subhe_fie region. A_a below hepatic region and I

orbit (Fig. 2). [ABV_ anterolateral bo_er of carapace. [ABV_

Suborbi_ spine. Spree on lower dm of orbit

Propodus (propodite). Second article from distal I

(Fig. 2). [ABW]

end of leg. Propodus of cheliped consists of I

palm and narrower fixed finger. [ABW] Sulens, sulei. Groove or furrow. [McL.]

Prosartema (dorsal eye brush). Long, thin, Suture. Weakly calcified areas of integument I

ciliated lobe arising dorsally from separating at ecdysis. [McL]

l

proximomesial border of first antennular

segment and extending anteriorly; found in Supraorbital spine. Spine above and behind orbit

Penaeidae. [ABW] (Fig. 2). [ABW] II

Protogastric lobe. Anterolateral lobe of subregion Telson. Terminal somite of abdomen. [ABW] I

of gastric region. [ABW] Tergite. Dorsal plate of segment. [ABW] II

Protopodite. Peduncle of an appendage; in Thelycurn. External seminal receptacle, variously il

unmodified form, consisting of one coxal and developed, lying on sternum of thorax and i_

one basal article. [ABW] formed by outgrowths from last and next to

lm

Pterygostomian region. Triangular space on last thoracic somites. [ABW] /

ventral surface of carapace, on either side of Urogastrie lobe. Posteromedian lobe or subregion |

buccal cavity inBrachyura. Region at of gastric region, sometimes called genital

anterolateral comer of carapace in shrimps, region. [ABW] II

[ABW] II

Pterygostomian spine. Spine at anterolateral

(anteroventral) comer or border of carapace

in shrimps (Fig. 2). [ABW] II

Institutional Abbreviations I

AHF -- former Allan Hancock Foundation MMS -- Minerals Management Service I

Collection (now part of the Crustacea SBMNH -- Santa Barbara Museum of Natural

collection of the Natural History Museum of

Los Angeles County). History I

CAS -- California Academy of Sciences SIO -- Scripps Institution of Oceanography

I

CMM -- Cabrillo Marine Museum, San Pedro, USNM _ United States National Museum of 1

California Natural History, Smithsonian Institution. II

LACM -- Natural History Museum of Los 1

Angeles County I

56 |
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I Methods of Preservation

I should be initially fixed in buffered formalin, available "full Ideally, decapods Commercially strength"

formalin represents a 37% solution of formaldehyde; a 1:20 mixture of this solution with water will result in

a very strong fixative that is commonly referred to as "5% formaldehyde" (actually 5% formalin). Stronger

I solutions be used but those exceeding 10% often result in brittle specimens. Decapods tend to cast

may very

off appendages when subjected to severe shocks, such as immersion in formalin. This tendency often can be

moderated by euthanizing them, either by using initially a slightly weaker solution (1-2%) of formalin and

I gradually increasing the percent of formalin to 5-10%, or by chilling them in ice water prior to fixation. The latter technique has the added advantage that color of the integument is retained. Smaller specimens (e.g. less

than 100 mm carapace width for crabs and 100 mm carapace length for shrimp and lobsters) may be placed

I directly into the fixative solution. For larger specimens, it is best to use a syringe to inject this solution into the major body cavities, gill chambers, and even into the joints of the appendages to assure proper fixation, prior

to placing the entire specimen in a 10% solution. Several workers (e.g. see Lincoln and Sheals, 1979: 63) then

i recommend addition of small amounts of glycerol to the fixative at this time to keep specimens from becoming too brittle. After fixation for at least 24 hours (and up to 3 or 4 days for large specimens), they should be

thoroughly rinsed in fresh water and transferred to 70% ethyl-alcohol for long-term storage. Specimens with

I detailed depositedcollection in a majorinformation and reputable (inclinstitution uding datethat of coisllectio committed n, depth, to the habitat, long method term careofofcapture, marineetc.) invertebrates should be

(e.g., the Natural History Museum of Los Angeles County).

!

List of Families, Genera, and Species in the MMS Collections

!

Order Decapeda

I Dendrobranchiata Penaeidea

i Family Sicy Sicyonia oniidaeingentis Ortmann, (Burkenroad, 1898 1938)

Pleocyemata

I Caddea

Family Crangonidae Haworth, 1825

I NNeocrangon eocrangon zacae (Chace, 1937)

resima (Rathbun, 1902)

Neocrangon communis (Rathbun, 1899)

I Crangon alaskensis Lockington, 1877

Metacrangon spinosissima (Rathbun, 1902)

i Family Alpheidae Rafinesque, 1815 Alpheus belIimanus Lockington, 1877

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!
i ,

Family Hippolytidae Dana, 1852 I

Eualus lineatus Wicksten and Butler, 1983

Spirontocaris sica Rathbun, 1902 i

HI

Spirontocaris holmesi Holthuis, 1947

Family Palaemonidae Rafinesque, 1815 I

Pseudocoutierea I ele gans Holthuis, 1951

Thalassinidea

Family Axiidae Huxley, 1879 I

Acanthaxius spinuIicaudus (Rathbun, 1902)

Calocarides quinqueseriatus (Rathbun, 1902) i

i

Calocarides sp.

Anomura i

Family Paguridae Latreille, 1803 i

Paguristes uIreyi Schmitt, 1921

Parapagurodes laurentae McLaughlin and Haig, 1973 i

Family Galatheidae Samouelle, 1819

Munida quadrispina Benedict, 1902 I

Pleuroncodes planipes Stimpson, 1860

Family Lithodidae Samouelle, 1819 m

Paralomis muItispina (Benedict, 1894) I

Brachyura

Family Homolidae De Haan, 1839 I

I

Paromolafaxoni (Schmitt, 1921)

Family Xanthidae MacLeay, 1838 I

Lophopanopeus bellus diegensis (Rathbun, 1900) I

Family Parthenopidae Alcock, 1895

Heterocrypta occidentalis (Dana, 1854) i

Family Majidae Samouelle, 1819

Erileptus spinosus Rathbun, 1893 l

Family Pinnotheridae De Haan, 1833 g

Pinnixa occidentalis Rathbun, 1893

I

Pinnixa scamit Martin and Zmarzly, 1994 i

i

I

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I Key to Families and Species

I Williams (1984) to the 48 families of known from the east coast of the

presented a key decapods

United States. While some families in that key have no west coast representatives, the west coast families are

a subset of those 48. Further, the key is updated to correct family names, authors, and dates, and is thus

I preferable to the older key of Schmitt (1921) or the various popularized guides to west coast fauna (e.g. Kozloff, 1987; Allen, 1980; Jensen, 1995, etc.). Because that key was constructed to help in the identification

of east coast species, it was never intended to be used in other geographic regions, and reliance upon it for

I identification of west coast species without some modification of the couplets could present problems.

The following key is a modified subset of Williams' (1984) key to families, and includes only those

families andspecies mentioned in this report. This is of course very misleading, in that if a species belonging

I to a family or genus not among those our report study area, not

in is found for the first time in this it will be

possible to identify it using the following key. In fact, even species known to occur in this area will not always

be identifiable using such an artificial key.For example, a worker attempting to identify commercially important

I penaeid of the crab Cancer will have no luck. Even penaeids and species of

shrinlp or species a genus though

the family Cancridae are commonly encountered in California waters, and occur in our study area, none was

found in this particular survey. The reader is referred to more specific keys to the particular families or genera

I in question, such as that of Zrnarzly (1992) for the pinnotherid Pinnixa or that of Wicksten (1990) for

genus

the shrirnp family I-Iippolytidae, or to more encompassing and more general guides to decapods outside this

immediate study, when similar problems present themselves. Specific couplets for separating species within

I families are taken from earlier published sources, such as Word andCharwat (1976) and Butler (1980), or are newly created by us.

I

1A. General form shrimplike, usually compressed. Pleura of second abdominal segment never overlapping

I those ............................................................................................... of first segment. First 3 pairs of legs chelate. [Suborder DeFamily ndrobranchiata; Sicyoniidae: infraorder SicyoniaPenaeidea] ingentis

1B. General form shrimplike, lobsterlike, or crablike; if shrimplike then with pleura of second abdominal

I segment overlapping those of first segment. Of first three pairs of legs, third not chelate .................

.............................................................................................................. Suborder Pleocyemata ...... 2

I 2A. Form shrimplike, usually with body compressed ........................................ Infraorder Caridea ..... 3

2B. Form lobstedike or crablike ........................................................................................................... 15

I 3A. First pair of legs chelate ................................................................................................................. 10

i 3B. First pair of legs subchelate ........................................................................ Family Crangonidae .. 4

4. Gastric region of carapace depressed below general level of the carapace .... Metacrangon spinosissima

i 5. Gastric region of not depressed below general level of 6

carapace carapace

I 6A. Carapace with only one median gastric spine ..................................................... Crangon alaskensis 6B. Carapace with two median gastric spines ........................................................................................ 7

I
!

7A. The rostrum tilts up at an angle of 45 degrees; the tip of the rostrum has a small flag-like structure I

(not evident in specimens less than 20 mm TL) ................................................. Neocrangon resima

7B. The rostrum may be tilted upward, but not at a 45 degree angle; flag-like structure at tip absent. 8 I

8. All abdominal segments except fwst andsecond carinate on dorsal surface, with two carinae separated III

by median sulcus on sixth abdominal segment ............................................. Neocrangon communis II

9. Only sixth abdominal segment carinate on dorsal surface, with two carinae separated by median

sulcus ................................................................................................................... Neocrangon zacae I

10A. Carpus of second legs entire .................................. Family Palaemonidae: Pseudocoutierea elegans m

10B. Carpus of second legs subdivided into two or more joints; first pair of legs with chelae asymmetrical. 11

'IB

11A. Chelae of first legs very large and distinct, at least on one side; eyestalks short and covered by

carapace ................................................................................ Family Alpheidae:Alpheus bellimanus II

11B. Chelae of first legs not obviously large and distinct; eyestalks not covered by carapace ...................

.............................................................................................................. Family Hippolytidae ...... 12 I

12A. Carapace without supraorbital spines ....................................................................... Eualus lineatus

12B. Carapace with two or more supraorbital spines ............................................................................. 13 B

13. Carapace with three median spines, which blend together with the rosWalteeth .... Spirontocaris holmesi I

14. Carapace with two median spines, which are sfightly anterior to the midline ......... Spirontocaris sp.

m

15A. Either lobster-like or crab-like; abdomen extended, bent upon itself, or flexed beneath thorax; last I

thoracic sternite free; uropods present; carapace not fused with epistome; first, second, or first 2 pairs

of legs chelate or subchelate ................................... Infraorders Thalassinidea and Anomum ..... 16 I

15B. Crab-like, abdomen permanently flexed beneath carapace; last thoracic sternite fused with preceding;

uropods rarely present, never biramous; carapace fused with epistome; In'st pair of legs chelate or m

subchelate .......................................................................................... Infraorder Brachyura ....... 27 I

16A. Abdomen asymmetrical .................................................................................................................. 17

I 16B. Abdomen symmetrical ................................................................................................................... 21

17. Abdomen membranous and with uropods adapted for holding body in hollow objects ................. 19 i

18. Abdomen asymmetrical and sclerotized, flexed under body ... Family Lithodidae: Paralomis muItispina

19. Third maxillipeds approximated at base; antennae setose, moth-like ................................................ B

................................................................................................ Family Diogenldae: Paguristes ulreyi I

20. Third maxillipeds widely separated at base by sternum; antennae sparsely setose ............................

Family Paguridae: Parapagurodes Iaurentae I

i
!

I 21A. Body subcylindrical; first 2 pairs of legs chelate or subchelate; abdomen extended ..........................

............................................................................. Infraorder Thalassinidea, Family Axiidae ....... 22

I 2lB. Body depressed; only first legs chelate; abdomen bent under thorax .......... Family Galatheidae .. 25

i 22A. B. Eyes pigale,mented without....................................................................................... pigment ............................................................................................................ Acanthaxius spinulicaudus23

I 23. Distoventral spines on of second leg Calocarides

present cal'pus sp.

24. Distoventral spines absent on carpus of second leg ............................. Calocarides quinqueseriatus

I 25. Lateroinferior edges of carapace greatly swollen so that epimeral structures, sides of carapace, are

visible in dorsal view; pereiopods flattened for swimming ............................ Pleuroncodes planipes

I 26. Lateroinferior edges not greatly swollen; pereiopods more cylindrical for walking

.......................................................................................................................... Munida quadrispina

I 27A. Linea homolica present on carapace laterally; 5th pereiopods carded dorsally or subdorsally .........

................................................................................................. Family Homolidae: Paramolafaxoni

n 27B. Linea homolica absent; 5th pereiopods carried laterally, as other pereiopods ............................... 28

28A. Body conspicuously narrowed in front; rostrum distinct ............................................................... 29

n 28B. Body moderate to broad in front; rostrum absent .......................................................................... 31

I 29. Chelipeds with fingers straight; hooked hairs present ................ Family Majidae: Erileptus spinosus i

30. Chelipeds with fingers deflexed; hooked hairs absent .......................................................................

i ............................................................................. Family Parthenopidae: Heterocrypta occidentalis

31A. Carpus of third maxilliped articulating at or near anterointernal angle of meres; lateral margins of

i mouth frame parallel or divergent .................... Family Xanthidae: Lophopanopeus bellus diegensis 3 lB. Carpus of third maxilliped not articulating at or near anterointernal angle of merus; lateral margins

of mouth frame convergent, becoming ellipsoid ................................. Family Pirmotheridae ....... 32

I

32. Propodus of pereiopod 4 nearly as broad as long .............................................. Pinnixa occidentalis

I 33. Propodus of pereiopod 4 approximately 2.6 times longer than broad ........................ Pinnixa scamit

!

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I 61
!

Descriptions of Species I

Order Decapoda Latreille, 1803 n

Suborder Dendrobranchiata Bate, 1888

Infraorder Penaeidea de Haan, 1849 l

Family Sicyoniidae Ortman, 1898 l

I1

Sicyonia ingentis (Burkenroad, 1938) l

Figure 2.3

m

Eusicyonia ingentis Burkenroad, 1938: I

Sicyonia ingentis.--Word and Charwat, 1976:19.--Perez Farfante, 1988:25, Fig. 41A-C.--Jensen, 1995:79.

Material Examined. One small (carapace length = 4.4 mm including rostrum, total length = 12.4 1

mm) juvenile, USNM Phase I voucher, 073-BSR, 03-TX.

Description. Postrostral carina with 1 tooth posterior to level of hepatic spine. Dorsomedial carina i

/

on first abdominalsegment produced into tooth smaller or only slightly larger than posterior tooth oncarapace.

m

Petasma with distal projections of dorsolateral lobules divergent andproduced in short filament. Thelycum m

with posterior component of median plate beating paired short lateral bosses cut by transverse suture (adapted 1

from Perez Farfante, 1988, key). iN

Color in life not reported. II

Commonly referred to as the ridge backed prawn, rock shrimp, Pacific rock shrimp, camaron de I

piedra, andcacahuete.

Biology. Commercially harvested between Santa Barbara and Ventura, California (Frey, 1971). Attains I

a total length of 157 mm (males) to about 180 nun (females) (Perez Farfante, 1988).

Taxonomic Remarks. The single specimen examined was small and in poor condition, with carapace,

telson, and appendages slightly damaged. Assignment to S. ingentis is therefore tentative, although S. ingentis B

is the only species in the genus reported from as far north as California. Sicyonia ingentis is one of only 4

El

eastern Pacific members of this genus that are of commercial importance (Perez Farfante, 1988), the other 3

being S. brevirostris Stimpson, 1874, S. penicillata Lockington, 1879, andS. disdorsalis (Burkenroad, 1934). I

However, other species in the genus, not of commercial importance, are also present (e.g., see Perez Farfante m

and Boothe, 1981). Spination on the dorsal keel of the carapace of our specimen agrees most closely with

descriptions of S. ingentis, based on the key of Perez Farfante (1988). i

m Type Locality and Type Specimens. Station 127, Dredge 1, off east coast of Cedros Island, west

coast of Baja California, in 76-120 m. Type andCotypes, Cat. No. 361,025, Department of Tropical Research,

New York Zoological Society (6 males, 4 females). I

iN

Distribution. Isla Mafia Madre and throughout the Gulf of California (Perez Farfante, 1988), including

Cedros Island, Baja California (Burkertroad, 1938), north to Santa Barbara, California, including Santa m

Catalina Island (Carlisle, 1969; Frey, 1971; Word and Charwat, 1976) and to Monterey Bay (Perez Farfante, n

1988). From 5 to 183 m, occasionally to 293-307 m (Perez Farfante, 1988).

!
!

I

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I Figure 2.3. Sicyonia ingentis (Burkenroad, 1938): A, lateral view; B, dorsal view of petasma; C, thelycum;

scale for B = lmm, scale for C = 2mm (after Perez-Farfante, 1988).

ill

i Perez Farfante The noand rthernBoothrange(1981:424), record forbased this species, on a single Monterey female Barchived ay, Califoatrnia, the California was fast giAcademy ven by a offoomScience ote sin

(CAS 009269). The present record is the second report of S. ingentis north of Pt. Conception, California,

which was the northern range of the species given by Frey (1971 ). I

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I 63
I

Suborder Pleocyemata Burkenroad, 1963 I

Infraorder Caridea Dana, 1852

I

Family Crangonidae Haworth, 1825

Neocrangon zacae (Chace 1937) I

i

Figure 2.4

Crago zacae Chace, 1937:136, Fig. 9. i

Crangon communis.--Rathbun, 1904 in part. I

Crangon zacae.--Word and Charwat, 1976:93, Fig. 5-3 I

Neocrangonzacae.--KurisandCarlton, 1977:554. Wicksten, 1980:39. WickstenandHendrick:x, 1992:6.

II

Material Examined. USNM 073-BSR-02-TX Phase I primary voucher, 1 specimen; SBMNH 058-

BSS-0I -TX Phase I secondary voucher, 1 specimen; MMS Phase ITvoucher, MIVAloha, Cruisel-3, Station

R-4, 91 m, 3 specimens, largest - female, carapace length 7.2 mm including rostrum; MMS Phase I1voucher, i

MIVAloha, Cruise 1-1, Station PJ-15, 155 m, 1 specimen. II

Description. Gastric region not depressed below general outline of carapace. Two spines on dorsal

midline of carapace, posterior one in front of midpoint of carapace. A pair of lateral spines in line with 1

anterior dorsal spine. Rostrum moderately ascendent, narrowly rounded at tip, not reaching as far forward as II

eyes. Abdomen with no carina on first five somites, sixth somite with parallel carinae divided by median

sulcus. Blade of antennal scale exceeded by outer spine. Chelae of first legs more than three times as long as i

wide.

Carapace to base of rostrum about one-fourth as long as abdomen. Gastric region not depressed

below the general outline of the carapace. Two spines in dorsal midline, posterior one much longer, arising in 1

front of middle of carapace. A strong lateral spine present in line with anterior dorsal spine. Anterior margin ill

with two spines, one at lower orbital angle, other at antero-lateral angle of carapace.

illll

Rostrum ascends at slight angle which varies somewhat, but never becoming as great as 45 °, beating i

pronounced dorsal sulcus, tapering to blunt point, never reaching as far forward as eyes.

w

The abdomen without carina on first five somites. Posterior portions of first two segments slightly I

elevated; elevations preceded by broad, shallow, transverse sulci. Sixth segment about one and two-thirds |

times as long as fifth, bearing pair of dorsal longitudinal carinae separated by median sulcus and bounded on

outer side by shallower sulci. Telson longer than sixth somite; with two pairs of dorsolateral spinules, am

The eyes large and black. II

Second segment of antennular peduncle about twice as long as third. Flagella extending to ends of

hairs fringing antennal scale; latter about three and one-half times as long as wide, with outer spine easily 1

exceeding blade in length.

Third maxillipeds extending slightly beyond antennular flagella. First legs exceed antennal scale but i

not reaching as far as terminal fringe of the latter. A spine present at outer angle of distal end of merus. Palm I[

almost three and one-third times as long as wide, and dactyl closes very obliquely. Second pair of legs shorter t

than the first. Third pair extending well beyond third maxillipeds and fourth falls just short of latter.

l

!
I

I

,,•

I A

I f

A

| _

I /

I

I

Figare 2.4. Neocrangon zacae (Chace, 1937): A, Female holotype, lateral view; B, female holotype, dorsal

view of carapace; C, paratype, antennal scale; D, paratype, chela of first pereiopod (from Chace,

I 1937).

I

I

I 65
I

Color in life: Body semi-translucent, mottled finely with greenish-brown and scarlet on dorsal and I

lateral surfaces; ventral surface white. Antennae banded with scarlet and white; eye mottled greenish and

black; uropods and telson same color as body; remaining appendages translucent white (after Chace, 1937). II

Biology. Wicksten (1980) states that specimens from 105 stations in southern California and Baja g

California, Mexico in the collections of the Allan Hancock Foundation were taken at depths ranging from

shore to 572 m from bottoms ranging from mud to coralline algae. I

g

Taxonomic Remarks. Chace (1937) states that this species is closely alfied toN. communis (Rathbun),

N. resima (Rathbun), and N. abyssorum (Rathbun). It is distinguished from the latter by the smaller eyes,

longer palm of the first leg and lack of a cafina on the fifth abdominal somite. It differs fromN, resima in the I

differently shaped and less ascendent rostrum and longer palm.N, zacae most closely resembles N. communis I

but differs in the total absence of a carina on any but the sixth segment of the abdomen.

m

proposed a separation of the genus Crangon into the subgenera Crangon s. s. and U

Zarenkov (1965)

Neocrangon. Kuris and Carlton (1977) elevatedNeocrangonto generic status and although Wicksten (1980)

and Wicksten and Hendrickx (1992) refer to the above species as Neocrangon zacae, Butler (1980) did not

feel that Zarenkov's diagnosis forNeocrangon was valid. I

Dr. M. Wicksten, Texas A &M University, feels that this species and the following one (N. resima)

are synonymous and has a paper submitted to that effect (pers. comm.). Without having seen that manuscript, m

we still feel comfortable with the recognition of the two species. II

Specimens ofN. zacae seen by us, including the material from the Santa Barbara Channel, appear to

differ consistently from specimens of N. resima by the setation on the extremity of the rostrum, which is lit

restricted to a distal tuft in N. resima but is more widely scattered along both sides of the rostrum inN. zacae. II

This distinction holds true also for local specimens (D. Cadien, pers. comm.). However, the illustration of the

holotype ofN. zacae (reproduced here as Fig. 4B) shows setae of the local "resima" form. Further investigation •

of this character might be warranted. II

Type Locality and Type Specimens. Station 125, D- 1, East of Cedros Island, 28"13'N., 115=07'W.,

88 m, on a muddy bottom, 27 March, 1936. Holotype female, Cat. No. 36,1096, Department of Tropical •

Research, New York Zoological Society. II

Distribution. Monterey Bay, California, U.S.A., to Cedros Island, Baja California Norte, Mexico;

off Gonzaga Bay, Gulf of California, Mexico; Clarion Island; north of Gorgona Island, Columbia (Wicksten m

and Hendrickx, 1992) m

Neocrangon resima (Rathbun, 1902) I

Figure 2.5

II

Crangon resima Rathbun, 1902:889; 1904:124, Fig. 65. J

Crago resima.---Schmitt, 1921:96, Fig. 64.--Goodwin, 1952:171. I

Crangon resima.--Butler, 1980:95.--Word and Charwat, 1976:87 I

Neocrangon resima.--Kuris and Carlton, 1977:554.--Jensen, 1987b:402, Fig. 19.16.

1

Material Examined. M/VAloha Station PJ-1, May, 1987, 1 ovigerous female, carapace length 10.8 I

including rostrum.

Description. First five abdominal segments smooth. First and second segments with thickened band •

along posterior margin, in front of which a slight transverse depression. Fifth segment with an obscure median J

carina, with short depression on either side at anterior end. Sixth segment has two prominent carinae and

corresponding median depression. Telson with a slight median sulcus. Eyes of moderate size. Second segment I

|
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I of antennular peduncle three times as long as third. Rostrum long, narrow, pointed, advanced beyond eyes,

ascending at an angle of about 45°, slightly curved, and prolonged downward in thin, compressed plate, which

I appears show nospatulate evidence inof side it. Hands view.shorter The development than inN. communis of this plate , aboutis three depndent times onas long age; asspecimens wide, anterior 20 mmargin long

also more longitudinal than inN. communis. Antennal scale about four-fifths length of carapace, exclusive of

I rostrum; Biology. spine exceeding Accordingblade (after Schmitt, 1921). to Schmitt (1921) three specimens of Neocrangon resima were taken at D5788

in San Francisco Bay, and were associated withNeocrangon communis, Crangon alaskensis, andMetacrangon

I spinosissima. Taxonomic Remarks. See remarks for Neocrangon zacae

Type Locality and Type Specimens. Off San Diego, California, 124 fathoms (227 m) Albatross

I Station 2935 (Rathbun, 1904).

Distribution. Monterey Bay, California, to San Domingo Point, Baja California, 44-266 fathoms

i (80 to 486 m) (Rathbun, 1904).

!

| A

mm_m_ m _m_mmm m _ m m

m m

| •/!

I

-</

JJ B E _--'<

I

Figure 2.5. Neocrangon resima (Rathbun, 1902): A, lateral view of carapace; B, antenna]scale; C, anterodorsal

I view of carapace; D, chela (A from Wordand Charwat, 1976;B-D from Rathbun, 1904).

I 67
m

Neocrangon communis (Rathbun, 1899) I

Figure 2.6 I

Crangon communis Rathbun, 1899:556:1904:112 (key), 123, Fig. 64.---de Man 1920.--Word and Charwat

1976:77.--Butler 1980:110, Plate 5D. i1

Crago communis.---SchmJtt, 1921:95, Fig. 63.---Goodwin, 1952:171. g

Sclerocrangon communis.--Derjugin and Kobjakowa, 1935:84.--Vinogradov, 1950:256.

Crangon (Neocrangon ) communis.--Zarenko v, 1965. I

m

Neocrangon communis.--Kuris and Carlton, 1977:554.--Jensen, 1987b:402, Fig. 19.17.

Material Examined. USNM 018-BSS-01-TX Phase I primary voucher, 1 specimen, carapace length 1

18.5mm including rostrum; SBMNH 014-BSS-01-TX Phase I secondary voucher, 1 specimen. m

Description. Body slender, depressed. Shell thin, pubescence on anterior part of carapace, and in m

abdominal sulci. Rostrum short, about 0.2 carapace length, median sulcus, ascending, tip rounded, i

Carapace spines: 2 median in anterior half of carapace, anterior spine slightly smaller; median dorsal

carina, ahead of anterior spine extending to posterior half of carapace; antennal moderate, with supporting 1

carina; branchiostegal strong, with carina; pterygostomian weak; hepatic strong; lateral margin of rostrum II

extending posteriorly over orbital fissure as ridge above and beyond hepatic spine. Eye moderately large;

comea well developed; weak tubercle. •

Antennule: peduncle short, second segment over twice length of first, latter with dorsal distal lobe; II

stylocerite short; inner flagellum longer than outer, and extending beyond antenna] scale.

Antenna: scale shorter than telson, spine exceeding lamella; basicerite, upper lateral lobe, lower i

oblique spine; peduncle long; flagellum about as long as body. II

Third maxilliped: long, slender; distal part of antepenultimate segment twisted; exopod long, with lash. m

Pereiopod 1 as long as third maxilliped, stout except for ischium and proximal portion of merus. 1

Merus with strong distal spine. Carpus with 2 weak distal spines. Propodus widens distally, fixed finger long

and stout. Dactylus obliquely transverse when flexed. Pereiopod 2 shorter than 1,slender, chelate. Pereiopod mid

3 longer than 2 and as slender. Dactylus slender, about 0.5 times propodus length. Pereiopod 4 shorter than 3, 1

stouter. Dactylus slender, about 0.65 times propodus length. Pereiopod 5 shorter than 4, slightly stouter.

Dactylus slender, about 0.8 times propodus length. I

Abdomen: all sulci filled with sparse short pubescence easily rubbed off. First and second somites, i

each with long transverse sulcus. Third to fifth somites each with wide median dorsal carina; third with an

anterior transverse sulcus through carina and a wide vertical sulcus below carina. Fourth somite with 2 l

transverse sulci, one at each end of carina. Fifth somite with an anterior transverse sulcus through carina, and II

a wide sulcus or depression on each side of carina. Pleuron of fourth somite with a moderate posteroventral

spine. Fifth pleuron with a moderate posterolateral spine. Sixth pleuron with 2 median dorsal carinae, a m

sulcus on outer side of each carina, and a weak posteroventral spine. Telson moderately wide, tapering to II

rounded tip. Median sulcus of telson with 2 pairs of dorsolateral spines. Inner uropod longer than outer with

neither reaching end of telson. i

Color in life: Medium gray background over most of body. More or lessuniform brown over branchial _l

region of carapace, ovedaid with pale yellow spots, and fine brown to charcoal dots over rest of carapace.

Abdomen has brown spots and blotches, light yellow spots, more intense along midlateral line of first to iN

fourth, and sixth somites; ventral margins of all somites bordered by magenta mottling. Magenta on pteopods, I

and chela of pereiopod I, which also has brown spots. Other pereiopods with magenta to rust patches; latter

color repeated distally on antennal flagellum. Brown spots on telson and uropods; outer margin of outer 1

uropod magenta (from Butler, 1980) I
I

I

I

.._+;:_,._i _" 7_

I

'

I

Figure 2.6. Neocrangon communis (Rathbun 1899): A, dorsal and lateral views (from Butler 1980); B, antennal

I scale; C, anterolateral view of carapace; D, anterodorsal view of carapace; E, chela (BI

I _9
!

Biology. Butler (1980) states that "this species may rank as the most common local shrimp [along the I

Pacific coast of Canada]. It occurs coastwide on mud bottoms from the shallow sublittoral to the lower

continental slope. On grounds where Pandalopsis dispar, Pandalus jordani, and P. borealis are trawled I

commercially, N. communis is the most abundant associated shrimp species. In addition to capture by bottom II

trawls, it also has been collected by midwater trawls, At one locality off Active Pass at maximum depths of

128-161 m, specimens were caught at intermediate depths from 20-67 m. 1

Ovigerous females, 9.1-13.9 mm carapace length, occurred in January, February, May, and June to m

September.

Known parasites are the branchial isopod,Arge/a pugettensis and the rhizocephalalan, Mycetomorpha i

vancouverensis." I

Taxonomic Remarks. Rathbun(1904) notedsome variationin rostralmorphology caused bybreakage n

and subsequent regrowth of the rostrum. See remarks for Neocrangon zacae, n

Type Locality and Type Specimens. Off Pribilof Islands, Bearing Sea, 94 m Albatross Station

3441. Holotype female length 64 mm, carapace length 16 mm, antennal scale length 10.5 mm. n

Distribution. Bering Sea to San Diego, California (including Puget Sound and Strait of Juan de I

Fuca), 20 - 309 fathoms (37 to 565 m) (Rathbun, 1904).

1

Crangon alaskensis Lockington, 1877 I

Figure 2.7 I

Crangon alaskensis Lockington, 1877:I70 in part.--Rathbun, 1904:112 (key), 114, Figs. 52, 53.--Butler,

1980:108, Plate 3E in part.--Kuris and Carlton, 1977:553.--Jensen, 1995:40, Plate 59. II

Crangon nigricauda.--Lockington, 1878:229, 465-480, in part.

Crangon crangon affinis.--Ortman, 1895:173 in part. In

Crangon vulgaris.wKingsley, 1899:709 in part. U

Crangon alaskensis elongataRathbun, 1902:888; 1904:112 (key), 115, Fig. 54.---de Man, 1920

Crago alaskensis elongata.--Schmitt, 1921:88, Fig. 58 I

m

Crago alaskensis.---Carlisle, 1969:237.

Material Examined. 1 specimen, MV Aloha Station R-8, Oct., 1988. n

Description. Our specimen more closely resembles the "elongata" form previously treated as a

subspecies by various authors. This form differs from the typical form as follows: longer and narrower

rostrum; outer antennular flagellum not reaching to end of antennal scale; longer antennal scale, as long as 1

carapace; faint dorsal median carina on fourth somite; tip of telson more acute.

U

Description ofC. alaskensis typical form: Body slender,depressed. Shellthin surface smooth. Rostrum •

short, about 0.2 times carapace length, with median sulcus, ascending, tip rounded. II

Carapace spines: 1 median in anterior third of carapace; antennal moderate; hepatic moderate, with

short oblique supporting carina, anterior to median; branchiostegal strong with supporting carina, almost to •

end of basicerite of antenna; pterygostomian moderate; lateral margin of rostrum extending posteriorly over

orbital fissure as ridge almost to center of carapace. Band around ventral margin of carapace, widening to

lobe adjacent to bases of pereiopods 1 and 2. •

II

I

70 !
!

I Eye moderately large, cornea well developed, weak tubercle.

Antennule: peduncle moderately long, third segment subequal to second; stylocerite short; outer

I flagellum extending to about end of antennal scale, inner longer. Antenna: scale 0.8-1.0 times carapace length, as tong as or slightly shorter than telson, tip of lamella

rounded, spine exceeding lamina somewhat; basicerite, 2 lateral spines; peduncle moderately long; flagellum

I shorter than body.

Third maxilliped: moderately long and slender, short exopod with lash.

I distal spine. Pereiopod Propodus I longer oblong, thanmoderately third maxitliped, long, 2.6-3.0 stout. Merus times average with innerwidth, spinefixed in proximal finger moderate, half, and aslender. strong

Dactylus transverse when flexed obliquely. Pereiopod 2 about as long as 1, slender and chelate. Pereiopod 3

i longer dactylusthan 2, slender, about 0.55 times length of propodus. Pereiopod 4 longer than 3, slightly stouter; slender, about 0.8 times propodus length.

Abdomen: ventral margins of pleura of first and second somites concave. Fifth somite with faint

I dorsal moderatemedian posteroventral carina andspine, short transverse and ventralsusulcus. lcus atTelson anteriornarrow, end. Sixth tapering somiteto acute with faint tip, dorsal mediandistally dorsalflattened sulcus,

with two lateral spines evenly spaced with respect to tip of telson. Inner uropod narrower, as long as or longer

I than outer, Color former reaching tip of telson (after Butler, 1980). in life: Light brownish gray background over almost entire shrimp, lighter on most appendages;

closely spaced dark gray and brown spots over body, pleopods, pereiopods, and all parts of antenna. Scattered

I spots of pale yellow, white, and light brown on carapace and abdomen. Pale yellow spots on antennal peduncle, proximal segments of pereiopods 2, 4, and 5, basipodites of pleopods. With a dense coveting of fine black dots

on posterior midtine of carapace, around posterior margin of sixth somite, and on telson and uropods. Tip of

I inner uropod orange (Butler, 1980, Plate 3E; Jensen, 1995, Plate 59). Biology. Very common along the west coast of the United States (Rathbun, 1904:115).

Taxonomic Remarks. Butler (1980) chose not to recognize the subdivision into two subspecies first

I suggested by (1902, 1904) specimens Canadian waters, instead attributing the slight

Rathbun for from

morphological differences between those "forms" to intraspecific variability. Rathbun (1904) discussed rostral

variation in the "elongata" form. The specimen illustrated here is from the Los Angeles Harbor area and

I adheres most to the "elongata" form.

closely

Type Locality and Type Specimens. For C. a. elongata, off Santa Barbara, California, 29 fathoms

i (53 m), Albatross DistributionStations . East part 2970, of Bering 2971 (Rathbun, Sea and alo1904:115). ng the Aleutian Islands, south to off Wdmington, California

(Rathbun, 1904, as C. alaskensis elongata). 3.25 to 41 fathoms (6 to 75 m) (Rathbun, 1904).

!

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I

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I

Figure 2.7. Crangon alaskensis I_x_kington, 1877: A, lateral view of ovigerous female; B, amemlateral view of

carapace; C, dorsal view af carapace; D, antennal scale; E, third maxiltiped; F, chela. Original

illustration from specimen taken from Los Angeles Harbor. I
!

I Metacrangon spinosissima (Rathbun, 1902)

I Figure 2.8

Crangon spinosissima Rathbun, 1902:891; 1904:112 (key), 130, Fig. 70.--deMan, 1920:217.--Wicksten,

1977:282.--Word and Charwat, 1976:89, Fig. 5-2.

I Crago spinisissima Schmitt, 1921:100, Fig. 69.--Goodwin, 1952:171.--Butler, 1950:50.

Metacrangon spinosissima Zarenkov, 1965:298.--Jensen, 1987b:400.--Butler, 1980:115, Plate lB.

I Material Examined. USNM 038-BSS-01-TX Phase I primary voucher, 1specimen, carapace length

6.2 mm including rostrum.

I pubescent. DescRostrum ription.short, Body0.15-0.2 stout, carapace depressed.length, Shellmedian thick, sulcus, mainlyascending, mgose, depresse tip acuted orportio rounded, n of Ccararapace a a

spines: 2 median, anterior larger, just posterior to rostrum, posterior inposterior half of carapace, the 2 joined

I bycarina; a median antennal carina; strong,submedian, ascending;about posorbital center moderate, of carapacconfluent e, with suwith pporting antennal; crinweak a; hepatic spine with belowsupporting antennal;

branchiostegal strong, with supporting carina; pterostomian weak; f'me carina extending from slight fissure

I bel wideni ow nanten g asnpronounced al spine posteriorly lobe adjacent acrosstodepression base of pereiopod in carapace. 2 or 3.Band around ventral margin of carapace,

Eyes small, corneas well developed, stalks taper distally, deflected obliquely.

I Antennule: Peduncle long, second segment longer than third, fast and second each with distal lateral spine; stylocerite moderately long, broad, pointed laterally; inner flagellum longer than outer, both exceed

antennal scale.

I Antenna: Scale shorter than telson, lamella exceeds spine; basicerite, 2 blunt lateral spines; peduncle

long; flagellum longer than body, with plumose setae.

Third maxiUiped: Moderately long, slender, except antepenultimate segment, setiferous; exopod long,

I with lash.

Pereiopod 1 longer than third maxilliped, stout. Merus with ventral longitudinal carina. Propodus

I widens flexed. Pereiopod distally, len2gthlonger 3.6-4.0thantimes 1,slender averageandwichelate. dth, fixedPereiopod finger mo3dlonger erate. than Dacty2,lusas obliq slender; uelydactylus l_ansversslender when

and about 0.44 times propodus length. Pereiopod 4 shorter than 3, moderately stout and setiferous. Dactylus

I fldactylus attened,flattened, about 0.9about times0.55propo timesduspropodus length. Pereiop length.od 5 shorter than 4, moderately stout and setiferous;

Abdomen: Pleura of first 3 somites each with 2 ventral spines. Fourth to sixth somite each with

I moderate of first andposteroventral third somites accommodate spine. Sixth somite pleuronwithofstrong second.spineFirstattodorsoposterior third somites margin. each withDepressions faint medianin pleura dorsal

carina. Dorsal anterior margin of first deeply concave. Second with dorsal transverse furrow, adjacent anterior

I mDorsal argin produced, median carina formingon sligh posterior t hoodhalf overoffurrow. fourth,Third lateralwitcarina h dorsalintransverse line withsul articular cus throuknobs. gh mediFifth an earina. with

transverse sulcus through anterior end of faint median dorsal carina, lateral carina extending to middle of

I somite obliquelyfromfromarticular articular knob. knobSixthtoward with 2dorsal prominent margin. dorsPosteroventral al carinae, shorter regions thanosomite, f sixth flared lateralstrongly carina extegivinding ng a

wing-like appearance in dorsal view. Telson narrow, tapering to acute tip. Median dorsal sulcus of telson with

I 2 pairs of widely spaced dorsolateral spines. Inner uropods longer than outer, and exceeding tip of telson.

!
I

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I

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__ - I_,,,_ --_ -__ I

__,_._, .:::_.... L',.:_._._ ...... •

I

B

'

I

I

_ I

o I

I

Figure 2.8. Metacrangon spinosissima (Rathbun, 1902): A, dorsal and lateral views (from Butler, 1980); B,

dorsal view of carapace (from Rathbun 1904); C, lateral view (from Rathbun, 1904); D, dorsal view I

of carapace; E, anterolateral view (D and E from Word and Charwat, 1976).

74 I
!

I Color in life: Background of body and appendages buff. Brownish mottling blending with background

on posterior branchial region of carapace. Other brown patches occur at bases of median spines and bordering

I found anterolat doresral allymargin on first. abdominal Light brownsomite patchandes lateral on pleura surfaces of firsoftfour to tthird h to sixth abdosomites. minal soThis mites. borders Darkerconspicuous areas are

light swath and rectangular patch near ventral margin of second pleuron. Light brown transverse bars are on

I telson rust. Latter and inner appendages, uropods. Pleopods anterolateml are light marginredoftocarapace, pink. Thirdandmaxill anterior iped and appendages, pereiopodswith1, 4,light and 5rust are setae light

(Butler, 1980, Plate 1B).

I allows itBtoiology assume . Butler a "defensive (1980:116) cataleptic notes that position the shape as inoArgis f firstlevior and secandondLebbeus abdominal schrencki somite.'" ofHethis alsoshrimp notes

that it is common offthe southwest coast of Vancouver Island and in the southern Strait of Georgia, is often

I trawled and August overinmud that bottoms region. (depths of 97-157 m), and has been found ovigerous in February, March, April,

Taxonomic Remarks. Distinguished from other species in the genus by having 1 or 2 ventral spines

I onmargin each of the first to fifth pleura and a transverse furrow on the second abdominal somite, with the adjacent produced as a hood over this furrow (Butler, 1980: 116).

Type Locality and Type Specimens. Off Point Arena, California, 51 fathoms (93 m) (Albatross

I Station 3351) (Rathbun, 1904).

Distribution. From Nootka Sound, Alaska (Butler, 1980), southward to Isla San Martin, Baja

i California (Wicksten, 1977), at a depth of 51 to 96 fathoms (93 to 176 m) (Rathbun 1904).

i Family Alpheidae Rafinesque, 1815

Alpheus beUimanus Loekington, 1877

I Figure 2.9

Alpheus bellimanus Lockington, 1877:34; 1878:470.--Kingsley, 1878a: 199; 1878b:59; 1883:111; 1899:716.--

I Bate, Rathbun, 1888:54.---Sharp, 1893:l12.---Coutiere, 1899:28.--Holmes, 1900:184, Plate 2. Fig. 41.-- 1904:108.--Word and Charwat, 1976:42.--Luke, 1977:4.--Brusca, 1980:252.--Wicksten,

1983:41; 1984:189.--Kim and Abele, 1988:13, Fig. 5.--Jensen, 1995:44, Plate 70.

I Crangon bellimanus.---Schnfitt, 1921:75, Fig.51.---Chace, 1937:188.

Material Examined. USNM 006-BRS-02-TX Phase I primary voucher, 3 specimens, largest - male,

I carapace length 9.3 mm; SBMNH 004-BRA-04-TX Phase I secondary voucher, 2 specimens.

Description. Major chela with sinuous inferior margin (distal to inferior transverse groove) sinuous

because of deep transverse groove; movable finger of major chela shallowly arched along proximal two-thirds

I of superior margin, and then regularly arched distally, tip stout, often bulbous; appendix masculina subequal

to or slightly overreaching distal end of appendix interna. Rostrum long, far overreaching middle of visible

part of first antennular segment; movable f'mger of minor chela laminate in male; merus of major first pereiopod

I bearing 6 to 10 small movable spines on inferior inner margin and a strong, acute immovable spine at the distal end, ischium likewise armed with 3to 4 spines on inner margin. Palm of major chela with deep transverse

groove on superior margin and with deep, distinct superior and inferior grooves on outer face. Stylocerite

I narrowing to a long sharp point, almost reaching to distal margin of first antennular segment. Basicerite with sharp lateral spine; spine broad at base. Ocular hood with sharp tooth directed slightly inward and downward

(modified from key and description of Kim and Abele 1988).

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I

I

' I

I

I

J I

I

I

Figure 2.9. Alpheus beIlimanus I_x_kington, 1877: male, carapace length 5.2 mm; A, anterolateral view; B,

anterodorsal view; C, carina below first antennular segment; D, right third maxilliped; E, outer i

face of major first pereiopod; F, inner face of same; G, outer face of minor first pereiopod; H, inner •

face of same; I, left second pereiopod; J, right third pereiopod; K, abdomen; L, telson and uropods; i

M, outer face of minor chela from first pereiopod of ovigerous female (carapace length 6.5mm); N,

right second pleopod (female, carapace length 4.6 mm)(from Kim and Abele, 1988). •

II

76 m
I

I Color in life: Body uniformly blood red changing to olive green after capture (Jensen, 1995:44, Plate

70), orange at tip of tail fan; tips of chelae and branchial regions orange; white mark on palm of major chela

I (Wicksten, 1983:42). Biology. Found in the lowest intertidal zones to 300 m, usually dredged on sand, rocks, soft mud, fine

gravel or among corallines (Kim and Abele, 1988).

I Taxonomic Remarks. Belongs to the Macrocheles (= Megacheles) of species of this

group very

diverse genus (Kim and Abele, 1988).

Type Locality and Type Specimens. San Diego, California (Kim and Abele, 1988). Types probably

I lost in the San Francisco fire of 1906 (M. Wicksten, pers. comm.)

Distribution. Monterey, California, south along outer coast of Baja California, Mexico; southern

i Gulf of Catiforrtia, Islas Socorro and Clarion; western Mexico; Panama; Chile; Colombia; Galapagos Islands (Kim and Abele, 1988).

i I Family Hippolytidae Dana, 1852

i Eualus lineatus Wieksten and Butler, 1983 Figure 2.10

I Spirontocaris (in part).

herdmani.--Rathbun, 1904: 100

Eualus herdmani.--Holthuis, 1947:11 inpart.--Hobson and Chess, 1974:579.--Word and Charwat, 1976:103

i Eualus lineatus in part.--But Wicksten ler, 198 and0:197- Butler, 1981983:1, (in part). Figs 1-2.--Jensen, 1987a:399; 1995:45.

Material Examined. USNM 016-BRA-01-TX Phase I primary voucher, 3 specimens; SBMNH

I 016-BRS-04-TX Phase I secondary voucher, 1 specimen.

Description. Rostrum short, reaching second segment of antennular peduncle, straight, with 3-6

I dorsal teeth and 1-3 ventral teeth, all of dorsal margin toothed. Carapace smooth, dorsum straight, small suborbital spine, moderate antennal spine with supporting

cafina, moderate pterygostomial spine.

I Abdomen smooth. Pleura of segments 1-3 rounded, pleura of fourth segment with weak posterolateral spine, fifth with strong posterolateral spine. Median dorsal margin of third segment strongly produced posteriorly.

TeIson with 3 pairs dorsolateral spines.

I Eyes moderately large, darkly pigmented.

Stylocerite reaching end of first segment of antennular peduncle. First segment with 3 moderate

i dorsal second,spines. with 2Sec spines. ond segInner mentflagellum about 0.5more timesthanlength2 times of firslength t, withof2outer. strong spines. Third segment shorter than

Basis of second antenna with sharp ventral spine, large blunt upper lobe. Scaphocerite oblong, shorter

I than caraMandible pace, lamella slightly longer than spine. Flagellum slightly exceeding body length. with slender incisor process, ending in small teeth. Molar process with spinules. Twojointed palp present. First maxilla with lower endite slender, upper broad; palp faintly bilobed, Second maxilla

I with with lower endite reduced, upper endite larger,bilobed; palp and scaphognathite well developed. First maxilliped exopod and 2-jointed palp; epipod bilobed, caridean lobe very small; endites of coxa and basis separated

by notch. Second maxilliped with exopod, podobranch, and epipod. Third maxilliped with exopod and epipod,

I exceeding antennular peduncle, stout and setose.

I 77
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First pereiopod shorter than third maxilliped, stout, with epipod. Fingers of chela less than 0.5 times I

length of palm. Carpus slightly shorter than palm. Merus about 2 times length of carpus. Ischium less than 0.5

times merus. Second pereiopod chelate, longer than first pereiopod, with epipod. Fingers shorter than length •

of palm. Carpus with 7 segments, the third and fourth the longest. Merus about 0.6 times length of carpus, I

slightly longer than ischium. Third pereiopod slightly longer than second, with epipod. Dactyl stout, about 0.2

times length of propodus, with 5 spines. Propodus with 16-19 spinules in 2-3 rows. Carpus about 0.4 times •

length of propodus. Merus about same length as propodus, with 3 spines. Ischium less than 0.5 times length II

of merus. Fourth pereiopod about as long as third. Dactyl stout, similar to that of third pereiopod, Propodus

with 15 spinules in single row. Merus with 2-3 spines. Fifth pereiopod as long as fourth. Dactyl about 0.2 •

times length of propodus, with 4-5 small spines. Propodus with 14-22 spinules in 1 or 3 rows. Merus with 0- l

1 spine. No epipods on pereiopods 3-5.

Second pleopod with appendix interna. Appendix masculina little more than half length of appendix •

interna, stout, apex truncated, with 8 long spinules. Uropods reaching end of telson, with long tooth on outer II

margin (from Wicksten and Buffer, 1983).

Color in life: Red diagonal lines on carapace and fh'st and second abdominal segments; red spots and i

blotches on third to sixth abdominal segments, telson, uropods, and protopodites of pleopods; smaller red

spots on all anterior appendages, including eye stalk and pereiopods (from Wicksten and Butler, 1983; Buffer,

1980; color Plate 1C, as Eualus herdmani), i

Biology. This species has been found in rocky rubble below 18 m (Jensen, 1995). Most specimens

have been taken by trawl. One specimen was collected by a SCUBA diver on the wreck of the Olympic, off

Long Beach, California. A first stage larva of the species has been hatched in the laboratory (Butler, 1980). i

The species is eaten by rockfishes of the genus Sebastes off Santa Catalina Island, California (Hobson and

Chess, 1974).

to be parasitized by Bopyroides hippolytes (Butler, 1980). i

Some individuals have been found

Taxonomic Remarks. Eualus lineatus is related to E. avinus (Rathbun), E. pusiolus (Kroyer), and

E. berkeIeyorum Butler in having epipods on the fh'st 3 pereiopods. InEavinus the rostrum is arched over the •

eye, and bears 12-14 upper teeth and 1-3 ventral teeth. The dactyl of the third pereiopod is slender and simple. II

Eualus pusiolus has a straight rostrum shorter than the second segment of the antennular peduncle, with 2-5

upper teeth and at most 1 lower tooth. The dactyl of the third pereiopod is stout. Eualus berkeleyorum has a •

straight rostrum with 8-11 dorsal teeth and 2-5 ventral teeth. The dactyls of its third pereiopods are slender |

and simple. It does not have a ventral spine on the pleura of the fourth abdominal segment (Wicksten and

Butler 1983).

Type Locality and Type Specimens. 1.5 mi. SW of Gulf Island, off Santa Cruz Island, California i

(33°56'00"N, 119°50'55"W), 89 m, mud and sponge bottom, 8 Nov 1941, VeIero III Sta 1435-41. Holotype

female total length 21.0mm; AHF type number 4129. •

D | istribution. Sitka, Alaska to Puget Sound (Butler, 1980);Santa Monica Bay, Palos Verde Peninsula,

Santa Catalina Island, and Point Loma California (Word and Charwat, 1976).

i

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I Figure 2.10, Eualus lineatus Wicksten and Butler, 1983: A, dorsal and lateral views; B, anterolateral view of

female (carapace length 3.1 ram); C, left third pereiopod of same; D, dactyl of left third pereiopod

of same; E, appendix mascutina of male (carapace length 4.1 mm)(from Wicksten and Butler,

I 1983).

I

i

II 79
I

II

Spirontocaris sica Rathbun, 1902 •

Figure 2.11 i

I

Spirontocaris sica Rathbun, 1902:894.--Rathbun, 1904:69, Fig. 25.---Schmitt, 1921:55, Fig. 32.--Holthuis,

1947:37.--Word and Charwat, 1976:154.--Butler, 1980:167, Plate 7D.--Wicksten, 1990:590.

Material Examined. USNM 055-BSS-01-TX Phase I primary voucher, 1 specimen, carapace length •

including rostrum 10.5 mm, rostrum length 4.7 mm. i

Description. Body slender and somewhat compressed. Rostrum moderately long, 0.9-1.2 times

am

carapace length, constricted at base, median section as broad blade, lower limb broader and tapering sharply,

distal portion as long ascending styliform process, tip acute. I

Carapace spines: 2 supraorbitals, widely space, with upper spine stronger; suborbital strong and I

rounded; antennal moderate with carina; pterigostomian weak or obsolescent.

Eye: Large and cornea well developed, i

Antennule: Peduncle long, third segment half length of second, dorsal distal spine present on each;

stylocerite moderately long and knifelike; inner flagellum longer than rostrum, outer extending slightly beyond

antennal scale, i

Antenna: Scale slightly shorter than telson, lamella exceeds spine somewhat; basicerite with two

lateral lobes; peduncle moderately long; flagellum longer than body. 1

Third Maxillaped: Moderately long, moderately stout with an epipod and an exopod. I

Pereiopod 1 shorter than third maxillaped and a little stouter, chelate, epipod present.

Pereiopod 2, chelate, slender and longer than 1, carpus with 7 segments. Pereiopod 3 longer and stouter than

2, merus with 5-9 spines, propodus with 4 spinules and a distal fringe of setae, dactylus slender. Pereiopod 4

slightly longer than 3, merus with 6-8 spines, propodus with 4-7 spinules, setae as in 3. Pereiopod 5 shorter

than 4, merus with 4-7 spines, propodus with 1-8 spinules, long setal band around distal end. 1

Abdomen: Ventral margin of pleuron of second somite broadly convex; posterodistal margins of

fourth and fifth deeply recessed at articular knobs; fifth with week posterolateral spine; sixth about as long as

telson, slender with moderate posteroventral spine; telson narrow, tapering to acute tip, 4 pairs of dorsolateral i

spines; uropods about same length as telson, extending to tip (after Butler, 1980). i

Color in life: Background of body milkish white to dull yellow. Carapace with red blotches, large m

specimens with red band along ventral margin, large red area on branchial region. Most have a pinkish dorsal 1

patch. Rostrum with a few red dots on the midrib. Antennule, basicerite and base of antennal scale pinkish.

lid

Lateral surface of ftrst five abdominal somites with fine red spots. Telson and uropods pinkish. Pereiopods

deep red (Butler, 1980: Plate 719). i

Biology. Buffer (1980) states that in almost all cases along the western coast of Canada this species

has been found in coastal inlets, and that as an adult may be pelagic rather than demersal. It is parasitized by i

the branchial isopod Bopyroides hippolytes. 1

Taxonomic Remarks. This specimen was sent to us with a tentative identification asS. dalli Rathbun,

1902, a species whose southern most range is Sitka, Alaska except for a single record from near Los Angeles •

(Word and Charwat 1976). We felt comfortable with this identification based upon rostral characteristics and 1

the key of Word and Charwat (1976). However, upon review of this manuscript both D. Cadien and M.

Wicksten suggested that this specimen probably is not S. dalli. In fact if rostra/characteristics are ignored this •

specimen seems to be S. sica, Rathbun, 1902 based on the keys of Schmitt (1921), Holthuis (1947), Buffer i

(1981 ), and Wicksten (1990). This specimen differs from typical specimens in that the rostrum is lacking the

long ascending styliform process and instead terminates in a short triangular point. Defining characteristics I

80 l
I Figure 2.11. Spirontocaris sica Rathbun, 1902: A, dorsal and lateral views; B, anterolateral view (A, from Butler, 1980; B, from Word and Charwat, 1976).

I

that place our specimen with S. sica are as follows: Carapace with 2 supraorbital spines; pleurae of abdominal

segments 1 and 2 rounded; epipod on first pereiopod only (after Holthuis, 1947). Spirontocaris holmesi (see

I below), on the other hand, has epipods on pereiopods.

both the first and second

Type Locality and Type Specimen. Holotype female: total length 58 mm; carapace length 24.8 mm

including rostrum; rostral length 12.8ram. Type locality: Santa Barbara Channel, 265 fins (485 m) Albatross

I station 3200.

Distribution. Restoration Bay, British Columbia to between San Benito Islands and Cedros Island,

I Baja California; 88-849 m (Wicksten, 1990)

I, 81
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Spirontocaris holmesi Holthuis, 1947 I

Figure 2.12 n

1

Spirontocaris bispinosa Holmes, I900:128.--Rathbun 1904:219, Fig. 23.---Schmitt, 1921:54, Fig. 30.--

Goodwin, 1952:170. n

HippoIyte bispinosus.--Williamson 1915. 1

Spirontocaris holmesi Holthuis, 1947:129.--Word and Charwat, 1976:149.--Butler, 1980:159 (key), 165,

Plate 6E.--Jensen, 1987a:399.--Wicksten, 1990:590. 1

1

Material Examined. USNM 007-BSS-01-TX Phase I primary voucher, 1 specimen, carapace length

including rostrum 12.4 nun, rostrum length 6.0 mm; SBMNH 103-BSS-01-TX Phase I secondary voucher, 1 •

specimen. 1

Description. Body slender and compressed. Shell thin and smooth. Rostrum moderately long, 1.0-

1.5 times carapace length, constricted at base. Median section of rostrum broad blade with lower limb broader •

and rounded. Distal portion slender, ascending, styliform process with a solitary distal ventral spine.

Carapace spines: 2 supraorbitals widely separated, upper spine stronger; suborbital moderate, pointed;

antennal moderate; pterygostomian weak or obsolescent. 1

I

Eyes large, corneas well developed.

Antennule: peduncle moderately long, third segment shorter than second, each with dorsal distal •

spine; stylocerite moderately long, inner flagellum shorter than rostrum, outer extending beyond antennal I

scale slightly.

Antenna: scale shorter than telson, lamella exceeds spine somewhat; basicerite with 2 weak lateral •

spines; peduncle long; flagellum longer than body.

Third maxilliped: moderately long, stout, with an exopod and an epipod.

Pereiopod 1 shorter than third maxilliped, a little stouter, chelate, with epipod. Pereiopod 2 longer n

than 1, slender and chelate, carpus with 7 segments, epipod present. Pereiopod 3 longer than 2 and a little E

stouter, merus with 6 or 7 spines, propodus with 4-8 spinules, distal fringe of setae present. Dactylus slender, III

about 0.6 times propodus length, tip acute. Pereiopod 4 about as long as 3 and as slender, merus with 5-8 II

spines, propodus with about 4 spinules, distal fringe of setae present. Dactylus slender, about 0.6 times II

propodus length, tip acute. Pereiopod 5 shorter than 4 and as slender. Merus with 4 or 5 spines, propodus with

3-6 spinules, distal fringe of setae present. Dactylus slender, about 0.55 times propodus length, tip acute. 1

Ill

Abdomen: venla'at margin of pleuron of second somite mainly straight or slightly concave. Fifth

somite posterolateral margin widely recessed at articular knob, and with moderate spine. Sixth somite with 1

moderate posteroventral spine, and small lobe above it. Telson narrow, tapering to rounded tip, with 3 or 4 !

pairs of dorsolateral spines. Uropods about same length, reaching to tip of telson. n

Color in life: Background of body yellowish wash; carapace, 2 oblique red lines starting in branchial •

region, meeting wide red bar along ventral margin adjacent to third maxilliped and pereiopods 1 and 2; 1

patches of red posterior to antennal spine; large patch of red dots in gastric and cardiac regions; rostrum

mainly transparent with fine red dots on lower limb; antenna, basicerite, and scale with red patches; peduncle 1

and flagellum pinkish; third maxilliped and pereiopods, red spots on milkish base; red blotches on abdomen, n

mainly on lateral surfaces; faint, f'me red speckling on telson and uropods; pleopods with distinctive red lines

III

on basiopodites. (from Butler, 1980: Plate 6E)

Biology. According to Butler (1980), this specis "almost invariably forms part of the catches of I

dredge or shrimp trawl hauls made at 70 m and deeper" in coastal areas of southwestern Canada. See Butler

(1980) for data on density, ovigery, and parasitic isopods infesting this species in that region. I1

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Figure 2.12. Spirontocaris holmesi Holthuis, 1947: A, dorsal and lateral views; B, anterolateral view (A, from

I Butler, 1980;B, from Word and Charwat, 1976).

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Taxonomic Remarks. Characterized by having a very slender body, bladelike rostrum with styliform

projection armed with solitary ventral spine, 2 widely spaced suborbital spines, and epipods on pereiopods 1

I and 2 (Butler, 1980).

Type Locality and Type Specimens. Puget Sound.

I Distribution. Yes Bay, Alaska, to San Diego, California (Schmitt, 1921; Butler, 1980).

I _
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Family Palaemonidae Rafinesque, 1815 I

Pseudocoutierea elegans Holthuis, 1951 I

If

Figure 2.13

Pseudocoutierea elegans Holthuis, 1951:182, Plate 57.--Holthuis, 1955:75, Fig. 46d.--Word and Charwat, I

1976; 171.--Wicksten, 1983a: 19; 1992:8.--Abele, 1976:71.

Material Examined. USNM 001-BRA-02-TX Phase I primary voucher, 3 specimens, largest - n

carapace length including rostrum 5.2 mm, rostrum length 2.9 mm.

m

Description. Genus description: Body more or less compressed. Distal part of rostrum cylindrical •

and unarmed. Basal part wing-like expanded and partly covers eyestalks. These wing-like expansions end l

anteriorly in sharp supraocular tooth. Carapace smooth. Antennal spines present, but postorbital and

pterygostomial spines lacking. Branchiostegal groove runs over length of lateral part of carapace. •

Abdomen with pleurae of third to fifth segments ending in sharp tooth. Posterior margin of telson II

bears three pairs of spines.

Scaphocerite is well developed. I

Mandible without palp, incisor process well developed. Laciniae of maxillula rather broad. No exopods

present on second and third maxillipeds. Only caridean lobe present on first maxilliped, but no flagellum is

visible. I

The carpus of first pereiopod not segmented. Second legs very unequal in size, but about equal in

shape. Last three legs rather slender. Dactylus is simple, without basal protuberance.

Second to fifth pleopods with appendix intema. An appendix masculina present on endopod of second •

pleopod of mate.

The uropodal exopod has the outer margin ending in tooth, which at inner side bears movable spine n

(after Holthuis, 1951:182).

Color in life: Not recorded (but see below).

Biology. Associated with and colored to match the red gorgonian Muricia californica where it co- I

occurs with Heptacarpus kincaidi (pers. comm., D. Montagne via D. Cadien). Ovigerous females among the

8 specimens examined by Holthuis ranged from 9 to 16 mm total length (Holthuis, 1951). Im

Taxonomic Remarks. This species is the sole member of the genus Pseudocoutierea Holthuis. I

Subsequent to the original description (Holthuis, 1951), wherein 8 specimens were listed from Allan Hancock

Expeditions in 1934, 1935, 1937, and 1941, there have been no reports of this species in the literature. •

An unfortunate error exists in the original description (Holthuis, 1951), where the illustrations of this II

species (Plate 55) were switched with those ofAnchistioides antiguensis (Plate 57).

Type Locality and Type Specimens. Holotype is an ovigerous female, 16 mm tong, from off Santa n

Catalina Island, California (Blake Station 1405-41) (AHF, now part of LACM collection) (Holthuis, 1951).

Additional paratypes exist in the LACM collections and at the USNM (AHF No. 411, according to Holthuis,

1951:185). I Distribution. Santa Catalina Island, California, south along the west coast of Baja, California, to the

Galapagos Islands, from 20 to 50 fathoms (37 to 91 m) (Holthuis, 1951:185). This report extends the range

northward to the Santa Barbara Channel. •

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Figure 2,13. Pseudocoutierea elegans Holthuis, 1951: A, anterolateral view; B, third pereiopod (both from

i Holthuis, 1951).

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Infraorder Thalassinidea Latreille, 1831 !

Family Axiidae Huxley, 1879 lII

Acanthaxius spinulicaudus (Rathbun, 1902)

Figure 2.14 I

Axius spinulicauda Rathbun, 1902:886; 1904:I49, Fig. 90

Axius (Paraxius) spinulicauda.--Borradaile, 1903:538. II

Axiopsis spinulicauda.---Schmitt, 1921:111, Fig. 74.--Hart, 1982:44, Fig. 8.--Kozloff, 1987:404.

Axiopsis (Axiopsis) spinulicauda.----de Man, 1925:69. !

1

Acanthaxius spinulicaudus.--Sakai and St. Laurent, 1989:66.

Material Examined. USNM073-BSR-03-TX Phase I primary voucher, 1specimen; MMS California 1

OCS Phase II, voucher lot, Cruise 3-1, station R-I, Rep:3, 35°05.8'N, 120°49.2'W, 91 m, one female, carapace

I

length = 17.8 mm, including 3.5 mm rostrum.

Description. Rostrum reaching middle of second joint of antennular peduncle, slightly deflexed, 1

longitudinally channeled; armed on either side with five to six teeth. Gastric region of carapace traversed by

five carinae, all of which fade out before reaching cervical groove; median one extending along basal third of 1

rostrum, and armed with four spines just behind line of the orbits; outer carinae a continuation of side margins 1

of the rostrum; outer and intermediate carinae unarmed; in front of the narrow median posterior lobe of the

1

carapace, the surface is compressed or pinched to form a short smooth ridge. Eyes black, of same diameter as 1

stalk, not reaching middle of basal antennular segment. Outer maxillipeds reaching end of antennal peduncle. II

m

Abdomen smooth above; pleura sculptured and pubescent (Schmitt, 1921).

Color in life: Carapace translucent pink with coral rostrum and ridges with greenish tinge laterally. •

Antennules and antennae pale pink and white with yellowish flagella. Eyestalks translucent, corneas black. 1

Chelipeds with shell-pink and white ischium and merus, coral red carpus and hand, and orange teeth. Walking

legs pink, coral and white. Abdomen with pleopods and tail fan orange with patches of pale orange or pink

(Hart, 1982). I

Biology. According to Hart(1982), in British Columbia this species burrows in subtidal mudsubstrate

especially in fjords. Very little concerning the natural history of this species has been published. •

Taxonomic Remarks. The illustrations of male and female major chela by Hart (1982, Fig. 8 c and

d) are mislabeled in that they are reversed from her description. We have taken this into account and corrected

the labeling in Fig. 12.1. 1

Type Locality and Type Specimens. Off Bodega Head, California, 62 fathoms (113 m), Albatross

Station 3172. Holotype female, carapace length (including rostrum) = 19.6 mm, rostrum length = 3.5 mm,

abdomen = 31.5 mm. 1

1

Distribution. Schmitt (1921) and Sakai and de Saint Laurent (1989) mention only the type locality

(off Bodega Head, California); there are additional unpublished records from other southern California localities.

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I Figure 2.14. Acfemale, anthaxriusight spcheliped; inulicaudaD, male (Rathbun, , right 1902): cheliped;A,E,matail le, flaaten.ra(A,C,D l view;modified B, carapace from dor Hartsal, 1982 view;, seeC,

text; B,E from Rathbun, 1904).

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Calocarides quinqueseriatus (Rathbun, 1902) I

Figure 2.15

I

Calastacus quinqueseriatus Rathbun, 1902:887; 1904:151, Fig. 91.--Schmitt, 1921:113, Fig. 76.--Balss,

1925:209.----Goodwin, 1952:175. ll

Calocaris (Calastacus) quinqueseriatus.--Borradalle, 1903:539.----de Man, 1925:l18.--Hart, 1982: 50, 1

Fig. 11.--Kozloff, 1987:404.

Calastacus rostriserratus Andrade and Baez, 1977:65, text-Fig. 1. II

Calocarides quinqueseriatus.---Sakai and St. Laurent, 1989:79.

Material Examined. Male, SBMNH Phase Isecondary voucher collection, 005-BSS-01-TX, carapace •

length (including rostrum) = 23.7 mm, abdomen length = 37.8 ram.

Juvenile, MMS California OCS Phase II voucher lot, Cruise 2-4, Station R-6, Rep. 2, 34°41.4'N,

120°57.9'W, 410 m, carapace length = 7.2 mm, abdomen length undetermined (damaged specimen). 1

1

Juvenile, MMS California OCS Phase II voucher lot, Cruise 1-2, Station R-6, Rep. 1, 34°41.4'N,

120°57.9'W, 410 m, carapace length = 5.9 mm, abdomen length = 9.3 mm. I

Description. Carapace surface smooth, cervical groove deep. Rostrum flattened, with toothed margins !

continuing as ridges on gastric area. Median ridge spined from mid-rostrum to gastric area. These ridges are

separated by 2 more short, spined ridges, making 5 in all. Eyestalk short and cornea without dark pigment. •

Antenna with relatively short projections or"thorns" on 2nd and 3rd segment of peduncle. Unequal, elongated 1

chelipeds with toothed margins and surface of hands covered with numerous small sharp granules. A slight

gape proximally between fingers of large cheliped but none on smaller. 1st walking leg short and chelate, with I1

spines on posterior margin of merus and ischium. Others long and slender with setae terminally. Abdomen 1

stout, pleura broad and lateral margins rounded. Tail fan with telson only slightly longer than 6th abdominal

l

segment: Lateral margins parallel and toothed, distal margin curved and with a median tooth. Two spined

ridges on telson and 1 on endopod of uropod. No pleopod on 1st abdominal segment. Slender biramous 1

pleopods on 2nd to 5th segments with appendix intema. In male 2nd pleopod has an appendix masculina as

I

well (Hart, 1982). 1

Biology. According to Schmitt's (1921) key, known only from depths of 200 fathoms (366 m) or !

greater, although in his list of distributions he included a depth of only 160 fathoms offSan Simeon Bay. Hart

(1982) lists the habitat type as abyssal mud, 288-2200 m. •

Taxonomic Remarks. Differs from C. investigatoris (now known asLophaxius rathbunae Kensley, 1

1989, family Calocarididae), which is known from Alaska to San Diego (Schmitt, 1921:113), in having a

carapace that is not granulate and in having 5 rows of spines posterior to the rostrum (Schmitt, 1921). •

Type Locality and Type Specimens. Off San Luis Obispo Bay, 200 fathoms (370 m), Albatross

Station 3196 (Schmitt, 1921). Type is a male, length of carapace and rostrum 28 mm, length of abdomen 41

mm (Schmitt, 1921). II

Distribution. Point Sur, California, south to the Santa Barbara Channel, including Anacapa Island

and San Nicolas Island, at depths greater than 160 fathoms (293 m) (Schmitt, 1921).

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Figure 2.15. Catocarides quinqueseriams (Rathbun, 1902): A, male, lateral view; B, male, right cheliped; C,

I male, carapace, dorsal view; D, tailfan (from Hart, 1982).

I Calocarides sp.

Figure 2.16

I Material Examined. Male, USNM Phase I primary voucher collection, 082-BSR-02-TX, carapace

length (including rostrum) = 22.3 mm, abdomen length = 30.5 mm.

(illustrated) Remthat arksdiffered . Amongsignificantly the specimens from that sentspecies to us asin several Calocarides morphological quinqueserriatus characters,wasmost onenotably specimenthe

shape of the first and second pereiopods and relative length of the carapace and abdomen. Consultation with

I Brian The Kensley, Smithsonian Institution, confn-msthat this is most likely an undescribed species of Calocarides. species complex along the west coast of the United States is currently undergoing investigation by J.

Martin and D. Cadien. Pending the results of their fmdings, which have indicated thus far that either additional

I west specimen coastisspecies referredremain to hereundescribed as Calocarides or thatsp.. the species displays considerable morphological plasticity, this

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Figure 2.16. Calocarides sp. A.: A, lateral view; B, dorsal view of restrain and anterior carapace; C, lateral view

of rostrum and anterior carapace; D, right cheliped; E, left cheliped; F, tetson and left uropod. !

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I Infraorder Anomura Milne Edwards, 1832

i Family Diogenidae Ortmann, 1892

Paguristes ulreyi Sehmitt, 1921

i Figure 2.17

Paguristes ulreyi Schmitt, 1921:125, Plate 18, Figs. 3-5, 7.--Hart, 1982:106, Fig. 37.---Gotshall, 1994:62,

I Plate Plate 128. 145.--Jensen, 1995:67,

Paguristes occator Listed as a synonym by Hart (1982) but not cited.

I Material Examined. USNM 013-BRC-02-TX Phase I 1 SBMNH 095- primary voucher, specimen;

BSS-01-TX Phase I secondary voucher, 2 specimens, largest - shield length 12.1 mm.

i acute Description. Anterior portionof carapace longer than wide; fronttridentate, rostriformtooth triangular,

, extending forward at least to base of the eye scales, exceeding lateral teeth by one-half their length;

lateratteeth blunt andsomewhat tuberculiform. Eye-stalks comparatively verylong and slender,length measured

i from orbital margin about as long as greatest width of anterior portion of carapace, or slightly longer; eye scales each with four to five spiniform teeth at tip; third segment of antennular peduncle exceeds cornea by

one-fourth its length. Antennae thickly long-haired beneath, few short hairs above.

I outer border Chelipeds of lower equal, facehairy; of merus memsinspine occasional d o upper specimens or aterislightly or edge rugose, and on inner and inboone rderspecimen of lower from face,

Monterey somewhat spinulose; carpus with five stout spines on upper inner edge; upper surface of hand well

I spined palm behind dark tipped spines, which are larger on the outer anterior edge of the immovable finger; inner edge of

the dactyl armed with three prominent spines, with two much smaller ones below on inner face,

and intermediate between them. Hands somewhat like those ofP. turgidus, one-third to sometimes nearly onehalf longer Ambulatory than wide,legs butvery withhairy fingerson linner ess acuminate. or anterior face of dactylus and propodus, anterior pair more so

than in any of the species listed in Schmitt, (1921); armature of anterior pair much as in P. bakeri (from

I Schmitt,Color 1921:125). in life: Carapace red-brown with 3 irregular lighter stripes on shield and 1 on each branchial

area. Chelipeds white with orange, light and dark red bands; fingers pink and white with black tips. Walking

legs rest and white, with black claws. Abdomen mottled red and white. Antennule white with dark red streaks and flagellum red ventrally. Antenna pink and red base; flagellum dark red with light areas atjoints and setae

light colored. Eyestalk white with dark red, almost black stripes dorsolaterally and ventrally; cornea black

I (after Hart, 1982; Jensen, 1995:67, Plate 128). Biology. This species is often found intertidally in rock crevices on exposed shores and subtidally in

rocky and graveled areas (Hart, 1982).

I Taxonomic Remarks. Paguristes uIreyi differs from P. perrieri in having the eye scales or

toothed

incised and not entire, longer antennular peduncles, which exceed the eye-stalks instead of being considerably

shorter, and in having the antennal flagella well haired beneath instead of sparsely so (Schmitt, 192I).

I Paguristes ulreyi differs from P. turgidus by the antennal flagellum being densely setose ventrally and

sparsely dorsally, eyestalks proportionately long and more slender, nearly as long as shield and hand with dark

i comeous teeth only at tips of fmgers. Brood pouch of female subrectangular (Hart, 1982).

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Figure 2.17. Paguristes ulreyi Schmitt, 1921:A, male,dorsal view; B,telson (from Hart, 1982). !

mm), fromTypeoffLoPoint calityLoma, and TCalifornia, ype SpecimAlbatross ens. Holotype Station(USNM 4304, 2550427) fathoms is a 59(46mmm) male (Schmitt, (carapace 1921).length Schmitt 22 I

gives additional measurements and mentions paratypes, which we assume are also housed at the USNM.

California, Distribution. Monterey to off San Diego, California, and San Geronimo Island, Baja I to a depth of 59 m (Schmitt, 1921).

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I Family Paguridae Latreille, 1803

i

I Parapagurodes iaurentae McLaugldin and Haig, 1973

Figure 2.18

I ?Pagurus [sp]:--Menzies and Miller, 1954:153.

Parapagurodes laurentae --McLaughlin and Haig, 1973:129, Figs. 4b, 9-11.

I Material Examined. USNM 006-BSS-01-TX Phase I primary voucher, I ovigerous female, shield

length 1.3 mm, lleggs (near hatching), diameter about 0.45 mm each.

I rows of Des widely cription. spaced,Palmstrong, of right tubular cheliped spines, withnotdorsal extending surfaceontoarmedfixedproxi finger. mallyPalmwith offourleftorcheliped five irregular with

dorsal surface havng single or double row of strong, tubular spines, not extending onto fnted finger. Palm of

left cheliEleven ped withpairdorsal surface having single or double row of strong tubular spines. s of phyllobranchiate gills. Third maxilliped with basis-ischium fusion incomplete; ischium

with crista dentata well developed, one accessory tooth; merus and carpus each with spine at dorsodistal

I margi not recurved. n. MaxillOcular ule withpeduncles proximalshort enditeor moderately tapered; endopodite short, stout, withwithexternal corneaelobedilated. moderatel Ocular y wel acicles l-developed, slender

to moderately broad, triangular or subtriangular, terminal spine submarginal. Antennal peduncles with one or

I two and third shortpereiopods setae on dowith rsodistal dactylsurface moderately of ultimelongate, ate segme slender, nt. Che notlstrongly ipeds uneq twisted. ual, right Fourthlarger pereiopods than left. subchelate Second

or not subchelate; dactyls usually with preungual process (cf. de Saint Laurent, 1970) on lateral face; propodal

I rasp sexualweakly tube; ornomode pairedratelypleopods, well-developed. pleopods M3ales through wit coxae 5 biramous of fifthwith pereiopods rami weakly symmetrical, deveoped, right orwithabsent.hort

Females with paized gonopores; no paired pleopods, biramous pleopods, pleopods 2 through 4 weakly or

I moderately margins generally well-devestraight, loped, pleopod slightly5concave, weakly developed or slightlyor abse oblique, nt. Uropods with a asymmetrical. row of small Telson spines with or spinules. terminal

Sternite of third pereiopods with anterior margin unarmed (from McLanghlin and Haig, 1973).

I Cadien, Color pers. comm.). in life: Carapace translucent gray. Pereiopods with two broad bands at mid propodus (D.

Biology. Often found infested with a bopyrid isopod of the genus Stegophryxus.

I According to Mclaughlin Haig (973), although Parapagurodes

Taxonomic Remarks. and 1 laurentae

is most closely related to P.makarovi, its superficial resemblance to an undescribed species of Pagurus may

cause mistakes in identification. To avoid such mistakes, distinguishing characters ofP. laurentae include the

I of sexual tubes in the males, the lack of spines the distal portions of the dorsal surfaces of the presence on

dactyls of the chelipeds, the moderately long and relatively slender dactyls of the second and third pereiopods,

and the lack of regular setation on the articles of the antennal flagella (after McLaughlin and Haig, 1973). In

I freshly preserved or living material, P.laurentae and P makarovi are easily separated by color pattern. In P.

Iaurentae red bands circle the propodus of the pereiopods, whereas in P. makarovi red stripes extend the

length of both the carpus and propodus (D. Cadien, pers. comm.).

Type Locality and Type Specimens. 2.5 mi SE of Seal Rocks, Santa Catalina I., California, 16'

33°

20" N, 118° I5' 20"W, Velero IH Station 1429-41,159-174 m. Holotype: Male (sheild length = 3.5 mm), AHF

4127. Paratypes listed in McLaughlin and Haig, 1973: Table 3.

i Distribution. Southem California, Channel Islands to west coast of Baja California, Mexico; Gulf of

California; 16 to 475 m (McLaughlin and Haig, 1973).

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Figure 2.18. Parapagurodes laurentae McLaughlin and Haig, 1973: male paratype - A, shield; B, mesial view of I

right cheliped: C, dorsal view of right cheliped: D, mesial view of left cheeped (modified from

McLaughlin and Haig, 1973).

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i Family Galatheidae Samouelle, 1819

I Munida quadrispina Benedict, 1902

Figure 2.19

i Munida quadrispina Benedict, 1902:269, Fig. 17.--Rathbun, 1904:166.---Schmitt, 1921:165, Fig. 105 (after

Benedict).--Hart, 1982:168, Fig. 66.--Kozloff, 1987:411, Fig. 19.20.--Gotshall, 1994:65, Plate

i 154.--Jensen, 1995:73, Plate 145.

Material Examined. USNM 004-BRA-04-TX Phase I primary voucher, 4 specimens, largest -

carapace length including rostrum 10.2 mrn, rostrum length 3.2 mm.

I Description. Abdomen unarmed. Rostrum long, slender, spine-like, laterally compressed, unarmed,

moderately serrate above and slightly so below. The straight, slender supraocular spines do not reach quite to

i the ends of the eyes and are united to the rostrum for nearly one-half their length. The marginal spines of the carapace vary from eight to ten on a side; there are six spines on the gastric area, four in a line behind the

supraocular spines, and one on each side near the hepatic region; the terminal spines of the line are very weak

and small, but one spine occupies the anterior branchial region; posterior margin unarmed. Chelipeds well set

I with spines and spinules, not hairy (after Schmitt, 1921).

Carapace longer than wide. 6 spines on gastric area; 4 in one line, and 1on each side of the ridge near

I the withhepnumerous atic regiospines n. 8 toand10narrow marginalfingers. spines.Walking Rostrumlegslongcompressed and compressed and armed and with minuteshort ly spined. spines. Chetipeds Abdomen

ridged but not spined except for minute spines on tail fan. Males with paired uniramous gonopods on 1st and

i 2nd abdominal Color segments. Female with small paired pleopods on segments 2 to 5. in life: Carapace and abdomen red-brown overall, ventrally lighter or white; ridges red with

grooves white, and blue spots in cervical groove. Chelipeds with red spines; fingers red with distinct white

I tips. 145).Walking legs with irregular light bands. Tail fan light colored (after Hart, 1982; Jensen, 1995:73, Plate

Biology. These animals inhabit rocky areas, gravel, mud, and sponge beds. They are predators on

I other Crustacea and can do considerable damage in shrimp traps (Hart, 1982).

Taxonomic Remarks. Commonly referred to as a squat lobster. Rathbun (1904:166) remarked that

this is "probably the species designated by Owen (1839?) as M. gregaria."

I Type Locality and Type Specimens. Holotype is a 35 mm long individual from off Cape Beale,

Vancouver Island, British Columbia, 66 fathoms (121 m), Albatross Station 2878 (Schmitt, 1921).

1904; ScDis hmitribution. Sitka, Alaska, to San Diego, California, 50 to 559 fathoms (91 to 1,022 m) (Rathbun, tt, 1921).

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Figure 2.19. Munida quadrispina Benedict, 1902: male, dorsal view (from Hart, 1982).

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Pleuroncodesplanipes Sfimpson, 1860 i

Figure 2.20

I

Pleuroncodesplanipes Stimpson, 1860:245.--Holmes, 1900:112.--Rathbun, 1904:166.---Schmitt, 1921:163, I

Plate 31, Fig. 2.--Boyd, 1967:394, Figs. 1-4.--Gotshall, 1994:64, Plate 153.--Jensen, 1995:74,

Plate 146. I

Material Examined. USNM 102-BSS-01-TX Phase I primary voucher lot; SBMNH 023-BSR-02-

TX Phase I secondary voucher lot. •

1

96 l
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I Figure 2.20. Pteuroncodesplanipes Stimpson,1860: male, dorsal view (from Boyd, 1967).

I Description. Carapace more or less convex, tapering anteriorly, transversely rugose, anterior edges of mgae fringed with closely set short hairs; with the exception of a few spinules behind the supraorbital teeth,

carapace is devoid of spines above; there is a spine at the rounded anterolateral angle, behind which there are

I a few spines on the lateral margin. Rostrum long and slender, scabrous above, and continued onthe carapace as a carina; supraorbital teeth spine-like, confluent at base with rostrum. Chelipeds slightly hairy; ambulatory

legs scabrous, with penultimate joints distinctly flattened and ciliated (Schmitt, 1921).

I Color in life: Red (Jensen, 1995:74, Plate 146)

Biology. One of the most common species in southern California, P.planipes is usually encountered

as a pelagic species in huge schools which occasionally wash ashore in vast numbers. Boyd (1967) however,

I species partially during two years wholly benthic after this time with

found that this is benthic its first and

densities measuring as high as 9 to 11individuals per m2.Animals were dredged and trawled at depths ranging

from 70 to 900 m on gray muddy sand and gray mud along the western coast of Baja California, Mexico.

I Boyd (1967) states that this species is a voracious omnivore that is in turn fed oceanic

upon by large game

I 97
I

i

fishes such as albacore, yellowfin tuna, and skipjack tuna, and is also fed upon by kelp-bed fishes. During its •

pelagic phase, P. planipes is a filter feeder near the surface. It spreads its setae-fringed legs to form a filter

basket and slowly sinks through the water column collecting particulates, small zooplankters, and perhaps •

larger phytoplankton, these are ingested as the legs are groomed, and the animal swims back to the surface to II

repeat its slow feeding descent (D. Cadien, pers. comm., based on his observations from the Texas A&M

University submarine Diaphus, in the Santa Maria Basin). Schmitt (1921) reports that a mass stranding •

which occured in 1859 was the source of specimens for the species description by Stimpson in 1860; subsequent I

reports of this phenomenon are numerous.

Taxonomic Remarks. This species is often referred to as the tuna crab or red crab. It is also included •

with those species known as squat lobsters.

Type Locality and Type Specimens. Schmitt (1921) gives two type localities: Monterey, California,

and a latitude and longitude of 24_N 130W¢,Pacific Ocean (Schmitt, 1921:163). 1

I

Distribution. From 90 miles southwest of San Francisco, California, to 150 miles southwest of Cape

St. Lucas, Lower (Baja) California (Rathbun, 1904; Schmitt, 1921). I

i

Family Lithodidae Samouelle, 1819

Paralomis multispina (Benedict, 1894) •

Figure 2.21 i

Leptolithodes multispinus Benedict, 1894:484.--Rathbun, 1904:165.

Paralomis multispina.--Bouvier, 1896:25.---Schmitt, 1921:159, Plate 23, 30,Figs. 7-8.---Goodwin, 1952:176, i

i

Fig. 8.1Hart, 1982:88, Fig. 28.--Kozloff, 1987:408.--MacPherson, 1988:76.

Material Examined. USNM 013-BRA-04-TX Phase I primary voucher, 1specimen, carapace length i

including anterior and posterior spines 3.02 mm, without spines 2.57 mm, carapace width with spines 2.57

ram, without spines 2.13 mm; SBMNH 016-BRA-01-TX Phase I secondary voucher, 1 specimen.

Description. Carapace about as broad as long; areolations well def'med. Median fine at summit of B

gastric region has a sharp spine about 4 mm in length. The lateral margins are armed with from twelve to i

sixteen spines about 3 mm in length. In young and in some adults there are small spines on the branchial

region. A semicircular line of six or seven spines marks the limits of the branchial and intestinal regions. !

Carapace thickly studded with blunt spines, each terminating in a flattened face or surface cut obliquely to the

surface of the carapace; this face is encircled by a fringe of short, stiff bristles. Rostrum consists of simple

median spine with two basal spines. Under the rostrum proper there is a very short, conical spine homologous n

with the subrostral spine ofLithodes; behind the spine are one or more spinules. Abdomen in male composed,

after the second segment, of several rows of leathery plates; second segment better calcified and harder.

Abdomen of female twisted to the right as in Lithodes. i

Chelipeds moderately slender, extending almost to distal end of carpal joints. Spines on inner margin

of carpal segments the most prominent. Ambulatory legs long and slender, thickly set with spines. Spines of

merus not so distinctly arranged in rows as on carpal and propodal segments; there is, however, a distinct row

on the upper margin. Spines of carpus arranged in eight more or less distinct rows; on the propodal segment J

the spines are arranged in six full rows and two half rows. There are four short rows of spines on the proximal

end of the dactylus. Dactyli compressed, slightly bent, a little twisted (Schmitt, 1921:159, after Benedict). i

II

Color in life: Carapace scarlet with spines dark and nodules fight. Chelipeds and walking legs scarlet

with lighter colored spines and tips of fingers. Eyestalks scarlet; corneas black. Antennal flagellum scarlet

(Hart, 1982). i

98 n
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i Biology. Hart (1982) reports habitat as deep sea mud.

Taxonomic Remarks. According to MacPherson (1988:76), there are 24 species of this genus in the

I Pacific (including Antarctic waters). Off California only two species occur, P multispina and P. verrilli (Benedict, 1894). The species are easily separated by the more spiny carapace and legs of P. multispina

(compare Plates 23 and 24 of Schmitt, 1921). The correct date of Benedict's description is apparently 1895,

I although the date appearing on the frontispiece of that volume is 1894 (Dawson, 1989). Type Locality and Type Slaeeimens. Off Queen Charlotte Islands, British Columbia, 876 fathoms

(1,602 m), Albatross Station 2860.

I Distribution. From off Shumagin Bank, Alaska, to off San Diego, California, 625 to 876 fathoms

(1,143 to 1,602 m) (Schmitt, 1921, after Rathbun, 1904); Japan to Gulf of California, deep water (Dawson,

i 1989).

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I Figure 2.21. Paralomis multispina (Benedict, 1894):male, dorsal view (from Hart, 1982).

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Infraorder Brachyura Latreille, 1803 I

Family Homolidae De Haan, 1839 I

m

Paromolafaxoni Sehmitt, 1921

Figure 2.22 I

Paromolafaxoni Schmitt, 1921, Plate 31, Fig. 7.----Rathbun, 1937: 68-69, Plate 18; Plate 19, Fig. 1; Table

19.-- Crane, 1937: 107.-- Church, 1971: 113 (color photograph, presumably this species).-- Luke, R

1977: 32.-- Wicksten, 1983b: 185, Fig. lb (drawing based on Church's (1971) photograph). Kuck I

and Martin, 1994: 178, Figs. 1-4.

[]

Material Examined. California: Santa Maria Basin, 250 m, MMS BioSurvey Station 1 C/D. One I

male, carapace length (including 4.9 mm rostrum) 35.8 mm; carapace width = 31.2 mm (USNM 001-BRC02-TX). I Type Locality and Type Specimens. Off Point Loma, California, 67 to 73 fathoms (123 to 134

meters), Albatross Station 4309, 03 March 1904. Type specimen USNM 53331, female.

1

Description. Carapace: Very short covering of setae over entire carapace, but not obscuring surface I

(i.e., not with "entire surface more or less obscured by a rather thick, short pubescence" [Schmitt, 1921]).

m

Supraorbital spines directed outward, with two (sometimes one; e.g., MMS 1 C/D) small, hooked spines. m

Branchial and hepatic regions distinctly inflated; body subrectangular in cross section, becoming more inflated 1

with increased size of crab. Lateral margins with row of 4-6 spines. Linea homolica visible along dorsolateral

m

margins (see Martin and Abele [1986: 587; 1988: 32] for discussion of lineae of decapods; see McLaughlin l

[1980: 167] for defmitions of types of tineae found in decapods). Small tubercles of varying number and size 1

covering carapace both dorsally and laterally, largest concentration on dorsal branchial region (Kuck and

Martin, 1994). 1

Chelipeds: Long, slender, equal in length; covered with dense, short setation; merus with row of 5-8 I

spines dorsally in females. Six specimens with variously sized, darkly pigmented, subcircular spot distally on

both sides of propodus (LACM 62-204. I has spots only on inner side of propodus; CMM 86.26.001 has spot 1

on inner side of right propodus only.) Nature of this spot, and why it appears in only some specimens of both

sexes, is unknown. Fingers with non-serrate, acute inner margins; dactyl crossing propodus inwardly at tip

(except one chela of specimen from MMS 1C/D); with patches of long, thick setae in most specimens; finger 1

pigmentation (preservation in 70% ethanol) ranging from white to brown to black.

Pereiopods: Pereiopods 2-4 with row of 15-22 corneous spines on inner (ventral) margin of dactyl;

merus with 6-9 dorsal and 3-12 ventral spines along margins; Pereiopod 5 dactyl with 5-8, and propodus with []

7-10 (proximal 5-6 in circular pattern) comeous spines on inner margin. Dorsal and lateral surfaces of propodus 1

densely covered with short setae; ventral surfaces of dactyl and propodus mostly devoid of setae.

Abdomen: Seven-segmented; large tubercle at posterior margin of first, and in center of second, •

segments; occasionally, small tubercle at anterior margin of first, large or small tubercle in center of third, and II

small tubercle on anterior margin of fourth, segments.

Male: Chelipeds more robust; larger specimens with propodus and carpus covered with prominent

tubercles, fewer and sometimes absent in smaller males and in females. Merus with 9 to 19 dorsal spines;

occasionally, additional parallel row of up to 11 spines in larger specimens. Abdomen with tubercle along

horizontally raised posterior border of sixth segment (also found in small female from LACM 41-33.9); •

anterior half to two-thirds of seventh segment generally narrower at base than in females. Pleopod 1 large,

robust, minutely crenulate at distolateral tip; overlapping ventral folds with long setae covering posterior

opening. Pleopod 2 as in Fig. 3c. Pleopods 1 and 2 becoming more robust and better defined with increased •

size of specimen.

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I Small individuals (e.g., LACM 41-33.9): Carapace with spines longer and sharper than in adults;

virtually no setae dorsally, except at tips of largest lateral, gastric, and hepatic spines; short hooked setae

I along irregularly somespaced margins; setae; rostrum, very fewsupraocular well-spacedspines, setae pereiopods, covering chelipeds. antennae,Supraocular and eyestalks spineswithdirected long, forward slender,

with small, hooked dorsal spines sometimes lacking (e.g., LACM 40-148.1). Chelipeds with 5 dorsal spines

I onfewer merus; and less fingers prominent withoutspines pigmentation. along dorsal Pereiopods and ventral 2-4margins proportionately of merus,longer than inandadults. morePereiopod slender, and5 dactyl with

with 6 and propodus with 8 spines.

I (preservation Color:inAll70%specimens ethanol) examined except forbyLACM Kuck and 66-371.1, Martinwhich (1991)hadappeared a whitishuniformly carapace.lightSeeto"chelipeds" dark cream

above for description of cheliped spots and finger pigmentation. Crane (1937) describes "color in life" as

I "entirely buff, except anterior part of carapace which is suffused with scarlet"(Kuck and Martin, 1991). Biology. Very little is known about the basic biology of P.faxoni, as is the case for many deep-sea

brachyurans. They have been recovered from vase-shaped hexactinellid sponges off Point Loma, California,

I The and were loath to leave the sponges' protective cover once collected (pers. comm., R. Velarde, via D. Cadien). reduced and subchelate fifth pereiopods suggest a carrying behavior, as has been seen in other members

of the genus (e.g., Wicksten, 1985, for P.japonica), with these legs holding some object(s) over the back of the

I carapace southern California" for protectioncarrying or camouflage. a sponge; Wicksten her drawing(1983b: was based fig. lb)onshows a colorwhat photograph is probably in Church a P.faxoni (1971)"frofoma

"10-inch-long [specimen] at a depth of 1,200 feet [366 m] off San Diego." Based on pereiopod morphology,

I Guinot and Richer de Forges (1981) and Wicksten (1985, 1986) inferred carrying behavior in all species of the family Homolidae. Similar behavior has been documented or inferred for other crabs with chelate or

subchelate posterior pereiopods (e.g., Dromiidae, Latreillidae, and Dorippidae).

I Only one ovigerous female exists in examined collections (LACM 62-205.1). A second was collected off San Diego, California, on 28 January 1992 by the Pt. Loma Biology Laboratory (OMP), but was discarded.

We know of only four P.faxoni that have been held successfully in aquaria (three females at the Cabrillo

I Marine Museum, and one male at the Scripps Institution of Oceanography). The only behavioral notes available are from observations by S. Vogel (CMM; pers. comm.) of crabs in aquaria carrying gorgonians with the fifth

pereiopods, and by R. McConnaughey (SIO; pers. comm.) of a crab that lived in an aquarium (5-10°C) from

I 05 May to October 1992. McConnaughey noted that the crab maintained the fifth pereiopods in an elevated position, but no obvious carrying behavior was observed (although shells and other items were introduced).

The species has also been found when trawled to still carry a gorgonian colony not otherwise represented in

I the trawl (and thus not likely to have been picked up after capture) (pers. comm., R. Velarde via D. Cadien). Taxonomic Remarks. Although the holotype (USNM 53331) is a female, and is the only specimen

mentioned by Schmitt (1921), his figure (Plate 31, Fig. 7) was erroneously referred to as a male in the

I accompanying figure legend. Rathbun (1937) used the same photograph, but correctly referred to it as a

female (holotype), and also included a photograph of the holotype in ventral view (clearly showing it to be

female). Rathbun (1937: 72, Table 18) listed the five previously known specimens, all of which were collected

I by the Albatross in 1904 and are housed at the USNM (R.B. Manning, pers. comm.).

The male first and second pleopods have been illustrated for relatively few species of Paramola.

Griffin (1965: Figs. 1-4) provided figures for P. spinimana, as did Serene and Lohavanijaya (1973: figs. 36-

I 37) for P.aIcockifaughni, and Guinot and Richer de Forges (1981: Figs. 5A, A 1 and 5C, C1) forP. profundorum and ?P.japonica Parisi, 1915. The general shapes of the pleopods of these species and those of P.faxoni are

quite similar, including the distinctive overlapping ventral folds of the fn-st pleopod (Fig. 3a; ventral view not

I shown in Serene and Lohavanijaya, 1973). The apex of the first pleopod is distinctive in these species, as in

many brachyurans, and most likely provides a good distinguishing characteristic among males of Paromola

species.

I

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Figure 2.22. Paramolafaxoni (Schmitt, 1921): Holotype female (carapace width 39.60 mm) USNM 53331,

dorsal view (from Kuck and Martin, 1994). 1

1

Distribution. All five specimens of P.faxoni listed by Schmitt (1921) and again by Rathbun (1937) II

were taken off Point Loma, California, in depths from 123 to 247 m. Wicksten (1985) examined another

specimen housed at the Allan Hancock Foundation (AHF), Los Angeles, California, where we found nine I

additional unreported specimens (Tables 1 and 2) distributed from Tajiguas, California (approximately 27 1

miles east of Point Conception, Santa Barbara County, CA), to Isla Cedros, Baja California, Mexico. The

only other published record for this species is a collection catalog by Luke (1977) documenting ten specimens Im

(sixmales, three females, and one juvenile) housed at the Scripps Institution of Oceanography and distributed •

from La Jolla, California, to Cabo Colonet, Baja California, Mexico. According to staff of the Monterey Bay

B

Aquarium Research Institute (MBARI), Monterey, California, there have been no sightings ofP. faxoniin the l

area during the past four years either by personnel or in extensive video footage (J. Berry, J. Connor, L. Lewis, 1

pers. comm.). Queries to the California Academy of Sciences (CAS; R. Van Syoc, pers. comm.), San Francisco,

California, and to the Universidad Nacional Autonoma de Mexico (UNAM; M. Hendrickx, pers. comm.), 1

Mazatlan, Mexico, did not result in any additional specimens. Additionally, Hendrickx (1992) did not list P. II

faxoni as occurring in the Gulf of California, Mexico. Therefore, it is likely that the geographic distribution of I

P.faxoni does not greatly exceed the boundaries defined by present specimen collections, although distribution

of the species west to Point Conception, CA (northernmost border of the California Province; e.g., Brusca and II

Wallerstein, 1979:73), is highly probable. 1

The report of Kuck and Martin (1991 ) extended the range ofP. faxoni northward to Tajiguas, California, m

and southward to Isla Cedros, Baja California, Mexico, and extends the depth range to 18-460 m. 1

11

102 I
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I Family Xanthidae MacLeay, 1838

I Lophopanopeus bellus diegensis ORathbun, 1900)

Figure 2.23

I Lophopanopeus diegensis Rathbun, 1900:37; 1904:184, Plate 9, Fig. 3; 1930:327, Plate 153, Figs. 6-7, 10,

text-Fig. 49.--Weymouth, 1910:52, Plate 12, Fig. 39.---Schmitt, 1921:245, text-Fig. 146, Plate 37,

i Lophopanopeus Fig. 5.--Johnson bellus diegensis and Snook, .--Menzies, t935:387.1948:7, Figs. 6-8, (male hypotype, AHF Station 1407-40).-

Allen, 1980:238, Plate 35, Figs. 402, 405.--Hart, 1982:200, Fig.81.--Jensen, 1995:19, Plate 7.

I Lophopanopeus bellus.--Charwat, 1977:71.

Material Examined. USNM 013-BRA, 04-TX Phase I primary voucher, 1immature female; carapace

I length immature= 3.3male, mm, carapace carapacelengthwidth= 2.8= 4.2mm,mm. carapace SBMNHwidth001-BRC, = 3.9 mm.02-TX Phase 1 primary voucher, one

Description. Ambulatory legs with dorsal and lateral surfaces, except on the lobations, covered with

I a pubescence; carpus markedly bilobed. Carpus of cheliped covered with irregular generally isolated elevated bumps; carpal number for bumps 4-6; a deep sulcus is located on dorsal surface of carpus near its anterolateral margin. Lobe at proximomedial part of dorsal surface of manus small. An enlarged tooth is present at

I the b proximal end of cutting edge of dactylus of major cheliped. Dorsal surface of carapace pubescent as inL.

. bellus. Morphology of tip of flagellum of male identical with that of L. b. bellus (after Menzies, 1948:7).

Original description: Upper margin of meral joints of ambulatory legs not spinulous. Hands with one

I or more lobes or teeth on upper margin. Carpus of cheliped very rough. Carpal joints of ambulatory legs strongly bilobed. Carpus of chelipeds covered with tubercles (Rathbun 1900:137).

Discussion of constant features: Only two characters, (1) the presence of irregular, elevated, isolated

I the dorsal surface of the of the and of bilobed bumps on carpus cheliped (2)the presence markedly cm-pal

segments of the ambulatory legs, serve to distinguish this species from L. bellus bellus (Menzies, 1948).

Color in life: Color and pattern variable ranging from purple and orange to brown, white, or mottled

I (Jensen, 1995:19, Plate 7).

Biology. This species has been collected from various habitats ranging from unprotected rocky shores

I tocourse potected sand tobayrock. s at dBelow epths ranging 20 fathoms from(37intertidal m) L. bellus to 100diegenensis fathoms (185is frequently m), on bottoms takenranging with L. from leucomanus mud to

leucomanus. Intertidal specimens are in general much larger than those found below 10 fathoms (Menzies,

I I948). Ovigerous females have been found from February to July and also in September.

Taxonomic Remarks. This subspecies ofLophopanopeus bellus was initially described by Rathbun

I (1900) as a distinct species,L, diegensis. Menzies (1948) considered it a subspecies ofL. bellus and described the differences between this and the other, more northern, subspecies, L. bellus bellus. However, Menzies

noted several problems, including areas of geographic and morphological overlap (see Menzies, 1948).

I In the two juveniles these collections, the pubescence of the pereiopods and carapace that Menzies (1948)

in

employed in his key is not readily evident. However, the distinctive nature of the bilobed crest on the carpus of

the walking legs, and the irregular raised bumps of the cheliped carpus, are evident even at these early

I developmental stages.

Adult specimens of/_. belIus diegenensis average approximately one half the size in carapace width

i of adults of L beIlus bellus.

I 103
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,

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,

D

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Figure 2,23. Lophopanopeus bellus diegensis Rathbun, 1900: A, anterodorsal margin of carapace; B, male, •

dorsal view; C, male, lateral view of fight cheliped; D, male holotype, dorsal view of right carpus;

E, male holotype, lateral view of second ambulatory leg; F, male holotype, lateral view of fourth

ambulatory leg (A, from Allen, 1980;B,C, from Hart, 1982; D-F,from Meazies, 1948). 1

1

l

Type Locality and Type Specimens. San Diego, 10 fathoms (18.3 m), holotype male (Rathbun's m

(1900) L. diegensis), USNM 4281. Hypotype, AHF Station 1407-40. m

Distribution. Lophopanopeus bellus diegenensis is the southern form of the species and ranges from I

Mission Bay to Monterey Bay, California. A sharp intergradation zone between L. b. diegenensis and L. b.

bellus is found between Point Conception and Point Sur, California, but specimens ofL. b. diegenensis have m

been identified from as far north as Washington and Alaska (Menzies, 1948). In some areas near Tacoma, I

Washington, in Puget Sound, I_,b. diegensis far outnumberL, b. beUus, with both forms occurring in a single

sample (D. Cadien, pers. comm.). This suggests to us that specific rather than subspecific status may be

warranted. I
I

Family Parthenopidae Alcock, 1895

I Heterocrypta oceidentalis (Dana, 1854)

Figure 2.24

I Cryptopodia occidentalis Dana, 1854:430.--Stimpson, 1857:458.--Lockington, 1877b:78.--A. Milne

Edwards, 1878:169.

: • Lambrusfrons-acutis Lockington, 1877a:31. Type locality, Santa Catalina Island, California; type not extant

II (destroyed in San Francisco fire).

Heterocrypta occidentalis.--Holmes, 1900:44.--Rathbun, 1904:170; 1925:559, Plates 204, 205; 1926:28.--

Weymouth, Schmit-t,1921:192, 1910:21, Fig.Plate 119. 2, Figs 4, 5.--Hilton, 1916:71, Fig. 7.--Nininger, 1918:36, Fig. 1.-- Johnson and Snook, 1927:363, Fig. 316.--Ricketts and Calvin, 1939:182,

Plate 35.--Garth, 1958:476, Plate Z4, Figs. 14 and 14a, Plate 55, Fig. 2.--Allen, 1980:236, Plate

I 32, Fig. 379.--Gotshall, 1994:69, Plate 165.--Jensen, 1995:34, Plate 53.

Material Examined. One juvenile, damaged, possessing only left pereiopod 1 (cheliped) and right

pereiopod 5, carapace length = 2.9 mm, carapace width = 2.8 mm, USNM 021-BSR, 02-TX Phase I primary

voucher.

Description. Size to 21.0 mm carapace length, 34.0 mm carapace width. Branchial ridges sinuous,

terminating posteriorly. atFingers lateralshort, angle,making and notacontinuing distinct angle, on gastric thoughregion, less than marked a right by two angle,ridges withcoming palm. Male to a point first

pleopod thick, capitate, contorted, tapering to "neck" then flaring; longer setae on lip of groove, in two rows

I inreflexed, concavitycorneous of terminal tp marginally portion, adenticulate; nd marginally.a subterminal Male secondcluster pleopodof denticles. little more than half the length of t-n-st,

Carapace broadly triangular; median region narrow, the flattened upper surface bounded by two

granulated ridges, converging to a point behind. Cardiac region furnished with a three-sided, pyramidal elevation, the edges usually granulated. Posterolateral regions large, and furnished with an S-shaped, granulated crest,

extending from near the posterior end of the median region to the acute lateral angles on the carapace; in front

I of Anterolateral the anteriormargins bend of this crest a pair of minute tubercles. Rostrum triangular, subacute, not depressed. straight or slightly concave in front, convex near the middle, the posterior portion

passing outwards and backwards, arching over the legs; the teeth on the anterior part small and irregular, but

I becoming posterior margin larger posteriorly, not producedthereoverfurnished the abdominal with secondary segments.denticles. IschiumPosterolateral of the maxillipeds marginssmooth, transverse; the

anterointernal angle produced; merus small, the surface concave and bearing a prominent tooth near the

I middle. Chelipeds long, Irigonal; the sides of the merus convex, the edge sharply granulate to dentate; carpus

with three of four granulate lines; hand about as long as the merus. The angles prominent and dentate and the

I sides concave; pollex short, deflexed; dactyl short, but longer than long axis of the hand. Ambulatory legs compressed, strongly carinated above; dactyls narrow, strongly sulcate, and corneous-tipped (Garth, 1958:476,

modified from Holmes, 1900:44).

I A long triangular concave area extends from the subhepatic region back to the afferent branchial openings. It includes the exognath and is surrounded by a fringe of hair; outside this area, and above the

ischium of the cheliped, a raised, level, oval area against which the inner surface of the manus contacts when

I the cheliped is flexed (modified from Rathbun, 1925:559).

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A !

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__I - • _ £ : .,,.,.i :%' B l

_" l

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I

Figure 2.:14. Heterocrypta occidentaIis (Dana, 1854): A, dorsal view; B, male, first pleopod; 12,male, second

pleopod (A, from Allen, 1980;B-C, from Garth, 1958). !

Color in life: Tips of tubercles white, with ridges bearing tubercles light purpleish, and remainder of

carapace mottled with numerous minute spots of white and purplish, giving a pink effect which often closely II

approaches white. Ambulatory legs usually a light yellow (Weymouth, 1910:21) (see Gotshall 1994, Plate

165; Jensen, 1995:34, Plate 53). A specimen collected on SCUBA off of Malibu by one of us (TLZ) had the I

following coloration; carapace white, raised regions gray with purple ridges.

Biology. This species has been found on sand, sand and shell, algae, gravel, mud, rock with kelp, I

algae, and coralline substrates. Often associated with sponges, worm tubes, and/or urchins. !

I

Collected from depths normally ranging from 4-57 fathoms ( 7-105 m) with extreme records being

intertidal and 95 fathoms (175 m).

Ovigerous females have been taken from February through May, and July through September. In !

February at San Diego, 38 of 44 females were with ova, and in April off Newport Beach, 51 of 51 females

were in this condition (Garth, 1958:476). Ill

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I Taxonomic Remarks. Garth (1958) states that this species is most abundant off Huntington Beach

and in Newport Harbor. Many large males were found in the 20 mm range, but none were found over 21 mm.

I rostrum Young unisder present 10 mmandaencrustations re considered oftobarnacles be immature, occurbutonindcarapaces ividuals andcanchelipeds be sexed ofat older 5 mm.specimens. Often a sharp

Type locality and Type Specimens. Monterey, California, carapace width = 31.7 mm, sex not

I stated, "probably in Yale Museum" (Garth, 1958, after Rathbun, 1925). Distribution. Gulf of Farallones to San Diego, California, and Los Coronados Islands, 13 to 36

fathoms (24 to 66 m) (Rathbun, 1904) (see Garth, 1958, for additional records).

I

Family Majidae Samouelle, 1819

I ErUeptus spinosus Rathbun, 1893

i Figure 2.25

Anasimus rostratus Rathbun, 1893:227; 1904:171, Plate 10, Fig. 4.--Weymouth, 1910:27.

I Anasimus spinosus.--Schmitt, 1921:196, Fig. 121. Erileptus spinosus Rathbun, 1893:227; 1904:171, Plate 10, Fig. 1; 1925:68, Plates 212, 213, Fig. 18.-

Holmes, 1900:21.--Weymouth, 1910:27, Plate 3, Fig. 7.---Crone, 1937:54.--Garth, 1958:91, Plate

I E, Fig. 8; Plate 5, Fig. 2.--Jensen, 1995:27, Plate 28, Fig. 6.

Material Examined. One ovigerous female, carapace length (including 1.4 mm rostrum) 7. I mm,

I carapace width = 4.7 mm. Specimen number 027-BRA, 02-TX, Phase I Voucher specimen, USNM. One ovigerous female, carapace length = 4.9 mm (including rostrum, broken at tip), SBMNH Phase I secondary

voucher collection, 006-BRA, 02-TX. One juvenile male, carapace length = 4.2 mm (including rostrum,

which California is broke OCSn Phase at tip)II,in voucher vial withtot, twoCruise other,3-4, smaller Station(unidentified R-4, rep.),3, m34°43.0'N, ajids, labeled 120°47.4'W, "Majid 91sp. m. 1," MMS

Description. Size to 11.5 mm (Rathbun, 1925). Carapace pyriform, convex, regions well defined;

I rostrum simple, slender; postorbital and supraorbital spines present. Basal article of antennae long and narrow, with spine at distal end. Anterointemal lobe of ischium of outer maxilliped large and strongly advanced;

merus subtriangular, the anterior margin the longest; outer, or principal, margin of next article nearly as long

I as outer margin of merus and almost straight until near distal end where it forms an elbow; terminal article unusually long.

Sexes markedly dissimilar. Rostrum a slender spine half as long as postrostral portion of carapace in

I male, shorter in female. Two median spines, one gastric, one cardiac; one long branchial spine. Postorbital spine small. Chelipeds very long, from three to four times length of carapace in male, only one and a half times

carapace length in female. A spine on first abdominal somite.

I Male: Carapace spinous; two spines on the median line, one onposterior part of the gastric region and

the other on the cardiac region; one long spine on the branchial region, with a small one in front of it and two

on the margin; a spine on the margin of the hepatic region and two very small ones arranged transversely on

gastric region; a spine on slender, spinulous on margins, onethe slender the orbital arch. Rostrum the about

half the length of the postfrontal portion of the carapace. Postorbital spine small, at some distance behind eye.

Abdomen bearing a spine on the first segment. Chelipeds nearly three times as long as carapace, granulate;

I merus one-spined above at the anterior margin; hand slender, slightly flattened vertically, increasing in width

toward the distal end; dactyl and pollex arched, gaping for one-half length. Ambulatory legs decreasing

regularly in length from the first to the fourth; fourth pair a little more than one-half the length of the first

I (Rathbun 1893).

i 107
U

Female: Carapace with two median spines, one on the posterior part of the gastric region and one on N

the cardiac region. Two spines on each branchial region and, almost in line with these, one on each protogastric

lobe, Lateral margins spinulous. Surface pubescent, rostrum slender, spinulous on the margins, curving upwards []

for its distal half. A prominent supraorbital spine. The first article of the abdomen carrying a spine; a slender l

spine above near the carpus; hand granulous; fingers nearly as long as palm, in contact. Ambulatory legs

slender, pubescent, decreasing slightly in length from the first to the fourth pair; dactyls spinulous below •

(modified from Rathbun, 1893 [ofAnasimus rostratus]). |

The length of the rostrum is subject to considerable variation. The females of the species, with their

shorter legs, are difficult to tell fromlnachoides laevis, but the ranges of these species only overlap in the Gulf n

of California (Garth, 1958:91).

Color in life: Cryptic, nondiscript tan carapace (Jensen, 1995:26, Plate 28). In alcohol, chelipeds and

legs conspicuously banded and marbled (Crane, 1937). n

[] Biology. This species has been recorded from sand, mud, rock, coral, and "coraUine" [algae], with

the majority of recordings (fully one third of the cases reviewed by Garth, 1958) coming from shell [debris] n

associated with sand. Velero III specimens came from depths ranging from 2 to 300 fathoms (4-554 m).

Ovigerous females have been recorded in every month of the year along the coasts of southern California and U

northern Baja California, although there is a seasonal trend with lows in the frequency curve in January, a rise

in April, and highs in August and September (Garth, 1958). n

I

Taxonomic Remarks. The rostrum for small females has never been drawn adequately; existing

figures do not depict adequate spination. Marked sexual dimorphism and variation within a given sex has lead !

to taxonomic confusion such that the different sexes were originally assigned to seperate genera. Specimens

are often covered with debris and attached material. U

Three MMS voucher lots bore the label "Majid sp. 1" and were not identified further. Collection data •

is the same for all vials, differing only in the "Rep:" field on the label. Of these, the lot mentioned above (Rep: |

3) contained three individuals, the largest of which is a small male E. spinosus. The other two specimens in

that lot are too undeveloped to allow confident identification. This is also the case with the other two lots []

(identical collection data as above, except that one vial is Rep: 1,the other Rep: 2), although the sole juvenile |

in the lot marked Rep: 2 (CL= 3.4 mm including strongly bifurcate rostrum), while clearly a majid, definitely

is not a young E. spinosus.

Type Locality and Type Specimens. Marked sexual dimorphism led to the sexes being treated by II

Rathbun (1893b) as separate species, the male assigned toErileptus spinosus, the female toAnasimus rostratus,

each with a designated holotype (see Garth, 1958). •

Type material: Holotype USNM 17341, male, carapace length = 10 ram, carapace width = 6 ram, off San

Diego, California, 36 fathoms (66 m), Albatross Station 2934. Female holotype (of Anasimus rostratus)

USNM 17340, northwest of Cedros Island, off Baja California, 58 fathoms (106 m). I

Distribution. Broadly distributed off southern California, from San Miguel Island and Point Santa

Barbara in the northern part of its range to Entrada Point, Magdalena Bay, Baja California, Mexico, and in

the Gulf of California from Rocky Point, Sonora, to Gorda Banks (Garth, 1958). A single specimen from n

south of the Ladrones Islands, Panama, was reported by Garth (1958). The present report extends the range

northward into the Santa Maria Basin.

!

!

!

108 I
I

I

I \

I A

I

I

I

•. ..:

!1

I! _ B

I

I

i

I

I Figure 2.25. Erileptus spinosus Rathbun, 1893: A, male; B, female (A and B after Rathbun, 1904, as Erileptus spinosus and Anasimus rostrams respectively).

!

!

; I 109
!

Family Pinnotheridae De Haan, 1833 !

Pinnixa occidentalis Rathbun, 1893 i

[]

Figure 2.26

Pinnixa occidentalis Rathbun, 1893:248, Fig. 12 (in part; not the specimen from San Diego); 1904:187, Plate I

7, Fig. 4; Plate 9, Figs. 6, 6a (in part; not specimens from Cape Fox); 1918:155, text Fig. 96, Plate

34, Fig. l.--Holmes, 1900:89.--Weymouth, 1910:56 text Fig. 3.--Schmitt, 1921:262, Fig. 156.-- m

Roesijadi, 1978:259 (larval physiology).--Hart, 1982:242, Fig. 101 (ecotogy).--Kozloff, 1987:417. 1

--Zmarzly, 1992:677, Fig. 1 (map), Fig. 12.

Pinnixa californiensis Rathbun, 1893:249.--Holmes, 1900:90.--Rathbun, 1904:187, Plate 7, Fig. 3.-- •

Weymouth, 1910:56 (in part; not young female from Pacific Grove). I

Material Examined. Two males, (USNM 071-BSS-01-TX, and SBMNH 018-BSS-01-TX) (one 1

from 91 m, no depth info on other specimen) [identification conf'urned by D. Zmarzly]; USNM Phase I i

primary voucher collection, 07 I-BSS-01-TX, 29 November 1983, one mature female, carapace length = 3.8

i

mm, carapace width = 7.7 mm; one juvenile, carapace length = 1.9mm, carapace width = 3.4; SBMNH Phase i

Isecondary voucher collection, 018-BSS-01 -TX, one male, (carapace length = 2.7 mm, carapace width = 5.2 i

ram), two females, three juveniles. i

Description, Large species to 19.5 mm carapace width, males, 20.5 mm carapace width, females ll

(Rathbun 1918). Carapace 2.0-2.2 times wider than long in adults, 1.6-1.9 times wider in juveniles. Dorsal I

surface pitted and highly irregular. Longitudinal median groove extending from frontal margin to mesogastric

region. Hepatic region delineated by grooves; cervical groove well developed, connection with deep gastric •

depression. Sharp transverse ridge in cardiac region; cardiac ridge formed of two crescent-shaped ridges, 1

appearing bilobed in male, not as acute or as bilobed in female. Anterolateral margin with granulate ridge.

Granules sometimes also present dorsolaterally on carapace, behind ridge. Ventral margin of anterolateral I

region granulate, with long plumose setae, i

Chelae robust, sexually dimorphic, distinctive among California species due to deflection and form of

fixed f'mger.In males, fixed f'mger short, about one-quarter length of palm, stout, and strongly deflexed; inner •

margin with large flat tooth proximally and small triangular tooth, occasionally worn fiat, distally. Dactylus l

becoming increasingly curved with age, strongly hooked in mature males; f'mgers gaping when closed. Dorsal

margin of dactylus smooth, sparsely setose; inner margin of dactylus typically toothless; some specimens with []

small triangular tooth at midpoint. Dactylar tooth usually present in small males and those regenerating 1

chelae; persisting in some mature males. Presence of dactylar tooth in mature males apparently unrelated to

size of animal or geographic occurrence. Dorsal margin of palm granulate; line of granules above ventral i

margin of palm on anterior face. l

Female chela more slender than that of male. Fixed finger longer, about 0.4 length of palm; obviously

deflexed, but less so than in male; inner margin paralleling ventral margin proximally; small triangular tooth •

at midpoint, followed by row of sharp transparent teeth that precede the upwardly curving distal tip. Dactylus i

of female not as strongly curved as in male; dorsal margin granulate, setose; inner margin with large triangular

tooth, occasionally worn flat, at midpoint, row of small transparent teeth between triangular tooth and distal II

tip, meeting row of similar teeth on opposing margin of fixed finger when fingers closed. Anterior face of i

propodus with several rows of granules running entire length just above ventral margin.

In adults, ambulatory legs fringed with plumose setae. Pereiopod 2 with ventral margin of merus _l

granulate to tuberculate; distal half of ventral margin of propodus with sharp spines; dactylus as long as

propodus, with longitudinal row of 4 or 5 spines. In ventral view, ischium with single tubercle on posterior

margin. On pereiopod 3, posteroventral margin of ischium with several small tubercles; dorsal and ventral I

110 i
I

I

I

,,/h _,,,,J'l ,,_ ",', ,,f,,_,"

l

Figure 2.26. Pinnixa occidentalis Rathbun, 1893: A, anterodorsal view of male; B, anterodorsal view of carapace

I face (juvenile of right male); gonopod C, anterior (A-E face fromofZmarzly, female cheliped; 1992). D, anterior face of male cheliped; E, abdominal

I

margins of merus and dorsal margins of carpus and propodus granulate to tuberculate; ventral margin of

I pandropodus 2 longitudinal with sharprowspines s of onshadis rptaslpinhaft. es onDactvyelus ntral asmargin. long as Onpropodus, pereiopodwith4,longi dorstudinal al and caventra rinal onmargins each sideof

merus heavily granulate to tuberculate; ventral margin of propodus bicarinate, carina serrate. Dactylus about

i as long as propodus, with 2 longitudinal cadnae laterally and 5 longitudinal rows of spines: 1 on dorsal

i tll
!

margin, 2 on ventral margin, and 2 laterally. On pereiopod 5, ischium and merus with granulate margins; I

dactylus as long as propodus. Dorsal and ventral margins of dactylus with rows of spines; setose. Distal tip of

dactylus of pereiopod 5 reaching will beyond distal edge of merus of pereiopod 4 when both legs extended, il

In specimens (greater than or equal to) 8 mm carapace width, ambulatory legs very slender; dactyls

also slender, cylindrical, nearly straight, as long as or longer than propodus on each leg. With maturity,

density of granules on alllegs increasing; size relationships of ambulatory legs changing dramatically; robustness •

of pereiopod 4 increasing allometrically

Male and female abdomen consisting of 7 free somites (after Zmarzly, 1992).

am

Biology. Found subtidaUy in burrows of the echiurid wormEchiurus echiurus alaskanus at northern !

extreme ofP. occidentalis range, also found free living (Hart, 1982; Zmarzly, 1992). On soft mud substrates

I

in Puget Sound (Lie, 1968). i

Taxonomic Remarks. Although southern California specimens tend to be slightly smaller than Alaska I

specimens (D. Zmarzly, pers. comm.), there appears to be little morphological difference between specimens

from these two areas. Hart (1982) cautioned that this may be a species complex in light of its wide geographic am

and bathymelric range. This was reitterated by Martin and Zmarzly (1994) when they recognized a new II

species in this complex (see below, P. scamit).

Type Locality and Type Specimens. South of Unimak Island, Alaska, 61 fathoms (112 m). Holotype ID

male, USNM 17474, carapace length = 9.5 mm; carapace width = 19.5 ram. I

Distribution. Alaska to Bahia Magdalena, west coast of Baja California, Mexico; intertidal to 238

fathoms (435 m) (Schmitt, 1921; Hart, 1982; Zmarzly, 1992). I

Pinnixa scamit Martin and Zmarzly, 1994

Figure 2.27 I

Material Examined. USNM 267500, and USNM 267501, see holotype and paratype below. I

II

Description. Carapace highly sculptured, with anterolateral ridge bearing pronounced and slightly

anteriorly-curved teeth; frontal margin with deep median cleft; cardiac ridge present, granular, not sharp. Ill

Fixed f'mger on major cheliped slightly deflexed; both fingers of major cheliped with single stout tooth located •

almost at midlength on inner margin. All ambulatory pereiopods with markedly serrate and setose dorsal and

i

especially ventral margins. Propodus of pereiopod 4 at least 2.5 times longer than wide (Martin and Zmarzly,

1994). I

Biology. Nothing is known.

Taxonomic Remarks. Most California species of Pinnixa White, 1846, have a ridge, which may be •

granular to tuberculate, along the anterolateral margin of the carapace. Exceptions include P.faba (Dana I

1851), P. longipes (Lockington 1876), P.tubicola Holmes, 1894, and P.weymouthi Rathbun, 1918, where an

anterolateral ridge is either absent or at best very weakly developed. Other exceptions are P.forficulimanus I

Zmarzly, 1992,P. minuscula Zmarzly, 1992, and P.littoralis Holmes, 1894, where the ridge is absent (Zmarzly I

1992). In the remaining California species of Pinnixa, this ridge is obvious in dorsal view, and sometimes is

as pronounced as in P.scamit. Juvenile stages of two species, P. occidentalis Rathbun, 1893, andP.franciscana II

Rathbun, 1918, sometimes exhibit acute teeth on the anterolateral ridge; however, this feature changes I

dramatically between the juvenile and adult stages. In addition, P.scamit has relatively slender legs with acute

marginal serrations as an adult, which is also characteristic only of juvenile stages in P. occidentalis and P. am

franciscana Thus P. scamit appears to have retained several "juvenile" characters in the adult stage (from I

Martin and Zmarzly, 1994).

!

112 I
iI

Figure 2.27. Pinnixa scamit Martin and Zmarzly, 1994: USNM 267500, and USNM 267501 (C only), western

i Santa Barbara Channel, California. A, female holotype, dorsal view of carapace and right side

appendages, pereiopods 1-3 drawn in situ, pereiopods 4 and 5 removed and figured separately (to

ensure correct proportions): B, same specimen, frontal view showing sculpturing of carapace and

subhepatic tooth (arrow): C, dorsal view of immature paratype; note that subhepatic tooth (arrow)

i ofandholotype, acute serrations left side,along outer anterolateral view; E, abdomen marginofareholotype, already ventral present view at thisillustrated size: D, third in situmaxi (firstlliped two

segments not visible, and segments not visible, and segment 3 shortened due to curvature of abdomen).

n Scale bar = 2.0 mm for A-C, E; 1.0 mm for D (from Martin and Zmarzly, 1994).

I

n 113
!

All other California species are easily distinguished from P. scamit by the length:width ratio of the I

propodus of pereiopod 4. In true P occidentalis Rathbun, the species to which P. scamit appears to be most

closely allied, the propodus of P4 is distinctly broader, nearly as broad as long. In contrast, in P. scamit the I

propodus of pereiopod 4 is approximately 2.6 times longer than wide. II

Type I.aw.alityand Type Specimens. MMS station data 071-B SS-01-TX, 34°29.040'N, 120°44.013"W,

western Santa Barbara Channel, just seaward of, and SSW of, Pt. Arguello, California; 311 m. Holotype •

female USNM 267500, carapace width 7.4 rnm, carapace length 3.7 mm 29 Nov 1983. Paratype juvenile,

USNM 267501 (sex indeterminate), carapace width 3.4 nun, carapace length 2.1 ram, same collecting data as

holotype (Martin and Zmarzly, 1994). I

Distribution. Presently known only from the type locality.

Acknowledgments I

We are especially grateful to the members of the Southern California Association of Marine Invertebrate I

Taxonomy for providing additional information and some unpublished data. Their assistance in compiling

this report, especially that of Don Cadien and Jim Roney, who also very kindly reviewed parts or all of the i

report, can not be overemphasized. At the same time, and because not all comments or suggestions could be I

accommodated, we take responsibility for the resulting product, and any existing errors are ours alone. We

also thank Mary Wicksten for reviewing some of the sections dealing with caridean shrimp, and Ron Velarde E

for his input and additional observations, often by way of Don Cadien. |

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120 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°05.3YW 98

2 35%7.70"N 121°06.52"W 200

3 35°27.07'N 121° 10.2(YW 291

4 35°26.56'N I21°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

1l 35°17.80'N 121°22.13'W 690

12 35°15.03'N 120057.3 I'W 98

t3 35°14.54'N 120°59.77'W 197

14 35°14.15'N 121°02.04'W 299

15 35°13.98"N 121°04.54"W 393

16 35°12.23'N 121°16.29'W 591

17 35°11.61"N 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

2 l 35006.1 I'N 120°44.82'W 49

22 35°05.85'N 120°50.23"W 99

23 35°05.60'N 120055.18"W 195

25 35°05.07'N 121°00.75"W 390

26 35°04.38'N 121°15.99"W 590

27 35°04.30'N 121°19.27'W 611

28 35°04.22'N 121°19.65'W 603

30 34°54.19'N 120°47.07"W 98

31 34°53.76'N 120°52.96'W 200

32 34°53.56"N 120°56.81"W 297

33 34°53.43"N 120°59.66'W 396

34 34053.15"N 121°04.40'W 492

35 34°52.96'N 121o10.30"W 548

36 34°52.77"N 121o15.37"W 492

38 34049.8 I'N 120052.66'W 197

39 34°49.53'N " 120°56.85'W 294

40 34°49.24'N 121°00.8 I'W 392

41 34°48.35'N 121°19.14"W 495

42 34°48.04"N 120°47.50'W 100

43 34°46.59"N 120°52.92"9/ 197

45 34044.9 I'N 120°59.59'W 395

46 34°41.22'N 121°13.56'w 597

47 34°41.99"N 121°10.81'W 378

48 34045.1 I'N ] 20°52.85"W 196

49 34°45.03'N 120°56.3 I'W 290

50 34°37.80'N 121°01.66"W 591

52 34°39.5CN 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

121
I

Table A.1 (Continued)

I

Station Latitude Longitude Depth I

(m)

!

58 34°34.35'N 120°45.18'W 99 m

l

59 34°33.65'N 120°47.18'W 216

60 34°33.25'N 120°48.34'W 275

61 34°33.0 I'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 I

64 34°33.15'N 120°40.90"W 59

65 34°31.27'N 120°43.27'W 107

66 34°30.46'N 120°44.55'W 201 •

67 34°30.29'N 120°45.50'W 282

68 34°29.24'N 120°45.99"W 390

69 34°22.88'N 120°54.20'W 927 I

l

70 34°29.67'N 120°43.70'W 200

71 34°29.04'N 120°44.0 I'W 306

72 34°28.4 I'N 120°44.76'W 401

73 34°28.2 I'N 120°36.80"W 98 m

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 I

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 m

81 34°21.26"N 120°28.83"W 294 •

82 34 °18.7 I'N 120°29.55'W 394 I

83 34 °17.20'N 120°30.2tYW 444

84 34 °13.54'N 120°31.19'W 394 I

1

85 34°25.88'N 120°16.3 I'W 113

86 34°24.45"N 120°17.02'W 197

87 34°21.6tYN 120°17.11"W 299

88 34 °17.89'N 120°16.86"W 393 •

l

89 34 °13.79"N 120°16.56"W 471

90 34°09.44"N 120°16.30'W 375

91 34 °11.73"N 120°07.43"W 540

92 34°08.70"N 120°07.50"W 444 •

93 34°07.63'N 120°07.5 I'W 357

96 34°22.9 I'N 120°05.42"W 296

94 34°24.54"N 120°05.47"W 96 m

95 34°23.70"N 120°05.47'W 198 •

97 34°22.28'N 120°05.49"W 393 I

98 34°12.87"N 120°05.59"W 561

99 34°11.22"N 120°05.86"W 540 m

I

100 34°08.67"31 120°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 •

I

104 34°59.45"N 120°56.49'W 294

105 34°59.23'N l20°59.60"W 392

106 34°58.95"N 121°04.42'W 492

107 34°58.65"N 121°15.08"W 573 •

I

108 34°58.21'N 121°17.88'W 492

Note: Sample labels from the Soft-substrate stations have several identification codes which include a station number, sample I

replicate number, and analysis type. These are as follows: 001 to 200 = the range of station numbers; BSS = Benthic •

Sediment Single (i.e., a non-replicated station); BSR = Benthic Sediment Replicate (three replicates taken at this station); BSV =

I

Benthic Sediment Variance (suhsamples); 01-09 = replicate numbers; TX = a taxonomy sample. Sample labels having the

designation BRA, represents a sample from rocks taken as part of the hard bottom survey. I

I

122 I
I

I ". _ _ _.ES'rERO I I

".2,,_,_

I

I _. _

s '"7 _ x _ CALIFORNIA ,-

': _o..._.; ( _.=.o,,

;.,,- \

19 ",18 _ 35o10"N i.

• Q,. ]_. R1 "_

I _- "., • " I

27 26

oe • • ,

28 ,,...,= 104 103 '. 102.

I 108 107 _e • • al 0 •

• • 105 ! OR8 J

35 34 " 32 31! _ 30" (PbSAL

40 39 38 o '_

4x * * :

• 4.3 42 [

n 45• * 0,':48 ° R4 (_LPUFIISIMA PI.

IOr'zn 46 47 49 i ,,R.5 • " ")

"--.':Y "-, • • R6 T 52

I .., 50 :. •

' _ " :" ". 6,_L_"_"_'_°

i '-, ......... 56 • 6,_."_.....

:" - _s o_

................... • ....... • 67 -- v 1L

: 62 . .,*e? _ _ _ I% CONCEPTION

I 63 '..e 72 71.......... .. e'/4 O65

........ : • ..... . 94

69.-' S6 75 "... 079 ..... .If6................. t9 _

• .........eso........ 200 --_o-....

-:. o_ 097

i -17 .sl

o78 082 • 88

rnI O83

089

i , 084 098 i 91 •

-- 34 ° 10'N ,' • _

/ KEY 090 9"2.• olCO

n "

- 93 • • I01

...' = Sit=S_=_t'_¢_oJ_-i_ (]'ha_ a)

I Figure A.1. Map showing location of soft-substrate stations from the Phase I Reconnaissance and Phase II

Monitoring Programs.

I

I

i 123
!

Table A.2. Location of soft-substral¢ stations taken during the Phase II Monitoring Program. i

m

Station Latitude _n_mde _p_ •

(m) []

i

R-1 35°05.83_ 1_°49.16_ 91 []

R-2 35°05.5_N 1_°53.4_W 161 i

R-3 35°05.3_N 121°_.9_W _9

R-4 34°43.01'N 120°47.39'W 92 i

R-5 34°42.69"N 120°50.83'W 154 | R-6 34041.40"N 120°57.90'W 410

R-7 34°52.90"N 121°10.3tYW 565

R-8 34°55.30_ 120°45.87_ _ •

R-9 34°53.68_ 120°59.12_ 410 II

PJ- 1 34°55.79'N 120049.9 )_ 145

PJ-2 34°55.32'N 120°49.59'W 142

PJ-3 34°56.26'N 120°49.58'W 138 •

PJ-4 34°56.26'N 120°50.24'W 150

PJ-5 34°55.32'N 120050.24'W 152

PJ-6 34054.7 I'N 120049.91 'W 148 i

PJ-7 34°55.79'N t20°48.60"W 123 •

PJ-8 34°56.87'N 120°49.9 I'W 142 J

PJ-9 34°55.79'N 120°51.23'W 169

PJ-10 34°53.63'N 120°49.91'W 147 i

PJ-11 34°57.95'N 120°49.91'W 136 !i PJ-12 34°55.58'N 120°49.9 I'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 •

Hi PJ- 15 34°55.79'N 120°50.57'W 155

PJ- t6 34°55.03'N 120°48.99"W 130

PJ-17 34°56.56'N 120°48.98'W 126

PJ-18 34°56.56'N 120°50.84'W 158 •

PJ- 19 34°55.03'N 120°50.84'W 167

PJ-20 34°50.38'N 120°49.9 I'W 148

PJ-21 35°0 t.23'N 120051.15"W 143 I

PJ-22 34°55.25'N 120°49.93'W 143 •

PJ-23 34°56.33'N 120°49.90'W 143 I

!

|

Table A.3. Sampling dates of MMS Phase II Monitoring Program.

Cruise Date I

1-1 October 1986

1-2 January 1987

1-3 May 1987 •

2-1 July 1987 II 2-3 October 1987

2-4 January 1988

2-5 May 1988 I

3-1 October 1988

3-4 May 1989 !1

I

,24 i
Table A.4. MMS Phase I - Locations of hard-substrate Iransects.

Station Beginning End Depth

Latitude Longitude Latitude Longitude (m)

l A/B 34°24.454 'N 120°01.876'W 34°24.464'N 120°00.878'W 69-73.5

t 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 110-126

2 C/D 34°10.984"N 120°28.09_'W 34° 10.780'N 120°27.554"W 120-123

4 A/B 34°27.539'N 120°40.364'W 34°28.162"N 120°40.189"W 168-237

6 A/B 34°30.246'N 120°35.555'W 54-63

6 C/D 34°30".42 I'N 120°3"4.315'W 54-63

13 A/B 34°42'.570'N 120°4'7.899'W 34°42.107'N 120°48.253'W 92-100

13 C/D 34°42.556'N 120°48.147'W 34°42.974'N 120°47.424'W 88.5-100.5

I4 A/B 34°43.589"N 120°49.093'W 34°42.826'N I20°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°45.912'1'4 120°49.726'W 91.5-123

17 A/B 34°49.382'N 120°50.768"W 34°49.60(YN 120°50.688"W 160.5-168

19 A/B 34°47.833"N 120°51.425'W 34°47.097'N 120°50.793"W 148.5-177

20 A/B 34°46.470'N 120°50.289"W 34°46.140"N 120°49.885"W 90-130.5

21 A/B 34°47.335'N 120°45.903'W 34°47.548'N 120°46.123"W 75-90

22 A/B 34°50.365'N 120°48.221'W 34°50.990'N 120°48.365'W 114-115.5

23 A/B 34°49.868'N 120°47.393"W 34°50.003'N 120°47.480"W 93-102

25 A/B 35°05.662'N 120°47.562"W 35°06.036'N 120°47.652'W 64.5-72

26 C/D 35°11.586'N 120°55.556'W 35°11.555'N 120°55.233"W 108-111

27 A/B 35°20.906'N 120°59.657'W 35°21.035'N 120°59.603'W 96-126

28 A/B 35°21.539'N 120°59.64 I'W 35°21.867"N 120°59.299'W 96-105

29 A/B 35°27.864'N 121°05.33 I'W 35°27.805"N 121°05.277'W 102-106.5

Table A.5. MMS Phase II - Locations of hard-substrate photosurvey stations.

Station Latitude Longitude Depth

(m)

PH-E 34°30.26'N t 20°42.76"W 119

PH-F 34°30.8 I'N 120°42.36'W 105

PH-I 34°29.96'N 120°41.68'W 107

PH-J 34°29.82'N 120°41.82'W 117

PH-K 34°29.37'N 120°42.26'W 160

PH-N 34°29.21 'N 120°42.05 'W i 66

PH-R 34029. II'N i20°42.67"W 213

PH-U 34°31.48'N 120°43.5 I'W 113

PH-W 34031.52"N 120°45.86'W 195

125
I

',, _. ESTERO I I I

_ _ CALIFORNIA I

• _N _ 35°10"N " I

I

I

::. ...,.._ I

_o_v. _-u ..... _ • k'_--_RGUELLO

• _-,,_ _'_ I

P_PW "_ / '"" • 2 ..............

"" / ................... 200 ...........

_-E • _-I __,

34°30' N PH-J _ i I

'

I

34o28,N I

120 °44'W

I

Figure A.2. Map showing location of hard-substrate stations from the Phase ] Reconnaissance and Phase II

Monitoring Programs. Phase II stations are indicated in the inset. I

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126 I
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I

The Department of the Interior Mission

I As the Nation's principal conservation agency, the Department of the Interior has responsibility

for most of our nationally owned public lands and natural resources. This includes fostering

sound use of our land and water resources; protecting ourfish, wildlife, and biological diversity;

I preserving the environmental and cultural values of our national parks and historical places; and providing for the enjoyment of fife through outdoor recreation. The Department assesses

our energy and mineral resources and works to ensure that their development is in the best

interests of aB our people by encouraging stewardship and citizen participation in their care.

I The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.

The Minerals Management Service Mission

I (__ As a bureau of the Department of the Interior, the Minerals Management Service's (MMS)

primary responsibilitiesare to manage the mineral resources located on the Nation's Outer

Continental Shelf (OCS), collect revenue from the Federal OCS and onshore Federal and Indian

i lands, Moreover and distribute those revenues.

, in working to meet its responsibilities, the Offshore Minerals Management Program

administers the OCS competitive leasing program and oversees the safe and environmentally

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 ensuringthe

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 responsive MMS strives to the topublic's fulfill itconcerns s responsibiliti and interests esthroughby the maintaining general guiding a dialogue principl withesallopotentially f: (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.
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