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I OCS Study

MMS 96-0002

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| TAXONOMIC ATLAS OF THE BENTHIC FAUNA OF

THE SANTA MARIA BASIN AND

WESTERN SANTA BARBARA CHANNEL

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I INAL REPORT Volume 9 of 14

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The Mollusca Part 2 m Gastropoda

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I Edited by: Paul H. Scott

James A. Blake

I ndrew L. Lissner

Submitted by:

I Science Applications International Corporation

10260 Campus Point Drive

San Diego, California 92121

I For:

I UMinerals .S. DepaMrtment of the Interior

anagement Service

Pacific OCS Region

770 Paseo Camarillo

I Camarillo, California 93010

Under Contract No. 14-35-0001-30484

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Minerals Management Service March 1996

I Pacific OCS Region

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DISCLAIMER i

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

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

nece not consiti ssarilyureflect the view of the Minerals Management Service. Mention of trade names does i

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

conformity with Minerals Management Service editorial standards.

TAXONOMIC DrscLAIMER I

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

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

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PROJECT ORGANIZATION

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PROGRAM MANAGER

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I DR.Aromsw L. Llss_a_ Science Applications International Corporation

10260 Campus Point Drive I San Diego, California 92121

I DEPUTYPROGRAMANAGER

I DR. JAMESA. BLAKE

ENSR Consulting and Engineering

I 89 Water Street

Woods Hole, Massachusetts 02543

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I PROmCTTECHNICALOFFICER

MR. FRANKJ. MANAC_.,O

i U.S. Department of the Interior

Minerals Management Service, Pacific OCS Region

I 770 Paseo Camarillo

Camarillo, California 93010

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Com'l_Ur_G Atrmoms

I DR. JAMESH. MCLEAN

DR. TEaRF_CEM. Gos_

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ACCESS NUMBER: 30484.9

TECHNICAL SUMMARY i

STUDY TITLE: TaxonomicAtlas of the Santa MariaBasinandWesternSanta BarbaraChannel.

REPORTTITLE: TaxonomicAtlasof the SantaMariaBasinandWesternSanta BarbaraChannel.Volume9. i

The Mollusca Part2 -- Gastropoda.

BB

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

SPONSORING OCS REGION: Pacific. I

APPLICABLE PLANNING AREA: Southem California.

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

COMPLETION DATE OF REPORT:March 1996.

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

CUMULATIVE PROJECT COST:$724,384. •

PROJECT MANAGER:Andrew L. Liasner.

AFFILIATION:Science Applications InternationalCorporation. i

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

i

PRINCIPALINVESTIGATORS" James A. Blake, Andrew L. Lissner. i

KEY WORDS: Marine invertebrates,Mollusca,Gastropoda,Prosobranchia,Opishtobranchia, taxonomy, n

California, Santa Maria Basin,Santa BarbaraChannel,continentalshelf. I

BACKGROUND: The TaxonomicAtlasof the Santa Maria Basinand westernSanta BarbaraChannel isan

extension ofthe benthicreconnaissance(PhaseI)andmonitoringprograms(PhaseII) that wereconducted i

by the MMSsince 1983.The organismsthatwerecollectedas partofthose programsprovidethe materialon

which the Atlas isdeveloped.Inordertofully documentthefaunacollectedby those programs,a seriesof 14

volumeswillbe preparedthat providekeys,descriptions,and illustrationsofthe benthic faunaofthe hardand •

soft substrate environments. A team of 40 experts onthe fauna has beenassembled to carry out this work I

and their contributionsare distributedamong the 14volumes.

OBJECTIVES:The objectivesofVolume9 aretoprovidekeys,descriptions,and illustrationsto the gastropod I

mollusksofthe SantaMaria BasinandwesternSanta BarbaraChannel. IB

DESCRIPTION: This compilationof the marinegastropodfauna of southernCalifornia is the first major •

treatmentoftheprosobranchgastropodssinceMcLean(1978)andfortheopisthobranchfaunasinceMcDonald

and Nybakken(1980). The presentworkconsistsofan introduction,laboratoryand fieldmethodology,a

glossary,and identification keys for both gastropodgroups. Fortysix families and 81 generaare diagnosed. IB

One of the genera is identified as new and here described for the first time. A total of 154 species are I

illustrated in photographsor line drawings. Of theseseven species are identifiedas new to science and are

here described for the firsttime.

BI

SIGNIFICANT CONCLUSIONS:The keys,detaileddescriptions,and illustrationsto the 154speciestreated I

inthisvolumerepresenta majorcontributiontogastropodmollusksystematicsfortheeasternPacific.Many

poorlyknownspeciesare newlydefinedwithnewillustrationsandphotographsT. he prosobranchgastropods i
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ACCESS NUMBER: 30484.9

I are placedina modemclassificationforthefirsttime. One newgastropodgenusand sevennew speciesare

described.

i STUDY RESULTS:The fauna Santa Maria Santa Channelis

of the Basinand western Barbara evaluated

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

sediments using box cores and from rocks using manipulatorarms of submersibles and remotely operated

I vavailabl ehiclese.Ttohethecolteamlectioonfsinvestig are orgaani tors. zeAdditiona dinto setsmloaterial f vouchfromersfrothembulk Phascollections e I and tl now that hav archived e beewith n mathede

NaturalHistoryMuseumof LosAngelesCountyisalsobeingexamined.Thetotalnumberofspeciestreated

I inthe entireTaxonomicAtlas(14 volumes)mayexceed 1,000species.

A totalof 154speciesin81 generaand46 familiesaretreatedas partof volume9. Thescopeof thisvolume

mira includesmaterialsfromtheMMS surveysaswellasadditionalmaterialsfromtheeasternPacificaccumulated

, bytheauthors.The authorshavealsoexaminednumeroustypespecimensof mostspeciesand alongwith

thecollectionstreatedhaveproduceda monographthatwillbe applicableto theentiremarineenvironment

of westernNorthAmedca. tn thisregard,thevolumebecomesthe mostimportantcompilationof data on

I contribution, northeasternthPaci is sfitudy c gasistropods a valuable in thcontribution e last two dtoecaunderst des. Inandaddingitithe on tdistribution o being anand impozoortangeography t taxonomicof

gastropods in the eastern Pacific. The ranges of many species are extended; others restricted. One new

i genusand sevennew speciesof prosobranchgastropodsarehere describedfor the first time.

STUDY PRODUCT:Scott, P. H.,J. A. Blake, andA. L. Lissner (Editors). 1996.TaxonomicAtlas of the Santa

Maria Basin and Western Santa Barbara Channel. Volume 9: The Mollusca Part 2 -- Gastropoda. A final

I rMinerals eportpreManagement paredby ScienceApplicationsInternationalCorporation,for the U.S. Department of the Interior, Service, PacificOCS Region,Camarillo,CA. OCS Study MMS 96-0002. Contract No.

14-35-0001-30484.

I

I "P.rsaffiliationmaybedifferentthanthatlistedforProjectManager(s).

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ACCESS NUMBER: 30484.9

III i IIO e I17 i I

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REPORTDOCUMENTATION1._.POnTNO. MMS 96-0002 2. _._c_l_t's Acc,s,_ m_

PAGE

4. T_ and Subtitle 5, Report Date

TaxonomicAtlasoftheSantaMariaBasinand WesternSantaBarbaraChannel. Volume March 1996

9. The MolluscaPart2 m Gastropoda. ¢,.

7. Author(s) 8, Porformln8 Organization Rapt, No.

PaulH. Scott,JamesA. BJakeand AndrewL. Lissner(editors)

; 9. Pen_rrntltll Orl_nizltion Name and Address 10, Projectrresk/Work Unit No.

ScienceApplicationsInternationalCorporation z_.Cont_,c_orc)Gr,._(mNo.

10260 Campus Point Drive

San Diego,California92121 (c_ 14-35-001-30484

(G)

112. SDonsorlnG Orl[anizatlon Nsme and Address 13. Type of Report & Period Covered

Final 9 of 14 volumes,

MineralManagementService,U. S. Departmentofthe Interior, 9/89 to 3/96 PacificOCS Region,770 Paseo Camarillo

Camarillo,California 93010 _4.

15. _pl_lermmts, l'y Nots, s

Contributingauthors: Dr. JamesH. McLeanand Dr.TerrenceM. Gosliner

Thiscompilationof the marinegastropodfauna of southernCaliforniaisthe first majortreatment oftheprosobranch

gastropodssinceMcLean(1978)andfortheopisthobranchfaunasinceMcDonaldandNybakken(1980). The present

workconsistsofanintroduction,laboratoryandfieldmethodology,a glossary,andidentificationkeysforbothgastropod

groups.The scopeofthisvolumeincludesmaterialsfromthe MMS surveysas wellas additionalmaterialsfrom the

eastem Pacificaccumulatedby the authors. The authorshave also examinednumeroustypespecimensof most

speciesandalongwiththe collectionstreatedhaveproduceda monographthatwillbeapplicableto theentiremarine

environmentof westernNorthAmerica.In thisregard,thevolumebecomesthe mostimportantcompilationof dataon

northeasternPacificgastropodsinthelasttwodecades. Inadditionto beingan importanttaxonomiccontribution,this

studyis a valuablecontributionto understandingthe distributionand zoogeographyof gastropodsin the eastern

Pacific. The rangesof manyspeciesareextended;othersrestricted.Fortysixfamiliesand81 generaare diagnosed.

One ofthe generais identifiedas newand here describedforthe firsttime. A total of 154 speciesare illustratedin

photographsor linedrawings. Ofthesesevenspeciesareidentifiedas newto scienceandare heredescribedfor the

firsttime.

17. Oocume.t krml_ s. Descriptors

Marine invertebrates, Mollusca, Gastropoda, marine snails,taxonomy, California,Santa Maria Basin, Santa

BarbaraChannel,continentalshelf.

k Ich._lflem/Open-Ended Tm

TaxonomicAtlas,Keysto species,faunalguide,marineecology,Mollusca,Gastropoda,marinebenthos.

I ¢. CO_kT! FhddiGroup

18. Avaibibil(t'y Statement 19. Security C/ross (This Report) 21. No. of PalreS

I Unlimited Unclassified 219

2o.S*cur_¢1=,(Thb_s) _. I=ric=

See ANSI-Z39.18) See Insffui:*lons on Reverse OPTIONAL FORM 272 (4-71)

i (Formerly NTIMS)

Doll)srtment Of.Corrdmerce
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Table of Contents I

1. Tm_ PROSOnRANCmA(by James H. McLean) I

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

Laboratory Methods ........................................................................................................................ 1 I

Scope ............................................................................................................................................... 2

Conventions ..................................................................................................................................... 3 I1

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Glossary ........................................................................................................................................... 5

List of Prosobranch Species ............................................................................................................ 8

Key to Families ............................................................................................................................. 10 1

Descriptions of Species ................................................................................................................. 13 1

Superfamily Acmaeoidea ........................................................................................................ 13

Family Lepetidae Gray, 1850 ........................................................................................... 13 1

lothia lindbergi McLean, 1985 .............................................................. 14 1

Family Lotfiidae Gray, 1840 ............................................................................................ 15

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Niveotectura funiculata (Carpenter, 1864) ............................................ 15 Ill

Superfamily Scissurelloidea ................................................................................................... 16 1

Family Scissurellidae Gray, 1847 .................................................................................... 16

Subfamily Anatominae McLean, 1989 ...................................................................... 16 1

Anatoma crispata (Fleming, 1832) ........................................................ 17 1

Superfamily Fissurelloidea ..................................................................................................... 18

Family Fissurellidae Fleming, 1822 ................................................................................. 18 I

Key to species of Fissurellidae ............................................................................ 18 1

Subfamily Emarginulinae Gray, 1834 ....................................................................... 19

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Scelidotoma bella (Gabb, 1865) ............................................................. 19 1

Cranopsis cucullata (Gould, 1846) ........................................................ 21 1

" Puncmrella cooped Carpenter, 1864 ..................................................... 22 m

Puncturella punctocostata Berry, 1947 ................................................. 22

Superfamily Trochoidea ......................................................................................................... 23 1

Family Turbinidae Rafinesque, 1815 ............................................................................... 23

Key to species of Turbinidae ............................................................................... 23

Subfamily Liotiinae H. & A. Adams, 1854 ............................................................... 24 n

Macrarenefarallonensis (A. G. Smith, 1952) ....................................... 24

Subfamily Colloniinae Cossmann, 1916 ................................................................... 26

Homalopoma draperi McLean, 1984 ..................................................... 26 1

Homalopoma paucicostatum (Dall, 187I) ............................................. 27

Homalopoma berryi McLean, 1964 ....................................................... 27

Homalopoma cordellensis McLean, new species ................................ 28 n

Family Trochidae Rafmesque, 1815 ................................................................................ 29 II

Key to species of Trochidae ................................................................................ 29

Subfamily Calliotropinae HAckman and McLean, 1990 ............................................ 30 I

Bathybembix bairdii (Dall, 1889) .......................................................... 30 1

Cidarina cidaris (Carpenter, 1864) ........................................................ 32

Subfamily Caltiostomatinae Thiele, 1924 ................................................................. 32 •

Calliostoma turbinum Dall, 1896 ........................................................... 33

Calliostoma variegatum Carpenter, 1864 .............................................. 34

Calliostoma platinum Dall, 1890 ........................................................... 34 •

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! Subfamily Solariellinae Powell, 1951 ....................................................................... 35

Solariella peramabilis Carpenter, 1864 ................................................. 36

m Solariella nuda Dall, 1896 ..................................................................... 37 Subfamily Halistylinae Keen, 1958 ........................................................................... 37

Halistylus pupoideus (Carpenter, 1864) ................................................. 37

Superfamily Cerithioidea ........................................................................................................ 39 Family Cerithiidae Fleming, 1822 ................................................................................... 39

Key to species of Cerithiidae .............................................................................. 39

Subfamily Bittiinae Cossmann, 1906 ........................................................................ 39

n Lirobittium rugatum (Carpenter, 1864) ................................................. 40

Lirobittium lomaense (Bartsch, 1911) ................................................... 41

LirobittiumfetelIum (Bartsch, 1911) ...................................................... 42

I Lirobittium larum 191 42 (Bartsch, l)

Lirobittium paganicum (Dall, 1919) ...................................................... 43

Family Turritellidac L6ven, 1847 .................................................................................... 43

I Turritella Carpenter, 1864 44 cooperi

Superfarnily Rissoidea ............................................................................................................ 45

Family Rissoidae Gray, 1847 ........................................................................................... 45

I Subfamily Rissoinae Gray, 1847 45 Key to species of Rissoidae ................................................................................. 45

Alvania rosana Bartsch, 1911 ................................................................ 45

AIvania tumida Carpenter, 1857 46 Family Vitrinellidae Bush, 1897 ...................................................................................... 46

Vitrinella eschnaurae Bartsch, 1907 ...................................................... 47

I Family Caecidae Gray, 1850 ............................................................................................ 48 Caecum crebricinctum Carpenter, 1864 ................................................ 49

Family Vanikoridae Gray, 1840 ....................................................................................... 49

I Key to species Megaloof Vanikoridae ............................................................................ 50 mphalus californicus (Dall, 1903) ............................................. 50

Megalomphalus sehmiederi McI.e.an, new species ............................. 51

I SupeFami rfamily Calyptraeiodea ................................................................................................... 51 ly Calyptraeidae Lamarck, 1809 .............................................................................. 51

Key to species of Calyptraeidae ......................................................................... 52

n Crepidula Crepipatellaglottidiarum dorsata (Broderip, Dall, 19051834) .........................................................

.................................................. 5254

Crepipatella orbiculata (Dall, 1919) ..................................................... 54

I Superfamily Family Ovulidae CypraeoidFleming, ea ....................................................................................................... 1822 ...................................................................................... 5555

Key to species of Ovulidae ................................................................................. 56

n Neosimnia Neosimnia loeb barbarensis eckiana (Dall, (Weinkauff, 1892) ...................................................... 1881) ........................................... 56

Superfamily Lamellarioidea ................................................................................................... 58

n FamilySubfamily TriviidaeTriviinae Troschel,Troschel, 1863 ...................................................................................... 1863 .......................................................................... 5858

Trivia ritteri Raymond, 1903 ................................................................. 58

U Superfami Familyly Nati Naticidae coideaForbes, .......................................................................................................... 1838 ........................................................................................ 5959

Key to species of Naticidae ................................................................................. 59

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Subfamily Polinicinae Finlay & Marwick, 1937 ....................................................... 59 •

Euspira pallida (Broderip & Sowerby, 1829) ....................................... 60

Calinaticina oldroydii (Dall, 1897) ....................................................... 61 •

Subfamily Naticinae Forbes, 1838 ............................................................................ 62 l

Cryptonatica affin is (Gmelin, 1791) ...................................................... 62

Superfamily Janthinoidea ....................................................................................................... 63 !

Family Epitoniidae Berry, 1910 ....................................................................................... 63 I

Key to species of Epitoniidae .............................................................................. 63

Epitonium berryi (Dall, 1907) ................................................................ 64 n

Epitonium hindsii (Carpenter, 1856) ...................................................... 65 Ii

Epitonium indianorum (Carpenter, 1864) .............................................. 66

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Epitonium sawinae (Dall, 1903) ............................................................ 67

Epitonium lowei (Dall, 1906) ................................................................. 67 II

Nodiscala spongiosa (Carpenter, 1864) ................................................. 68

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Supeffamily Eulimoidea ......................................................................................................... 69

Family Eulimidae Troschel, 1853 .................................................................................... 69 II

Key to species of Eulimidae ................................................................................ 69

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Eulima raymondi Rivers, 1904 .............................................................. 70 i

Pseudosabinella baked (Bartsch, 1917) ................................................ 71 II

Balcis micans (Carpenter, 1864) ............................................................ 72 el

BaIcis oldroydae (Bartsch, 1917) ........................................................... 73

PoIygireulima rutila (Carpenter, 1864) ................................................. 74 I[

Vitreolina columbiana (Bartsch, 1917) .................................................. 75 n

VitreoIina macra (Bartsch, 1917) .......................................................... 76

Vitreolina yod (Carpenter, 1857) ........................................................... 76 i

Haliella abyssicola Bartsch, 1917 ......................................................... 77

Superfamily Muricoidea ......................................................................................................... 78

Family Muricidae Rafinesque, 1815 ................................................................................ 78 n

Key to species of Muricidae ................................................................................ 78

Subfamily Ocenebrinae Cossmann, 1903 .................................................................. 80

Ocinebrina lurida (Middendorff, 1848) ................................................ 80 1

Ocinebrina barbarensis (Gabb, 1865) ................................................... 81

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Austrotrophon catalinensis (Oldroyd, 1927) ......................................... 82

Subfamily Trophoninae Cossmann, 1903 ................................................................. 83 l

Boreotrophon triangulatus (Carpenter, 1864) ....................................... 84

Boreotrophon multicostatus (Eschscholtz, 1829) .................................. 85

Boreotrophon bentleyi DaU, 1908 .......................................................... 85 n

Boreotrophon avalonensis Dall, 1902 ................................................... 86 []

Boreotrophon eucymams Dall, 1902 ..................................................... 86

Boreotrophon apolyonis (Dalt, 1919) .................................................... 87 •

Boreotrophon raymondi (Moody, 1916) ................................................ 87

Boreotrophon keepi (Herflein & Strong, 1937) ..................................... 89

Boreotrophon kabati McLean, new species ........................................ 89 •

Boreotrophon tolomius (Dall, 1919) ...................................................... 90 II

Boreotrophon hazardi McLean, new species ...................................... 91

Boreotrophon pedroanus (Arnold, 1903) .............................................. 92 •

Boreotrophon stuarti (E. A. Smith, 1880) ............................................. 92 !

Scabrotrophon maltzani (Kobelt & Ktister, 1878) ................................ 94

Scabrotrophon cerritensis (Arnold, 1903) ............................................. 94 •

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n Scabrotrophon lasius (Dall, 1919) ......................................................... 96

Scabrotrophon grovesi McLean, new species ..................................... 96

n Scabrotrophon Ocenotrophon painei clarki (McLean, Dall, 1903)new......................................................... species ........................................ 9798

Family Turbinellidae Swainson, 1835 ............................................................................. 99

i Key to species of Turbinellidae .......................................................................... 99 Subfamily Ptychatractinae Stimpson, 1865 ............................................................... 99

Exilioidea rectirostris (Carpenter, 1864) ............................................. 100

i Exilioidea kelseyi (Dall, 1908) ............................................................. 101 Family Buccindae Raf'mesque, 1815 .............................................................................. 102

Key to species of Buccinidae ............................................................................ 102

Neptunea amianta (Dall, 1890) ............................................................ 102 i

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1 Neptunea tabu/am (Baird, 1863) .........................................................

103

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Family Nassariidae Iredale, 1916 ................................................................................... 103

Key to species of Nassariidae ........................................................................... 104

I Nassarius (Hinds, 1844) 104 perpinguis

Nassarius rhinetes Berry, 1953 ............................................................ 105

Nassarius insculptus (Carpenter, 1864) ............................................... 106

I Family Fasciolariidae Gray, 1853 107 Fusinus barbarensis (Trask, 1855) ...................................................... 107

Family Columbellidae Swainson, 1840 ......................................................................... 108

I Key to species of Columbellidae 108 Subfamily Pyreninae Suter, 1919 ............................................................................ 108

Astyris gausapata (Gould, 1850) ......................................................... 109

n Astyris permodesta (Dall, 1890) .......................................................... 110

Amphissa reticulata Dall, 1916 ............................................................ 111

Amphissa bicolor Dall, 1892 ................................................................ 112

I Amphissa undata (Carpenter, 1864) ..................................................... 112 Family Cystiscidae Stimpson, 1865 ............................................................................... 113

Plesiocystiscus myrmecoon (Dall, 1919) ............................................. 113

I Superfamily Cancellarioidea ................................................................................................ 115 Family Cancellariidae Forbes & Hanley, 1851 .............................................................. 115

Key to species of Cancellariidae ....................................................................... 115

I Subfamily Cancelladinae Cancellaria cooperii Forbes &Gabb, Hanley, 18651851 ........................................................................................................... 115 116

Cancellaria crawfordiana Dall, 1891 .................................................. 116

I Subfamily Admetinae Admete californica Troschel, (Dall, 1866 ..................................................................... 1908) ........................................................... 117 118

Admete gracilior (Carpenter, 1869) ..................................................... 118

I Superfamily Key Conoidea to species .......................................................................................................... of Turridae and Conidae ........................................................... 1191

Family Turridae Swainson, 1840 ................................................................................... 122

I Subfamily Cochlespi AntipIanesxinaecatalinae Powell,(Raymond, 1942 ................................................................. 1904) ............................................... 1223

Antiplanes thalea (Dall, 1902) ............................................................. 124

m Antiplanes Carinomrrisbriseis fortis Dall, Bartsch, 19191944 ...............................................................

......................................................... 125 126

Rhodopetoma diauIax (Dall, 1908) ...................................................... 126

i Rhodopetoma renaudi (Arnold, 1903) ................................................. 128
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Subfamily Cmssispirinae Morrison, 1966 ............................................................... 128 n

Pseudotaranis strongi (Arnold, 1903) ................................................. 129

Pseudotaranis hyperia (Dall, 1919) ..................................................... 129 •

Family Conidae Fleming, 1822 ...................................................................................... 130 m

Subfamily Clathurellinae H. & A. Adams, 1858 ..................................................... 130

Borsonella bartschi (Arnold, 1903) ..................................................... 131 •

Borsonella coronadoi (Dall, 1908) ...................................................... 132 |

Borsonella omphale Dall, 1919 ........................................................... 133

Borsonella merriami (Arnold, 1903) ................................................... 134 !

Borsonella hooveri (Arnold, 1903) ...................................................... 134 •

Borsonella pinosensis Bartsch, 1944 ................................................... 135

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Ophiodermella inermis (Reeve, 1843) ................................................. 136 1

OphiodermelIa cancellata (Carpenter, 1864) ...................................... 137 1

OphiodermelIa fancherae (Dall, 1903) ................................................ 138

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Subfamily Mangetiinae Fischer, 1884 ..................................................................... 138

Kurtzina beta (Dall, 1919) ................................................................... 139 i

Kurtzia arteaga (Dalt & Bartsch, 1910) .............................................. 139

Crockerella lowei (Dall, 1903) ............................................................ 141

Crockerella crystallina (Gabb, 1865) .................................................. 142

Crockerella eriphyle (D_l, 1919) ........................................................ 143 1

Crockerella tridesmia (Bevy, 1941) .................................................... 143

Crockerella evadne (Dall, 1919) .......................................................... 144 1

Crockerella conradiana (Gabb, 1869) ................................................. 144 R

Crockerella seotti McLean, new species ............................................ 145

Crockerella philodoce (Dall, 1919) ..................................................... 146 I

Crockerella cymodoce (Dall, 1919) ..................................................... 147 II

CrockereIla castianira (Dall, 1919) ..................................................... 147

Subfamily Daphnellinae Deshayes, 1863 ................................................................ 148 !

Daphnella clathrata Gabb, 1865 ......................................................... 148 I

Acknowledgments ....................................................................................................................... 149

Literature Cited ........................................................................................................................... 149 i

2. THE OPIS'rnOBRANCnIA(by Terrence M. Gosliner)

Introduction ................................................................................................................................. 161 i

Morphology ................................................................................................................................. 162

Collection and Fixation ............................................................................................................... 163 n

Laboratory Methods [] .................................................................................................................... 163

Glossary ....................................................................................................................................... 164

List of Opisthobranch Species ..................................................................................................... 165 m

Key to Opisthobranch Species .................................................................................................... 166

Descriptions of Species ............................................................................................................... 168

Family Acteonidae ......................................................................................................... 168 •

Rictaxis punctocaeIatus (Carpenter, 1864) .......................................... 168 II

Acteon sp.............................................................................................. 171

Family Hydatinidae ........................................................................................................ 173 •

Parvamplustrum sp.............................................................................. 173

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i Family Cylichnidae ........................................................................................................ 173

Cylichna diegensis (Dall, 1919) ........................................................... 174

I Acteocina Acteocina harpa eximia (Dall, (Baird,1871) 1863)...............................................................

........................................................... 177 177

Family Philinidae ........................................................................................................... 179

Philine Philine polystrigma (Dall, 1908), new combination ............................ 180 sp. 1 .......................................................................................... 183

Philine sp. 2 .......................................................................................... 183

i Family Aglajidae ............................................................................................................ 186 Aglaja ocelligera (Bergh, 1893) .......................................................... 186

Aglaja sp............................................................................................... 187

Family Gastropteridae .................................................................................................... 187

I Gastropteron pacificum Bergh, 1893 ................................................... 187

Family Retusidae ............................................................................................................ 189

Retusaxystrum Dall, 1919 ................................................................... 189

I Volvulella Dall, 1919 191 californica

VolvuleUa panamica Dall, 1919 ........................................................... 191

Family Diaphanidae ....................................................................................................... 192

I Diaphana californica Dall, 1919 192 Family Pleurobranchidae ............................................................................................... 193

Berthella californica (Dall, 1900) ........................................................ 193

U Pleurobranchaea californica MacFarland, 1966 196

Family Aldisidae ............................................................................................................ 196

Aldisa sanguinea (Cooper, 1863) ......................................................... 196

n Family Chromodorididae 199 Cadlinaflavomaculata MacFarland, 1905 ........................................... 199

Family Potyceratidae ...................................................................................................... 201

n Holoplocamus sp.................................................................................. 201 Family Goniodorididae .................................................................................................. 201

Okenia sp. ............................................................................................. 202

I Family Dendronotidae Dendronotusfrondosus .................................................................................................... 202 (Ascanius, 1774) ............................................ 202

Family Tritoniidae .......................................................................................................... 204

I Tri Tritonia toniafestiva diomedea(Steams, Bergh, 181894 73) ............................................................

........................................................... 220504

Family Arminidae .......................................................................................................... 205

n Family Tergipedidae Armina...................................................................................................... californica (Cooper, 1863) ...................................................... 205 207

Cuthona rolleri Behrens and Gosliner, 1988 ....................................... 207

I Family AeolidiidAeolidiella ae ......................................................................................................... chromosoma (Cockerell and Eliot, 1905) .......................... 209

CerberiIla mosslandica McDonald and Nybakken, 1975 .................... 210

I Literature Cited ........................................................................................................................... 210

APPENDIX................................................................................................................................................... 214

!

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n
I

List of Figures I

Figure 1.1. Lepetidae, Lottiidae. A. lothia lindbergi McLean, I985:183 m, off Point Pifios, Monterey

County, California (LACM 2063, holotype); length 6.2 mm, 3 views. B. Niveotectura •

funiculta (Carpenter, 1864): 29 m, off Point Fermin, Los Angeles County, California (LACM

65-2.14); length 19.3 mm, 2 views .................................................................................. 14

Figure 1.2. Scissurellidae. A. Anatoma crispata (Fleming, 1832): 511-530 m, NW of Tanner Bank, i

California (LACM 77-250.23); height 4.0 mm................................................................ 17 m

Figure 1.3. Fissurellidae. A. Scelidotoma bella (Gabb, 1865): 30 m, Monterey Bay, California (USNM •

162062, holotype ofSubemarginula yatesii); length 51 mm, 3 views. B. Cranopsis cucullata I

(Gould, 1846): 12-38 m, Carmel, Monterey County, California (LACM 60-24.71); length

31.8 mm, 3 views. C. Puncturella cooperiCarpenter, 1864:86-117 m, Santa Catalina Island, •

California (LACM 40-155.17); length 7.2 mm, 3 views. D. Puncturellapunctocostata Berry, II

1947:39 m, Carmel, Monterey County, California (LACM 60-24.72); length 3.1 mm, 3

views ................................................................................................................................. 20 •

Figure 1.4. Turbinidae. A. Macrarenefarallonensis (A. G. Smith, 1952): 64-66 m, Santa Rosa Island,

California (LACM 41-197.2); height 8.6 mm, 2 views. B. Homalopoma draperi McLean,

1984:20-30 m, Isthmus Cove, Santa Catalina Island, California (LACM 2001, holotype); I!

height 4 mm, 2 views. C. Homalopoma paucicostatum (Dall, 1871): 9-15 m, N side Santa !

Cruz Island, California (LACM 63-5.30); height 4.4 ram, 2 views. D. Homalopoma berryi

McLean, 1964:21 m, N side San Nicolas Island, California (LACM 72-101.33); height 3.7 I_

mm, 2 views. E. Homalopoma cordellensis McLean, new species: 37-55 m, off Point Sur,

Monterey County, California (LACM 2779, holotype); height 7.2 mm, 2 views ........... 25

Figure 1.5. Trochidae. A. Bathybembix bairdii (Dall, 1889): 757 m, off San Clemente Island, California l"

(USNM 94865, lectotype); height 50 mm, 2 views. B. Cidarina cidaris (Carpenter, 1864):

265-274 m, SE of Santa Catalina Island, California (LACM 40-146.1); height 23.3 ram, 2 1

views. C. Calliostoma turbinum Dall, 1896:183 m, off Point Conception, Santa Barbara !

County, California (USNM 122578, holotype); height 12 mm, 2 views. D. CaIliostoma

variegatumCarpenter, 1864:73 m, off Long Point, Santa Catalina Island, California (LACM !1

53-69.1); height 22.9 mm, 2 views .................................................................................. 31 1

Figure 1.6. Trochidae.A. Calliostomaplatinum Dall, 1890:752 m, off Santa Barbara Island, California

(USNM 96558, holotype); height 32 ram, 2 views. B. Solariella peramabilis Carpenter, I[

1864:88-93 m, Cortes Bank, California (LACM 41-112.2); height 16.3 mm, 2 views. C.

Solariella nuda Dall, 1896:421-695 m, E of Long Point, Santa Catalina Island, California

(LACM 40-130.5); height 11.3 mm, 2 views. D. Halistylus pupoideus (Carpenter, 1864): •

22 m, off Point Vicente, Los Angeles County, California (LACM 19731); height 5.7 mm. II

...................... ..,.. .... . ........................ ......., ..... ...o. ................................................................ 35

Figure 1.7. Cerithiidae, Turritellidae. A. Lirobittium rugatum (Carpenter, 1864): 95-101 m, off East I

Point, Santa Rosa Island, California (LACM 41-168.8); height 10.9 mm. B. Lirobittium m

lomaense (Bartsch, 1911): 165-185 m, off Long Point, Santa Catalina Island, California

(LACM 39-104.13); height 10.3 mm. C. Lirobittiumfetellum (-Bartsch, 1911): 64 m, off II

Santa Barbara Point, Santa Barbara County, California (LACM 5-20.8); height 8.9 mm. D. el

Lirobittium Iarum (Bartsch, 1911): 59 m, off Whites Point, Los Angeles County, California

(LACM 48-14.2); height 10.5 ram. E. Lirobittium paganicum (Dall, 1919): 410 m, off !

Purisima Point, Santa Barbara County, California (SBMNH 142800, Santa Maria Basin, II

sta. R-6); height 7.8 mm. F. Turritella cooperi Carpenter, 1864:46-51 m, S of Fraser Point,

Santa Cruz Island, California (LACM 41067.3); height 35.8 mm. ................................. 41 I

|
!

i Figure 1.8. Rissoidae, Vitrinellidae, Caecidae. A. Alvania rosana Bartsch, 1911: 64-183 m, off Newport

Beach, Orange County, California (LACM 65-63.4); height 2.1 mm. B. Alvania tumida

I Carpenter, 1.2 mm. C. 1857:27-34 Vitrinella eschnaurae m, NE endBartsch, Isla Cedros, 1907:BSan aja Pedro, CaliforLos nia (LACM Angeles 71-156.21); County, California height

i (USNM 127559, holotype); diameter 2.3 mm, 3 views. D. Caecum crebricinctum Carpenter,

! • 1864: San Diego, California (USNM 14930, lectotype); length 6.2 mm........................ 47

Figure 1.9. Vanikoridae, Calyptraeidae. A. Megalomphalus califomicus (Dall, 1903): 8-21 m, Isla

Asuncion, Baja California Sur (LACM 67-66.49); height 4.6 mm. B. Megalomphalus

I schmiederi McLean, new species: 37-55 m, off Point Sur, Monterey County, California (LACM 2777, holotype); height 2.3 ram. C. Crepidula glottidiarum Dall, 1905: San Pedro,

California (USNM 183455, syntype); length 20.8 mm, 2 views. D. Crepipatella dorsata

I (B142801, mperip,Santa 1834): Maria109Bm, asin, off Poi sta.n6); t Bulength chon, S14.9 an Lumm, is Obispo 2 views.CoE.unty, Crepipatella California orbiculata (SBMNH

(Dall, 1919): 591 m, off Point Buchon (SBNHM 142802, Santa Maria Basin, sta. 16);

I length 13.2 mm, 2 views.................................................................................................. 53

Figure 1.10. Ovulidae, Triviidae. A. Neosimnia loebeckiana (Weinkauff, 1881): 41 m, off Huntington

Beach, Orange County, California (LACM 49-135.7); height 14.4 mm. B. Neosimnia

i (Dall, 1892): m, Avalon, Island, (SBMNH

barbarensis 91 off Santa Catalina California

34497, holotype of Neosimnia catalinensis); height 23 mm. C. Trivia ritteri Raymond, 1903:

74 m, off W end, Anacapa Island, California (LACM 50-17.4); height 11.2 mm................ 57

I Figure 1.11. Naticidae. A. Euspira pallida (Broderip and Sowerby, 1829): 256 m, SE of Santa Catalina

Island, California (LACM 39-131.6); height 10.8 ram. B. Calinaticina oldroydi (Dall, 1897):

I Off affinisSanta(Gmelin, Catalina1791): Island,256California m, SE of(CSanta AS 064315 Catalinaholotype); Island, height California 35 mm. (LACMC. Cryptonan 39-131.5); 'ca

height 9.5 mm ................................................................................................................... 60

i Figure 1.12. A. (Dall, 1907): Off Santa Catalina Island, California Epitoniidae. Epitonium berryi (USNM

109502, holotype of Epitonium sawinae var. catalinense); height 13.5 mm. B. Epitonium

hindsii (Carpenter, 1856): San Pedro, Los Angeles County, California (USNM 46222; height

I 18.2 ram. C. Epitonium indianorum (Carpenter, 1864): 12-38 m, Carmel, Monterey County, California (LACM 60-24.73); height 37.8 ram. D. Epitonium sawinae (Dall, 1903): Off

Newport, Orange County, California (USNM 110638, holotype of Epitonium

I acrostephanus); height 20 mm. E. Epitonium lowei (Dall, 1906): 210 m, off E1 Capitan, Santa Barbara County, California (LACM 118106); height 30.5 ram. F. Nodiscala spongiosa

(Carpenter, 1864): 46-49 m, E of Santa Barbara Island, California (LACM 39-79.9); height

H 9.8mm .............................................................................................................................. 65

i

i

Figure 1.13. Eulimidae. A. EuIima raymondi Rivers, 1904:46-53 m, off Abalone Point, Orange County,

i Californi 82-91 m,a (LACM 40-104.53); height 12.2 ram. B. Pseudosabinella bakeri (Bartsch, 1907): Isthmus Cove, Santa Catalina Island, California (LACM 70-15.7); height 5.6

ram. C. Balcis micans (Carpenter, 1864): 66-86 N, Off Platt Point, Santa Cruz Island,

i California (LACM 41-83.22): height 12.8 mm. D. Balcis oldroydae (Bartsch, 1917): on sea cucumbers, off San Pedro, Los Angeles County, California (LACM 151631): height 8.6

mm. E. Polygireulima rutila (Carpenter, 1864): 131 m, off Santa Catalina Island, California

i (LACM 54-76.9); height 7.0 mm..................................................................................... 71

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Figure 1.14. Eulimidae. A. Vitreolina columbiana (Bartsch, 1917): 27 m, off San Pedro, Los Angeles i

County, California (LACM 65-35.22); height 6.8 ram. B. Vitreolina macra: (Bartsch, 1917):

82 m, off Ship Rock, Santa Catalina Island, California (LACM 65-6.22); height 4.9 mm. C. •

Vitreolina yod (Carpenter, 1857): 366-375 m, off SE side San Nicolas Island, California |

(LACM 77-132.8); height 3.2 mm. D. Haliella abyssicola Bartsch, 1917:392 m, off Point

Sal, Santa Barbara County, California (SBNHM 142803, Santa Mafia Basin, sta. 40); height •

10.2 mm ............................................................................................................................ 75 i

Figure 1.15. Muricidae. A. Ocinebrina lurida (Middendorff, 1848): 12-38 m, Carmel, Monterey County,

California (LACM 60-24.66); height 14.1 mm. B. Ocinebrina barbarensis (Gabb, 1865): i

86 m, S of Long Point, Santa Catalina Island, California (LACM 41-179.8); height 20.1

ram. C. Austrotrophon catalinensis (Oldroyd, 1927): 46 m, off San Pedro, Los Angeles

County, California (CAS 063306, holotype); height 100 mm......................................... 81 i

i

Figure 1.16. Muricidae. A. Boreotrophon triangulatus (Carpenter, 1864): 55 m, off White's Landing,

Santa Catalina Island, California (LACM 1028, holotype of Trophon albospinosus), height !

21 mm. B. Boreotrophon multicostatus (Eschscholtz, 1829): 29 m, off Santa Catalina Island, i

California (USNM 109165, holotype of Boreotrophon peregrinus); height 23 ram. C.

I

Boreotrophon bentleyi Dall, 1908:37 m, off San Diego Harbor, California (USNM 110648, i

holotype); height 19.5 mm. D. Boreotrophon avalonensis Dall, 1902:146 m, off Avalon, i

Santa Catalina Island, California (USNM 109109); height 16.5 mm. E. Boreotrophon

avalonensis Dall, 1902:40 m, Cortez Bank, California (CAS 102992); height 19 mm. F. i

Boreotrophon eucymatu$ Dall, 1902:227 m, off San Diego, California (USNM 109089, i

holotype); height 21 mm [not 27 mm] ............................................................................. 84

Figure 1.17. Muricidae. A. Boreotrophon apolyonis (Dall, 1919): 395-620 m, Santa Barbara Islands, i

California (USNM 209303, lectotype); height 21 mm. B. Boreotrophon apolyonis (Dall,

1919): 390 m, Tanner Bank, California (LACM 76- 313.1); height 14.0 mm. C. Boreotrophon

raymondi (Moody, 1916): 390 m, Tanner Bank, California (LACM 76-313.3); height 14.9 •

rnm. D. Boreotrophon raymondi (Moody, 1916): Off China Harbor, Santa Cruz Island, I!

California, 234 m (Thelma Crow collection); height 18.8 mm. E. Boreotrophon keepi (Strong

and Hertlein, 1937): 55-91 m, off W end San Nicolas Island, California (CAS 065961); i

height 26.9 mm................................................................................................................. 88 Ii

Figure 1.18. A. Boretrophon kabati McLean, new species: 439-488 m, off Long Point, Santa Catalina

Island, California (LACM 2782, holotype); height 17.0 mm. B. Boreotrophon tolomius B

(Dall, 1919): off San Miguel Island, California (USNM 222441, holotype); height 19 mm.

i

C. Boreotrophon hazardi McLean, new species: 219 m, off Isla Vista, Santa Barbara County,

California (LACM 2783, holotype); height 20.0 mm. D. Boreotrophon pedroanus (Arnold, i

1903): 270 m, off Gaviota, Santa Barbara County, California (LACM 65582); height 14.5 II

mm. E. Boreotrophon stuarti (E. A. Smith, 1880): 112 m, off Santa Barbara, California

(USNM 122582, holotype ofBoreotrophon smithi); height 41 mm................................ 90 E

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!

I Figure 1.19. Muricidae. A. Scabrotrophon maltzani (Kobelt and Kiister, 1878): 265-274 m, SE of Santa

Catalina Island, California (LACM 40-146.6); height 21.5 mm. B. Scabrotrophon cerritensis

I (Arnold, mm C. 1903): 270 m, off San Clemente Island, California (LACM 150961); height 21.5 Scabrotrophon lasius (Dall, 1919): 366-375 m, off SE end of San Nicolas Island,

California (LACM 77-132.4); height 22.9 mm. D. Scabrotrophon grovesi McLean, new

i species: 79-140 m, off NE side of Santa Catalina Island, California (CAS 074962); height 28.1 mm. E. Scabrotrophon cIarki McLean, new species: 213 m, off Monterey, Monterey

County, California (LACM 2774, holotype); height 43.4 mm. F. Ocenotrophonpainei (Datl,

i 1903): 91 m, off Avalon, Santa Catalina Island, California (USNM 109306, holotype); height 14.2 mm................................................................................................................. 95

Figure 1.20. Turbinellidae, Buccinidae. A. Exilioidea rectirostris (Carpenter, 1864): 426 m, Santa Barbara

I Channel, 91 m, off California San Diego,(USNM California213359); height 33.7 mm. B. Exilioidea kelseyi (Dall, 1908): (USNM 110631, holotype); height 33.5 ram. C. Neptunea

amumta (Dall, 1890): 757 m, off Santa Barbara Islands, California (USNM 96556, holotype);

I height (LACM72150502); ram. D. Neptunea height 91.2tabulata mm .................................................................................(Baird, 1863): 137 m, off Anacapa Island, California 100

Figure 1.21. Nassariidae. A. Nassarius perpinguis (Hinds, 1844): San Pedro, Los Angeles County,

I (CAS 064364, syntype var.bifasciatus); height mm.

California of 21.2 B. Nassarius rhinetes

Berry, 1953:68 m, Monterey Bay, California (USNM 212133); height 33 mm. C. Nassarius

insculptus (Carpenter, 1864): 174-247 m, off Point Loma, San Diego County, California

I (USNM var. eupleura, holotype); height mm. (Trask,

209046, 15 D. Fusinus barbarensis

1855): 91 m, S of Santa Catalina Island, California (SBMNH 118915); height 98 mm.....

............. .............................................................................................. ............................. 105

I Figure 1.22. Columbellidae. A. Astyris gausapata (Gould, 1850): 82-91 m, off Long Point, Santa Catalina

Island, California (LACM 41 -185.4); height 10.5 mm. B. Astyris permodesta (Dall, 1890):

I Amphissa 505 m, off reticulata Santa Barbara Dall,Islands, 1916:113-335 Californiam, Od offSNMPoint96526, Loma, lect Sanotype); Diego height County,12.5California mm. C.

(USNM 210004, lectotype); height 16 mm. D. Amphissa bicolor Dall, 1892:563 m, off

I Amphissa Point Sur, undata Monterey County, California (USNM 106877, syntype); height 13.8 mm. E. (Carpenter, 1864): 68-73 m, S side Santa Cruz Island, California (LACM

40-165.31); height 12.0 mm ........................................................................................... 110

I Figure 1.23. MargineUidae, Cancellariidae. A. Cystiscus myrmecoon (Dall, 1919): 591 m, off Point Buchon, San Luis Obispo County, California (SBNHM 142804, Santa Maria Basin, sta.

16); height 3.9 ram. B. Cancellaria cooperi Gabb, 1865:124 m, off Santa Barbara, California

I (Beach, USNM Los 106860); A height 56 mm. C. Cancellaria crawfordiana Dall, 1891: Off Redondo ngeles County, California (LACM 72505); height 49.8 mm. D. Admete

californica (Dall, 1908): 1103 m, off San Diego, California (USNM 110626, lectotype);

I height County,15.8California mm. E.Admete gracilior(Carpenter, 1869): 69 m, off Point Fermin, LOsAngeles (LACM 57-90.1); height 11.4 mm .................................................. 114

Figure 1.24. TCalifornia urridae. A. Antiplanes catalinae (Raymond, 1904): 229 m, off Santa Catalina Island, (USNM 192087, lectotype); height 25.1 mm. B. Antiplanes thalea (Dall, 1902):

461 m, off San Luis Obispo, California (USNM 122568, holotype); height 40 mm. C.

Antiplanes thalea (Dall, 1902): 97 m, off Santa Rosa Island, California (USNM 109198,

I holotype ofA. santarosana); height 36 mm. D. Antiplanes briseis Dall, 1919:150-161 m,

Avalon Bay, Santa Catalina Island, California (LACM 40-123.7); height 25.0 ram. E.

Carinoturrisfortis Bartsch, 1944:227-229 m, N of Anacapa Island, California (LACM 4lI 52.3); height mm............................................................................. .......................

14.3 123

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!

Figure 1.25. Turridae. A. Rhodopetoma diaulax (Dall, 1908): 62 m, off Islas Los Coronodos, Baja I

California (holotype, USNM 110609); height 19.0 mm. B. Rhodopetoma diaulax (Dall,

1908): 150 m, off Santa Rosa Island, California (holotype of B. rhodope, USNM 212361); m

height 19 mm. C. Rhodopetoma renaudi (Arnold, 1903): 159-174 m, SE of Santa Catalina l

Island, California (LACM 41-209.12); height 12.3 mm. D. Pseudotaranis strongi (Arnold,

1903): 511-530 m, Tanner Bank, California (LACM 77-250.22); height 13.1 ram. E. •

Pseudotaranis hyperia (Dall, 1919): 177-190 m, off San Nicolas Island, California (LACM B

40-99.6); height 15.0 mm ............................................................................................... 127

Figure 1.26. Conidae. A. BorsoneIla bartschi (Arnold, 1903): 214-234 m, SE of Santa Catalina Island, i

California (LACM 40-125.11); height 21.0 mm. B. Borsonella coronadoi (Dall, 1908):

229-240 m, Cortes Bank, California (LACM 41-114.3); height 22.0 mm. C. BorsoneUa

omphale Dall, 1919:274-311 m, off Pyramid Cove, San Clemente Island, California (LACM i

39-126.3); height 19.8 mm. D. Borsonella merriami (Arnold, 1903): 104 m, S of Santa []

Rosa Island, California (LACM 41-172.12); height 18.1 mm. E. Borsonella hooved (Arnold,

1903): 511-530 m, NW of Tanner Bank, California (LACM 77-250.24); height 21.1 mm. F. II

BorsoneUa pinosensis Bartsch, 1944:119-165 m, off Emerald Bay, Santa Catalina Island,

California (LACM 39-5.7); height 24.8 mm.................................................................. 132

1.27. Conidae. A. OphiodermeUa inermis (Reeve, 1843): 18 m, off San Diego, California (USNM i

i

Figure

I

110644, holotype of halcyonis); height 23 mm. B. Ophiodermella cancellata (Carpenter,

1864): 198-199 m, off Long Point, Santa Catalina Island, California (LACM 41-164.21); i

height 8.5 mm. C. Ophiodermellafancherae (Dall, 1903): 86-95 m, NW of Anacapa Island, []

California (LACM 41-47.17); height 10.0 mm. D. Kurtzina beta (Dall, 1919): 102 m, off

i

Point Afio Nuevo, San Mateo County, California (USNM 206554, holotype); height 5.0

mm. E. Kurtzia arteaga (Dall and Bartsch, 1910): 22-27 m, off Point Loma, San Diego •

County, California (USNM 150993, holotype of var. ropen); height 6.5 mm .............. 136 aiD

Figure 1.28. Conidae. A. Crockerella lowei (Dall, 1903): 95 m, S of East Point, Santa Rosa Island, •

California CLACM 41-326.5); height 9.9 ram. B. CrockereUa lowei (Dall, 1903): 96 m, off n

San Miguel Island, California (LACM 75-412.2); height 9.2 ram. C. Crockerella crystallina

(Gabb, 1865): 101 m, E of Santa Rosa Island, California (I.,ACM 41-327.6); height 8.6 ram. []

D. CrockereUa eriphyle (Dall, 1919): 247-256 m, SW of Balboa, Orange County, California l

(LACM 41-6.3); height 11.8 mm. E. Crockerella tridesmia (Berry, 1941): 196 m, E of

Long Point, Santa Catalina Island, California (LACM 41-146.6); height 10.2 ram. F. i

CrockereUa evadne (Dall, 1919): 91 m, offRedondo Beach, Los Angeles County, California il

(LACM 17351); height 9.0 mm..................................................................................... 141

Figure 1.29. Conidae. A. CrockereUa conradiana (Gabb, 1869): 55 rn, off Santa Catalina Island, California i

(LACM 151660); height 10.4 mm. B. Crockerella scotti McLean, new species: 100 m,

Santa Barbara Channel, California (LACM 2776, holotype); height 11.5 mrn. C. Crockerella

philodoce (Dall, 1919): 439-488 m, off Long Point, Santa Catalina Island, California (LACM II

41-182.3); height 9.5 mm. D. Crockerella cymodoce (Dall, 1919): 59 m, off Goleta, Santa

Barbara County, California (LACM 75-463.2); height 12.6 mm. E. Crockerella castianira

(DalI, 1919): 104-113 m, SE of Bennett Point, San Miguel Island, California (LACM 41- _l

176.7); height 11.6 mm. F. Daphnella clathrata Gabb, 1865:219 m, off Isla Vista, Santa

Barbara County, California (LACM 74-70.1); height 19.9 mm .................................... 145

!

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xviii i
!

Figure 2.1. Living animals A. Rictaxis punctocaelatus (Carpenter, 1864). B. Aglaja ocelligera (Bergh,

1893). C. Gastropteron pacificum Bergh, 1893. D. Berthella californica (Dall, 1900). E.

I Pleurobranchaea californica MacFarland, 1966. F. Diaphana californica Dall, 1919...... 171

Figure 2.2. A. Rictaxis punctocaelatus (Carpenter, 1864), juvenile shell, × 40. B. Acteon sp., shell, ×

30. C. Parvamplustrum sp., preserved animal, × 80 ...................................................... 172

I Figure 2.3. A. RictaxispunctocaeIams (Carpenter, 1864), radular teeth, x 1000. B., C. Acteon sp, B.

jaw rodlets, x 3000. C. radular teeth, x 4000. D. Parvamplustrum sp., radular teeth, x 2000.

i ........................................................................................................................................ 174 Figure 2.4. A. Cylichna diegensis (Dall, 1919), shell, x 20. B. Acteocina eximia (Baird, 1863), shell, x

10. C., D. Acteocina harpa (Dall, 1871), C. shell, x 40. D. Gizzard plates of holotype, x 25.

I ........................................................................................................................................ 177

Figure 2.5. Cylichnadiegensis (Dall, 1919).A. Jaw rodlets, x 3000.B. Lateral teeth, x 1000. C. Rachidian

I teeth, x 2000. D. Gizzard plate, x 150........................................................................... 178

i I Figure 2.6. Acteocina harpa (Dall, 1871). A. Rachidian teeth, x 4000. B. Lateral teeth, x 3000. C., D.

Acteocina eximia (Baird, 1863). C. Lateral teeth, x 1000. D. Gizzard plates, x 20...... 180

I Figure 2.7. A-C. PhiIinepolystrigma (Dall, 1908). A. Preserved animal, x 25.B. Central nervous system,

x 40. C. Penis, x 40. D-F. Philine sp. 1, D. Preserved animal, x 15. E. Shell, x 20. F. Penis,

i 40 .................................................................................................................................... 183 Figure 2.8. A., B. Philinepolystrigma. A. Lateral teeth, x 700. B. Gizzard plate, x 50. C., D. Philine sp.

1, C. Lateral radular teeth, x 1200. D. Gizzard plate, x 70 ............................................ 184

I Figure 2.9. A. Philine sp. 1,shell sculpture, x 300. B. Philine sp. 2, radular teeth, x 700. C. Gastropteron

pacificum Bergh, 1893, radular teeth, x 300.................................................................. 186

I Figure 2.10. A. Philine sp. 2, preserved animal, x 80. B. Aglaja sp., preserved animal, x 40. C.-E. Aglaja ocelligera (Bergh, 1893). C. Preserved animal, x 5. D. Shell, x 10. E. Penis, × 10...... 187

i Figure 2.11. A, B. Gastropteron pacificum Bergh, 1893. A. Shell, after MacFarland, 1966. B. Penis, x

25. C. Diaphana caIifornica shell, x 10. D. Retusa xystrum Dall, 1919, shell, x 50. E.

Volvuella califomica DaU, 1919, shell, x 25. F. VolvueIla panamica Dall, 1919, shell, x 30.

............ .................. ...................... ............ .................................................... ..... ............... 190

I Figure 2.12. A.,B.RetusaxystrumDall, 1919. A. Shellsculpture, x5OO. B. Gizzardplates, x4OO. C.,D.

Diaphana californica Dall, 1919. C. Rachidian teeth, x 1500. D. Lateral teeth, x 1500 ....

I ........................................................................................................................................ 192

Figure 2.13. A, B. BertheUa califomica (Dall, 1900). A. Shell sculpture, x 300. B. Jaw rodlets, x 1500.

C. Pleurobranchaea californica MacFarland, 1966, jaw rodlets, x 1200. D. Berthella

I californica (Dall, 1900), teeth, x californica

inner radular 200. E. Pleurobranchaea

MacFarland, 1966, radular teeth, x 500......................................................................... 196

I Figure 2.14. A. preserved Berthellaspecimen. californicaC. (Ventral Dall, 1900 view), shell, of preserved x 30. B.,specimen, C. Holoplocax 40. musD.spOkenia . B. dorsal sp., view ventralof

view of preserved specimen, x 65. E. Cerberilla mosslandica McDonald and Nybakken,

I 1975, living animal. F. Tritonia diomedea Bergh, 1894, living animal ........................ 197

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Figure 2.15. Living animals. A. Aldisa sanguinea (Cooper, 1863), B. Cadlinaflavomaculata MacFarland, I

1905. C. Tritoniafestiva (Stearns, 1873). D. Dendronotusfrondosus (Ascanius, 1774) ....

........................................................................................................................................ 199 I Figure 2.16. A, B. Aldisa sanguinea (Cooper, 1863), radular teeth, x 500. C., D. Okenia sp. C. Inner

lateral teeth, x 1500. D. Outer lateral teeth, x 3000 ....................................................... 200

Figure 2.17. A.-C. CadIinaflavomaculata MacFarland, 1905. A. Jaw rodlets, x 5000. B. Cenlxal portion I

of radula, x 1000. C. Outer lateral teeth, x 1000. D. Holoplocamus sp., radular teeth, x

400 .................................................................................................................................. 202 I Figure 2.18. Radular teeth. A. Dendronotusfrondosus (Ascanius, 1774), x 700. B. Cuthona rolleri

Behrens and Gosliner, 1988, x 1000. C. Tritoniafestiva (Stearns, 1873), x 1000.D. Tritonia

diomedea Bergh, 1894, x 700 ........................................................................................ 205 I

Figure 2.19. Living animals. A. Armina californica (Cooper, 1863). B. Cuthona rolleri Behrens and

Gosliner, 1988. C. Aeolidiella chromosoma (Cockerell and Eliot, 1905) ..................... 208 I Figure 2.20. A., B. Armina californica (Cooper, 1863). A. Jaw rodtets, x 1000. B. Central radular teeth,

x 700. C. Aeolidiella chromosoma (Cockerell and Eliot, 1905), radular teeth, x 500. D.

Cerberilla mosslandica McDonald and Nybakken, 1975, radular teeth, x 700 ............ 210 I

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I Acknowledgments

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

Interior, Minerals Management Service, Washington, D.C., under Contract No. 14-35-0001-30484. Ms.

I Made volume.Murphy of the Santa Barbara Museum of Natural History provided production assistance for this

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i List of Acronyms

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,

I transect A/B.

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

I transect C/D.

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

! California, USA.

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

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

I MMS United States Minerals Management Service.

NHML The Natural History Museum, London, United Kingdom.

I SCAMIT Southem 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.

UCMP Univesity of California Museum of Paleontology, Berkeley, California, USA.

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I

| 1. THE PROSOBRANCHIA

! u,

James H. McLean _

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i Introduction

The Subclass Prosobranchia, also known as Streptoneura, is used here in its traditional sense, although

I it is now understood to be a paraphyletic group, rather than one that is monophyletic and includes all of its descendants. As used here it includes the recently recognized suborder Patellogastropoda and the recently

recognized suborder Vetigastropoda (Patellogastropoda and Vetigastropoda are part of the old suborder

I Archaeogastropoda), and the suborders Mesogastropoda and Neogastropoda (which together are also known as Caenogastropoda). These are the gastropods in which the effect of torsion is retained in the adult members---

the mantle cavity retains its anterior position. The gill is a typical etenidium, for the most part bipectinate in

I Pate that lare logastropoda treated here, andhave Vetigastropo external dshells. a, and monopectinate in Caenogastropoda. Most members, including all

Some families that have previously been treated as prosobranchs, including the families Pyramidellidae,

I Architectonicidae, and Rissoellidae, are now considered primitive members of the Heterobranchia along with the more derived groups Opisthobranchia and Pulmonata; the latter two groups are united as Euthyneura (see

Gosliner, Chapter 2).

I Gastropod classification is currently a state likely to some

in of flux and is remain unsettled for time.

A traditional classification was outl/ned by Vaught(1989), but changes basedon cladistic analysis of anatomical

charactershave been proposed by recent authors, including Haszprunar (1988, 1993) and Ponder and Waren

I 1 Further modifications A detailed treatment of phylogeny and classification is omitted ( 988). are anticipated.

here because this work is intended primarily as an identification manual.

I Prosobranch gastropods represent the largest group of marine mollusks; in numbers of species they surpass those of all other groups combined. Species can usually be identified on shell characters alone. The

morphology of the radula differs among species in some families, providing additional characters for species

l differentiation; in other families it may be useful only at the family level. i

i Laboratory Methods

Identification of shelled gastropods is possible without using specialized techniques for relaxation or

I preservati and shouldon,bealthusedoughforthe purposes methodsother described than identification by Gosliner inof Cspecies. hapter 2Small-shelled of this volumespecimens are equallycanapplic be fixed able

intact in 75% ethyl alcohol or 10% buffered formalin for 24 hours, followed by washing and transfer to 75%

I ethyl order alcoh enableol.theThefixative entire btoodies penetrate of larger through specime thnearly s canncoil otsbeoffithexdshell. unessShells the shell are best is crapreserved cked in ainvice, a dryin

state; even those kept in buffered alcohol will suffer corrosion after a number of years. Specimens previously

I fixed decomposition. in formalinTheorradulalcoho(and l cantobesome washed extent, andother dried;organs) the initial can still fxabetionstudied allowats the a later tissdate ues byto dry rehydrating without

the specimen, followed by preparation using conventional methods.

I 1 Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA 90007

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Scope I

I

The objective of this work is to provide an illustrated identification manual for the prosobranch •

gastropods occurring offshore at depths of 50 to 500 m in the vicinity of the Santa Mafia Basin and the U

western Santa Barbara Channel. For the most part, it also applies to those of the southern California Bight,

the entire area from Point Conception to the Mexican border. Itis intended to provide coverage for species not i

included in my earlier field guide Marine Shells of Southern California (McLean, 1969; revised, 1978), in U

which I treated the species likely to be found in the intertidal zone and in the rocky sublittoral zone accessible

by diving. A total of 126 species are treated herein, including 7 new species. Major revisions of the genera

Lirobi_'um, Boreotrophon, Scabrotrophon (new genus), Borsonella, OphiodermeIla and Crockerella are

accomplished here.

Although my original intention had been to treat the heterobranch Pyramidellidae, which are well !

represented in the offshore habitats of southern California, they are not treated here because I have not as yet

U

studied them to the same extent as the groups that are treated in this work.

There is little material in museum collections and few published records for the offshore gastropod I

fauna from the Santa Maria Basin and the entire area north of Point Conception. The collection produced by

the Santa Maria Basin project thus fills a major gap. However, the voucher collection does not include many m

species of moderate to even large size that are expected to occur there, considering thattheir known distributions •

include the Monterey Bay area and the southern California Bight. I have therefore included species not in the

voucher collection and have attempted to treat most of the offshore species not covered in Marine Shells of

Southern California that are well represented in collections and known to occur between Monterey Bay and

southern California. In order to increase the general utility of this work, there are some included species that

u

are yet known only from the southern California Bight. Such species can be expected to occur north of Point m

Conception because the offshore fauna is less subject to the thermal barriers that may delimit the northern and

southern faunas atPoint Conception. However, budget limitations have made it necessary to omit some of the

m

offshore species known only from the southern California Bight. Other species omitted are those of the lower

continental slope below 500 m and the few broad ranging, true abyssal species, which have been poorly

sampled off southern California.

This work coincides with work of mine on the entire prosobranch and shell-bearing heterobranch In

gastropod fauna ranging from Alaska to the outer coast of Baja California. Detailed accounts for approximately I

1,100 species are in preparation, both for an annotated checklist and for an illustrated manual to follow. The

checklist, which is nearing completion, will provide a current summary of my understanding of the systematic m

arrangement and synonymy of all names for shelled, benthic gastropods that have been proposed for the n

faunal provinces between Alaska and Baja California, giving geographic distributions and depth ranges for

each species. Additional genera necessary for the species included here and in the larger context have been

published (McLean, 1995), although one further new genus is introduced herein. Taxonomic innovations I_

made here have also been done in the larger context of consideration of the entire fauna. Species that had to be

omitted from the present work will therefore soon be fully treated, both in the checklist and the illustrated

manual. I

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I Conventions

I Superfamilies

Traditional superfamilies are given in centered heading but are not diagnosed.

I Families and Subfamilies

Families and subfamilies are diagnosed and notes on biology and general remarks with references to

I major revisions are included. These treatments are not exhaustive and not be consistent from one family

may

to the next. The objective here, however, is to lead the reader to the most recent accounts of the taxonomy and

phylogeny of the group.

I Genera

An understanding of generic concepts depends upon a comparison to the type species, which is a

I matter of importance to taxonomists. Type species and the manner of their designation are therefore cited

here. The type species is the one species in the genus that is representative of the genus. If the type species was

explicitly indicated in the original description the designation is called original designation (OD). If only one

I species was original account no statement to designate a type species,

included in the and there is it becomes

the type species by monotypy (M). If, however, more than one species was included and the author of the

genus did not select a type species, then the first subsequent author who does so is cited as designator by

I subsequent designation (SD). Type species are by original combination, genus species

cited their the and in

which they were first proposed, although this may have little beating on the genus to which they are now

assigned. In addition, the faunal affinity of the type species is indicated.

I Diagnosis. Brief description of the genus, intended to be applicable to all members.

Biology. Information about feeding and reproduction. If this section is omitted, see the heading under

I the family. Remarks. Notes that apply to all species of the genus, including references to papers that are restricted

in scope to the genus.

I Species

Synonymies. All synonymies include the citation of the original description for the nominate species

I and its synonyms. Additional references in the synonymies are not complete but are intended to give major illustrated works, including the first illustration for a species originally described without figures.

References to the three volume work of Oldroyd (1927) are given because she provided a useful

I reprint of the original description and copies of original figures. Note that pagination for Oldroyd is that from the bottom of the page, continuous for the three volumes. References to Abbott's American Seashells (1974)

serve to indicate some recent name changes.

I To make it easier to follow, each in is treated in its various before

permutations name asynonymy

going on to the next synonym.

i Material Examined. Gives station numbers and number of specimens for examined lots in the voucher collections for the Santa Maria Basin project. For maps showing positions of the stations see the appendix at

the end of this volume. According to the contract for the preparation of this series, the Phase I and Phase II

i voucher specimens are being divided between the collections of the Santa Barbara Museum of Natural History

and the United States National Museum of Natural History.

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The number of examined lots in the LACM collection are also given to provide a quantitative measure •

of abundance for each species. Lots are the curated units in reference collections of mollusks; each lot can

contain one or more specimens from the same collecting station. •

Description. Brief descriptions are based on examination of specimens and study of previous literature. II

Size is indicated in mm for specimens considered to be mature, with a lower and upper limit.

Type Locality and Type Specimens. The type locality is the locality cited in the original publication, 1

in some cases slightly abbreviated here for consistency. For some species, in which only museum catalog U

numbers for type material was originally included, I give further information from the original label.

mill

Type specimens cited here are of three kinds. Holotype: The single specimen used in the original !

description. Syntypes: The original specimens used in an original description--indicating that a single specimen II

was not designated asthe holotype by the original author. Lectotype: A single specimen from a syntype lot that

has been designated by a later author to be the primary specimen of comparable standing to a holotype. 1

References to lectotypes require citation of the author and date of the publication in which the designation was

It

made. A number of lectotype designations for species and synonyms described by Dall are made herein.

Paratypes and paralectotypes are not cited in the present work, except for the new species.

u

Depth records for the primary types are given in the original form, usually in fathoms. If this information

is missing, none was provided in the original description. l

Institutional abbreviations are found on after the List of Figures in the preface (page xx). I

Catalog numbers for types at the CAS have been updated to the new 6-digit numbering system.

Distribution. Distributions are given north to south. States are cited, but not countries (e.g., Canada 1

and Mexico). To assist in locating the end points, the latitude is given to the nearest half degree (e.g., 33°N,

35.5°N).

Northern and southern records are based on specimens in the LACM collection, unless the record is mR

m

held in another museum, in which case the museum and catalog number is cited. For some species, the

distributional record may be based on a published record, which is then cited by author, date, and page. In

some cases the LACM catalog number of a distributional record is given in brackets.

Many of the end points represent new distributional records, but no effort is made to distinguish new

limits because the baseline records in the literature for eastern Pacific gastropods are so out of date that there •

is no recognized source of information. |

Habitat. Species are noted as being characteristic of either hard or soft bottoms and an estimation of

the upper and lower limits of bathymetric occurrence is indicated in meters. These estimates are obtained from 1

museum labels and take into account the original depths for type material. Extreme records from collections 1

are ignored, as the objective is to increase the predictive value of the information.

Remarks. Comparisons to related species and notes relevant to the taxonomic history are given in 1

this section. Changes to previously understood synonymies and classification are mentioned here.

If a species is often assigned to an established subgenus, this is mentioned here. Subgenera are usually

omitted in the preparation of faunal lists and they are therefore not used in the formal headings in this work. 1

g

Illustrations. Locality information in the captions is abbreviated and does not include geographic

coordinates from the LACM labels. The size for each figure was calculated from a plot of shell height against

figure height, in order to show larger shells at a larger size than those of small-shelled species. B

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I Literature Cited

I The Literature Cited is intended to be complete for all references to publications in the text and the

entries in the synonymies for which the pagination is given. It does not include references to the authors of

i famil purposes ies, ofgenera comparison; and the type designations. It does not include references to species cited in the text for in such cases, the date is part of the name of the species, rather than a referral to the

Literature Cited.

I by subscripts Multiple (e.g.,public a,b).aIttioisnas simple by th matter sam auth to determine or in a single the year corect in the reference Literature by checking Cited arethenotpagination distinguishedand

figures for synonymy entries to see if they coincide with the inclusive pagination for each reference.

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Glossary

! Anal notch. Notch on outer lip near suture, Body whorl. The largest and terminal spiral whorl

corresponding to position of anus. of the shell.

I Anal sinus. Same as anal notch. Caleffieafion. Formation of shell from calcium

carbonate.

Angulation. Carina at midwhorl position.

I Anterior notch. Notch at base of shell for the Callus. Thick deposit of shell material.

siphon of the living animal. Canal. Extensions of the aperture; anterior or

I Aperture. Opening of the last whorl, through scanal iphonmay al canal be called protectsthetheanalsipho notchn;. posterior

which the head and foot of the living animal

emerge. Canedlate sculpture. Formed by axial ribs and

I Aphallie. Lacking a penis or copulatory organ.

spiral cords intersecting at tight angles.

Carina. Prominent spiral keel or ridge.

Apex. Early whorls or tip of the spire; this is the

I end. Chitin. Uncalcified protein forming periostracum posterior

Apical. Pertaining to the apex. and operculum.

I Clathrate. Same as cancellate.

Aspinose. Lacking spines.

Columella. Shell pillar surrounding the axis of

Axial lamellae. Axial ribs of thin, bladelike form. coiling.

I Axial ribs. Sculpture parallel to the axis of coiling. Concentric. Sculpture of a limpet that corresponds

Axis of coiling. Imaginary line aroundwhich the to growth tings.

I shell coils. Cord. Ridgelike spiral sculpture.

Base. Anterior end of the shell, the lower partto Costae. Thin, strongly projecting axial ribs.

the left of the aperture if adextral shell is

I held Crenulate, Having aregularly notched edge,

apex up.

usually from terminations of spiral sculpture.

Beading. Sculpture produced where axial ribs are

I crossed by spiral cords. Deck. calyptraeids Shelf of sheU. material in aperture of

Biearinate. Having two carina per whorl.

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Denticle. Tooth shaped projections on inner and/or Notch. Indented margin of shell. •

outer lip. Odontophore. Cartilage associated with radula.

Dextral. Shell coiling in which the aperture is on 1

the right when the shell is viewed with apex Operculum. A horny, sometimes calcified, N

at the top. structure attached to the foot that may seal

the aperture of the shell when the animal

Direct development. Larval stage passed within retracts. U

egg capsule. Outer lip. Outermost margin of the aperture, the

Dorsum. Back side of gastropod shell, opposite growing edge of the shell. I

aperture. Parietal wall. Broad inner lip area of the body

1

Fold. A ridge or plication on the columella, whorl. 11

II Foramen. Hole in shell of Fissurellidae marking Paucispiral operculurn. Operculum having few

position of anal siphon, whorls.

m

Fossa. Broad trenchlike depression. Paucispiral protoconeh. Protoconch having only a I

few whorls or turns.

Fusifom. Elongate shell profile with spire and

canal of similar length. Penultimate whorl. The next to last whorl, m

Growth lines. Simple axial sculpture resulting Periostraeum. Horny or fibrous outermost layer of

from successive growth increments on the the shell, composed of conchiolin. •

outer lip. Peristome. Outline of the aperture.

II

Iudsed sculpture. Sharply cut grooves.

Periphery. Outermost part of the whorl that m

Inner lip. Inner margin of the aperture, projects farthest from the axis. I

Intercalary cords. Cords of lesser strength Peritreme. Combined growing edge of aperture,

between primary cords, including inner and outer lips. I

Involute. Shell growth in which the length of the Planktotrophie. Larval stage that is freeaperture increases, concealing the spire swimming and feeding in the plankton, n

Lameliae. Sculpture of thin scales. Pustule. Small unit of knobby sculpture.

Lateral teeth. Teeth on either side of the rachidian Plait. A fold or plication on the columella. •

tooth. Plicae. Folds. Columellar plicae are folds on the II

Lip. The growing edge of the shell, columella.

mm

Lip lirae. Spiral ridges within aperture at outer lip. Plicate: Folded or twisted. I

Marginal teeth. The outermost radular teeth in the Ptenoglossate. Type of radula with similar teeth in

row. the row. I

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Multispiral operculum. Operculum having many Protandric hermaphrodite. Sex change from male

whorls or turns, to female. •

Multispiral protoeonch. Protoconch having many Protoeonch. The larval shell. U

whorls or turns. Protractive. Axial sculpture in which the fibs

Nacre. Pearly interior layer of the shell (if present), project forward. U

Nodose. Beating tubercles or knobs. Raehidian tooth. The central radular tooth in the

row. !

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I Radial. Sculpture of a limpet that radiates from the Spiral cords. Raised ridges that encircle the shell

apex. at right angle to coiling axis.

I Radula. Ribbon bearing teeth used for rasping Spire. Whorls at apical end of the gastropod, food. posterior to the aperture.

i Retraetive. Axial sculpture in which the ribs (and Striae. Sculpture of minute grooves or ridges. growing edge) curve backward. Subsutural. Referring to area just below suture

Selenizone. Lamellar deposition marking previous when apex is held up.

Suture. Line marking the junction of the whorls.

positions of the anal notch.

Semilunate. Half moon outline of aperture Tabulate. With shelflike area below suture.

i pro outerduced lip. by straight columeUa androunded Teleoeoneh. The shell except for the protoconch.

Sinistral. Shell coiling in which the aperture is on Umbilicus. Open cavity atthe base of the shell,

I the left when the shell is viewed with apex at within the axis of coiling.

the top. Umbilical ehiul_ Narrow depression, rather than

Siphon. Anterior extension of mantle used for open cavity.

I water intake. Variees. Pleural of varix.

Siphonal canal. Anterior extension of shell

I surrounding the siphon. Varix.repArlesenting arger thantheordoutinar er ytipaxial duringrib,a resting

Siphonal faseiole. Lamellar deposition marking growth stage.

I previous positions of the siphonal canal. Veliger. Larval stage of gastropods with velar Siphonal notch. Indented margin of shell for lobes.

protrusion of siphon. Volution. A whorl of a shell or turn in an i

II

I Shoulder. Uppermost projecting area on the whorl, operculum.

Slit. Indented margin of shell for water intake in Whorl. A turn or coil in a spirally coiled shell. The

I Scissurellidae. body whorl is the last and most inflated; the penultimate is the next to the last whorl.

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List of Prosobranch Species I_

(new species in bold)

Family Lepetidae Family Caecidae I

Iothia lindbergi McLean, 1985 Caecum crebricinctum Carpenter, 1864

Family Lottfidae Family Vanikofidae •

Niveotectura funiculata (Carpenter, Megalomphalus californicus (Dall, []

1864) 1903)

Megalomphalus schmiederi McLean,

Family Scissurellidae new species

Anatoma crispata (Fleming, 1832) !

Family Calyptraeidae

Family Fissurellidae Crepidula glottidiarum DalI, 1905 •

Scelidotoma bella (Gabb, 1865) Crepipatella dorsata 03roperip, 1834)

Cranopsis cucullata (Gould, 1846) CrepipateIla orbiculata (DaI1, 1919)

Puncturella cooperi Carpenter, 1864 ..

Puncturella punctocostata Berry, 1947 Family Ovulidae

Neosimnia loebeckiana (Weinkauff, 1881) I

Family Turbinidae Neosimnia barbarensis (Dall, 1892)

Macrarenefarallonensis (A. G. Smith, []

1952) Family Triviidae II

Homalopoma draperi McLean, 1984 Trivia ritteri Raymond, 1903

Homalopoma paucicostatum (Dall, ..

1871) Family Naticidae l

Homalopoma berryi McLean, 1964 Euspira pallida 03roderip and Sowerby,

Homalopoma cordellensis McLean, 1829)

new species Calinaticina oldroydi (Dall, 1897) •

Family Trochidae Cryptonatica affin is (Gmelin, 1791) !

Bathybembix bairdii (Dall, 1889)

Cidarina cidaris (Carpenter, 1864) Family Epitoniidae

Calliostoma turbinum Dull, 1896 Epitonium berryi (Dall, 1907) 1

Calliostoma varie gatum Carpenter, Epitonium hindsii (Carpenter, 1856)

1864 Epitonium indianorum (Carpenter, 1864)

Calliostoma platinum Dall, 1890 Epitonium sawinae (Dall, 1903) []

Solariella peramabilis Carpenter, 1864 Epitonium lowei (Dall, 1906) 1

SolarieIla nuda Dall, 1896 Nodiscata spongiosa (Carpenter, 1864)

Halistylus pupoideus (Carpenter, 1864)

Family Eulimidae I

Family Cerithiidae Eulima raymondi Rivers, 1904

Lirobittium rugatum (Carpenter, 1864) Pseudosabinella bakeri (Bartsch, 1907)

I_,irobittium lomaense (Battsch, 1911) Balcis micans (Carpenter, 1864) []

Lirobittium fetellum (Bartsch, 1911) Balcis oldroydae 03artseh, 1917) 1

Lirobittium larum 03artsch, 1911) Polygireulima rutila (Carpenter, 1864)

Lirobittium paganicum (DaJl, 1919) Vitreolina columbiana 03artsch, 1917)

Vitreolina macra (Bartsch, 1917) 1

Family Turritellidae Vitreolina yod (Carpenter, 1857)

Turritella cooperi Carpenter, 1864 Haliella abyssicola Bartsch, 1917

i

Family Rissoidae Family Muricidae I

Alvania rosana Bartsch, 1911 Ocinebrina lurida (Middendorff, 1848)

Alvania tumida Carpenter, 1857 Ocinebrina barbarensis (Gabb, 1865)

Austrotrophon catalinensis (Oldroyd, 1

Family Vitrinellidae 1927)

_trinella eschnaurae Bartsch, 1907 Boreotrophon triangulatus (Carpenter,

1864) I
!

I Boreotrophon multicostatus (Eschscholtz, Family Cystiscidae

1829) Plesiocystiscus myrmecoon (Dall, 1919)

Boreotrophon bentleyi Dall, 1908

I Boreotrophon avalonensis Dall, 1902 Family Cancellariidae Boreotrophon eucymatus DaU, 1902 Cancellaria cooperii Gabb, 1865

Boreotrophon apolyonis (Dail, 1902) CancelIaria crawfordiana Dail, 1891

I BoBoreotrophon reotrophon keepi raymondi (Moody, 1916) Admete califomica (Dall, 1908) (Strong and Hertlein, Admete gracilior (Carpenter, 1869)

1937)

Boreotrophon kabati McLean, Family Tumdae

I new species Antiplanes catalinae (Raymond, 1904) Boreotrophon tolomius (Dall, 1919) Antiplanes thalea (Dall, 1902)

Boreotrophon hazardi McLean, Antiplanes briseis Dall, 1919

I Bornew eotropho specniespedroanus (Arnold, 1903) RhCaodo rinopetoma turrisfortdiaulax is Bartsc (Dalh,

l, 191908 44 )

Boreotrophon stuarti (E. A. SJrfith, Rhodopetoma renaudi (Arnold, 1903)

1880) Pseudotaranis strongi (Arnold, 1903)

I Scabrotrophon maltzani (Kobelt and Pseudotaranis hyperia (Dall, 1919) Kiister, 1878)

Scabrotrophon cerritensis (Arnold,1903) Family Conidae

I ScScabrotrophon abrotrophon lasi grovesi us (DMcL all, ea191n,9) BoBorsrsooneIla nella bacoronadoi rtschi (A(Dall rnold,, 11903) 908)

new species Borsonella omphale Dall, 1919

Scabrotrophon clarki McLean, Borsonella merriami (Arnold, 1903)

I Borsonella hooveri (Arnold, 1903)

new species

Ocenotrophonpainei (Dall, 1903) BorsonelIa pinosensis Bartsch, 1944

OphiodermeIla inermis (Reeve, 1843)

I Family Turbinellidae Exilioidea rectirostris (Carpenter, 1864) Ophiodermella 1864) cancellata (Carpenter,

Exilioidea kelseyi (Dall, 1908) Ophiodermellafancherae (Dall, 1903)

', Kurtzina beta (Dall, 1919)

[ 1 Family Buccinidae Kurtzia arteaga (Dall and Bartsch, 1910)

Neptunea amianta (Dall, 1890) Crockerella lowei (Dall, 1903)

Neptunea tabulata (Baird, 1863) Crockerella crystallina (Gabb, 1865)

I Crockerella eriphyle (Dall, 1919)

Family Nassariidae Crockerella tridesmia (Berry, 1941)

Nassarius perpin guis (Hinds, 1844) Crockerella evadne (Dall, 1919)

Nassarius rhinetes Berry, 1953 Crockerella conradiana (Gabb, 1869)

I Nassarius insculptus (Carpenter, 1864) Crockere Crock Ua scotti McLean, new species erella philodoce (Dall, 1919)

Family Fasciolariidae Crockerella cymodoce (Dall, 1919)

I Fusinus barbarensis (Trask, 1855) Daphnella Crockerellaclathrata castianiraGabb, (Dall,1865 1919)

Family Columbellidae

Astyris gausapata (Gould, 1850)

I Astyris Amphissapermod reticulata esta (DDall, all, 11916 890)

' Amphissa bicolor DaU, 1892

I Amphissa undata (Carpenter, 1864)

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Key to Families I

(based on shell characters of species treated herein)

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Because shell characters are often convergent among prosobranch families, few can be defined on

shell characters alone. The key that follows is not based on relationships and applies only to the species I

treated herein; other species in some families may not conform to the distinctions used here. A key based on 1

phylogenetic distinctions would also be based on characters of radula and anatomy, but would be more

difficult to use for identification purposes. 1

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1A. Shell of limpet form ......................................................................................................................... 2

lB. Shell spirally coiled or tubular ......................................................................................................... 5 I

2A. Shell simple cap-shaped ................................................................................................................... 3 1

2B. Shell with foremen, notch at margin, or interior deck ...................................................................... 4 |

Apex one-forth shell length from anterior margin ............................................................... Lepetidae I

3A.

3B. Apex nearly central .............................................................................................................. Lottiidae

m

4A. Shell with notch or hole ................................................................................................. Fissurellidae 1

4B. Shell with interior deck ................................................................................................ Calyptraeidae 1

5A. Shell elongate, tubular ......................................................................................................... Caecidae

5B. Shell spirally coiled .......................................................................................................................... 6 I

6A. Shell with deep slit at periphery ................................................................................... Scissurellidae •

6B. Shell not with deep slit at periphery ................................................................................................. 7 l

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7A. Aperture circular or nearly so, rounded anteriorly, lacking siphonal canal or notch ....................... 8 1

7B. Aperture elongate, with siphonal canal or notch ............................................................................ 16

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8A. Shell interior nacreous ..................................................................................................................... 9 I

8B. Shell interior not nacreous ............................................................................................................. 10

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9A. Operculum calcified on outer surface ............................................................................... Turbinidae

9B. Operculum not calcified on outer surface .......................................................................... Trochidae I

10A. High-spired, shell height more than 3 times aperture length .......................................................... 11 I

10B. Low-spired, shell height less than 3 times aperture length ............................................................ 13 I

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I 11A. Axial ribs present ............................................................................................................. Epitoniidae

1lB. Axial ribs lacking ........................................................................................................................... 12

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12A. Spiral sculpture present ................................................................................................. Turritellidae

i 12B. Spiral sculpture absent ...................................................................................................... Eulimidae

13A. Shell medium to large, over 8 mm diameter ....................................................................... Naticidae

I 13B. Shell minute, under 5 mm diameter ............................................................................................... 14

I 14A. 14B. Shell Shell umbilicate not umbilicate .............................................................................................................................. ............................................................................................................ Rissoidae15

I 15A. Axial sculpture present ................................................................................................... Vanikoridae

15B. Axial sculpture lacking ................................................................................................... Vitrinellidae

I 16A. Aperture as long as shell ................................................................................................................ 17

i 16B. Aperture less than length of shell ................................................................................................... 18

17A. Spiral sculpture lacking ....................................................................................................... Ovulidae

I 17B. Spiral sculpture present ....................................................................................................... Triviidae

I 1818BA.. CCol olumumell ellaa smoot with plic h .......................................................................................................................... ations .............................................................................................................. 2190

I 19A. Sculpture lacking ............................................................................................................... Cysticidae

19B. Axial and spiral sculpture present ................................................................................ Cancellariidae

I 20A. Canal short or lacking .................................................................................................................... 21

i 20B. Canal extending past length of aperture ......................................................................................... 23

21A. Aperture not deeply notched ............................................................................................. Cerithiidae

2lB. Aperture with deep siphonal notch ................................................................................................. 22

I 22A. 22B. Fasciole Fasciole wlacking ith deepdeepfosfossa sa ..................................................................................................

......................................................................................... Columbellidae Nassariidae

I 23A. Outer lip with deep anal sinus ................................................................................ Turridae, Conidae

23B. Outer lip not with deep anal sinus .................................................................................................. 24

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24A. Sculpture spinous or lamellar ............................................................................................ Muricidae

24B. Sculpture not spinous or lamellar .................................................................................................. 25 I

25A. 25B. Scul Sculpture pture spiral axial and onlyspiral ......................................................................................................... .............................................................................................................. Buccinidae26 I 

26A. Periostracum smooth, glossy ......................................................................................... Turbinellidae I

26B. Periostracum rough, fibrous ........................................................................................ Fasciolariidae

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I Descriptions of Species

I Superfamily Acmaeoidea

I Lepetidae, According Acmaeidtoae,theandrecent Lottiidae. classifiThe cationdistinction of Lindbergbetween (1988:55), the Lottiidae there are and threethefamilie Acmaeidae s in the swas uperfamil detailedy:

by Lindberg (1986:142), along with other genetic changes among the patellogastropod limpets of the northeastern

I Pacific.

Family Lepetidae Gray, 1850

I Diagnosis. Shell of limpet form, with horseshoe-shaped muscle scar opening anteriorly; apex anterior,

anterior slope concave, posterior slope convex, color usually white. Intermediate shell layers foliated and

concentric crossed lamellar. Eyes and nuchal cavity gill lacking. Radula with 2 or 3 pairs of lateral teeth, first

I pair medially fused, forming a tooth; marginal pairs, developed.

central teeth 2 well

Biology. Lepetid species live in cold water faunal provinces, some occurring in shallow water at high

latitudes, others are deeply submergent at low latitudes. The only paper on the biology of a lepetid species is

I that of Yonge (1960).

Remarks. The Lepetidae are commonly known as the blind limpets, as the eyes of other

I patellogastropods are lacking. Genera and species of Lepetidae are few. The family was reviewed worldwide by Moskalev (1977).

Because differences in shell sculpture and external anatomy provide few genetic characters, the classification

i is primarily based on radular characters. There are two main expressions of the fused inner lateraltooth: one having a long andtaperedpoint, and one having cutting edge extending straight across. There is also abasic

difference in marginal teeth, one kind hasthe cusp of the marginal teeth smooth andthe other has a comblike

I fringe. Combinations of these two traits enables the definition of four genera on the basis of these characters.

Genus lothia Gray, 1850

I lothia Gray, 1850. Type species (M): PateIlafulva MiiUer, 1776. Northeastern Atlantic.

i Diagnosis. Shell small, white, red-orange, or white with gray markings; sculpture of broadly spaced, imbricate radial ribs. Apex inclined anteriorly; protoconch erect, rounded, subspiral; apex of adult shell worn

away. Interior termination of muscle scar posterior to apex. Animal with paired labial appendages. Fused

i inner lateral tooth of radula elongate and sharp pointed, outer pair of laterals greatly reduced and closely adjacent to fused inner laterals; marginal teeth narrow, inner surface with comblike fringe.

Remarks. lothia differs from Lepeta Gray, 1847, in having an erect, non-spiral protoconch, the

I termina fringed tirather on ofthan the msmooth uscle. scar behind the apex rather than in front of it, the marginal teeth narrower and

The massive, sharp-pointed central tooth (a fusion of the inner lateral teeth) may function to pierce

I the tests of encrusting bryozoa, which often occurs on rocks bearing lothia.

lothia species occur offshore, usually at depths greater than those accessible to divers.

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lothia lindbergi McLean, 1985 I1

Figure 1.1A

Iothia lindbergi McLean, 1985:336, fig. 1. 1

Material Examined. California, Santa Mafia Basin, Phase I: sta. 2, 200 m (4); sta. 6, 109 m (7). .i

Other material: 22 lots in the LACM collection. I

Description. Shell thin, translucent. Anterior slope concave to straight, posterior slope convex. Ill

Sculpture of concentric growth lines and widely spaced, irregular, imbricate ribs. Exterior and interior of shell 1

white. Radula as for genus Length 5-10 mm.

1

Type Locality and Type Specimens. Off Point Pifios, Monterey County, California, 180 m. Holotype:

LACM 2063. 1

Distribution. Queen Charlotte Islands, British Columbia (54°N), to Islas San Benitos, Baja California

(28°N)" Habitat. Rocky bottoms, 40-900 m. I

Remarks. This species differs fromlothiafulva (Miiller, 1776) in the northeast Atlantic primarily in

lacking the reddish orange coloration of that species. No members of the genus are known to occur in the 1

Arctic Ocean.

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Figure 1.1. Lepetidae,Lottiidae.A. Iothia lindbergi McLean,1985:183 m, off PointPifios,MontereyCounty, I

California (LACM2063, holotype); length 6.2 ram, 3 views. B. Niveotecturafuniculta (Carpenter, 1

1864): 29 m, off Point Fermin, Los Angeles County, California (LACM 65-2.14);length 19.3ram,

2 views.

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I Family Lottiidae Gray, 1840

I shell varioDiaguslnoy scisolored. . ShellShell of limpecomprised t form, with of prismati horseshoe-shaped c outer layers muscand le scar crososed-lamellar pening anterioinner rly; apex layers. anterior, Eyes

present, nuchal cavity gill usually present on left side. Radula with 3 pairs of mineralized lateral teeth; marginal

t teeth either 2 pairs, 1 pair, or lacking. Biology. Lottiid limpets graze on surficial algae and diatoms. They are broadcast spawners.

Remarks. Lottiid limpets axe abundant in numbers of species and individuals in the intertidal zone of

I the eastern Pacific.

Generic characters include details of shell structure and the number of marginal teeth and their

i morphology.

Genus Niveotectura Habe, 1944

_1 Niveotectura Habe, 1944. (OD): Patella Gould, 1859. Northwestern Pacific. aypespoeie_ pallida

Diagnosis. Shell medium to large, white; sculpture of strong radial ribs; lateral teeth 3 pairs; marginal

I teeth lacking.

Biology. The species of Niveotectura have the lateral teeth with blunt tips suited for grazing on

coralline algae.

I Remarks. There are two living species, the type species, which is a relatively large species common

in shallow water of the northwest Pacific, and the smallerNiveotecturafuniculata, occurring offshore in the

northeastern Pacific. The genus was treated in detail by Lindberg and Marincovich (1988), who described a

i fossil species from Alaska.

1 Niveotecturafuniculata (Carpenter, 1864) B

Figure lAB

I Scurria mitra var.funiculata Carpenter, 1864:650.--Palmer, 1958:123, pl. 17, figs. 24, 25 [holotype:].

l

Acmaea mitra vat.funiculata: Oldroyd, 1927:747.

I Acmaeafuniculata: Hanna and Smith, 1931:21, pi. 2, figs. 1,3,4.--Lindberg, 1981:96, fig. 96A.

Niveotecturafuniculata: Lindberg, 1988:143.

i Scurria Scurria mmititrara var. var.tenuiscul tenuisculpta pta "Carpe Carpenter, nter," 18Dall, 66:346 1871:242.--Palmer, [nomen nudum]. 1958:124, pl. 18,figs. 11-13 [syntype].

Ill Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

l examined: 47 lots in the LACM collection.

Description. Shell of medium to high profile, apex subcentral. All slopes straight. Sculpture of

I broad, equal inflatsizeradial andrishape; bs. Extecriusps or ablunt; nd interiobasal r coplates lor white, simple. sometimes Lengthtinted 15-25withmm.pink. Radular teeth approximately

Type Locality and Type Specimens. Monterey, California. Holotype: USNM 14799.Scurrla mitra

i var. tenuisculpta: Neah Bay, Clallam County, Washington; syntypes: USNM 15490.

[] Distribution, Shumagin Islands, Alaska (55°N, 160°W), to Bahia Magdalena, Baja California Sur,

Mexico (25 °N).

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Habitat. 20-70 m, rock bottoms with coralline algae, below the thermocline in California. I

Remarks. This species was first assigned to Niveotectura by Lindberg (1986:143). Superficially it

resembles Acmaea mitra Rathke, 1833, in the family Acmaeidae, the shell of which differs in structure of its I

shell layers and in lacking the radial sculpture. Although the distribution of Acmaea mitra extends into the l

sublittoral, Niveotectura funiculata replaces it at greater depths, where it is the only species of Lottiidae

known to occur. There is variation in shell height and strength of the radial ribbing. A detailed account of this []

species was first given by Hanna and Smith (1931). Palmer (1958:124) discussed the nude name Scurria []

mitra var. tenuisculpta, the validation of which she credited to Dall (1871).

I

Superfamily Scissurelloidea I

Family Scissurellidae Gray, 1847

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Diagnosis. Shell small, turbinate, umbilicate; lip with either slit or foramen; operculum multispiral. I

Shell structure of lamellar aragonite, interior lacking nacre. Ctenidia paired, the left larger than the fight,

ctenidial filaments finger-shaped. Left kidney a papillary sac; columellar muscles paired. Cephalic lappets []

and neck lobes lacking, cephalic tentacles setose, epipodial tentacles setose (except first 2 pairs); mantle 1

margin with small papillae and setose pallial tentacles projecting through slit. Rachidian tooth of radula broad

at base, tip narrow, with finely denticulate, overhanging cutting edge; first 4 lateral teeth cusped like rachidian

and with broad basal areas overlaping adjacent laterals; fLfthlateral larger and having long, finely denticulate I

tip; marginal teeth numerous, with long, serrate tips.

Biology. Scissuretlids occur on soft bottoms offshore and on rock and gravel bottoms in shallow I

water. What little is known of the biology of scissureUids is based on knowledge of Anatoma crispata, as

detailed by Fretter and Graham (1962, 1976). That species feeds on detrital material; its reproductive biology

is unknown, l

The most recent account of scissurellid anatomy is that of Haszprunar (1989).

Remarks. The Scissurellidae have small white shells with an open slit or foramen, which serves as an m

excurrent opening for water currents passing through the mantle cavity. Scissurellids resemble the much l

m

larger pleurotomariid slit-shells. Most scissurellid species occur offshore, where they are often overlooked

because of their small size. lira

The Scissurellidae have long been recognized as a family. Based on anatomical differences from the l

Pleurotomariidae, the group has recently been elevated to the superfamily level (Haszprunar, 1989; McLean,

1989), an action more recently supported by studies of pleurotomariid sperm morphology by Healy and l

Harasewych (1992). |

McLean (1989) offered a subfamilial classification of Scissurellidae based on gill and protoconch

characters, recognizing four subfamilies. In addition to the typical subfamily, three were proposed as new: •

Anatominae, Temnocinclinae, and Sutilizoninae, the last two restricted to the hydrothermal-vent habitat. l

Subfamily Anatominae McLean, 1989 I

Diagnosis. Genera of Anatominae have the protoconch sculpture finely reticulate, the ctenidia •

bipectinate and the radula with an enlarged pturicuspid ( fourth lateral tooth). l

Remarks. Anatominae differ from Scissurellinae in protoconch sculpture. Protoconchs of the

scissurelline genera Scissurella d'Orbigny, 1824, and Sinezona Finlay, 1927, differ in having sculpture of !

strong axial fibs. l

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I Genus Anatoma Woodward, 1839

Anatoma Woodward, 1839. Type species (M): ScissureUa crispata Fleming, 1832. Circumboreal.

I Description. Shell turbinate, umbilicate; spire elevated. Selenizone at the periphery, slit open onefifth circumference of last whorl. Sculpture cancellate; protoconch smooth.

I Remarks. Species occur in all oceans, primarily offshore on soft bottoms. The type species has the [ largest sized shell in the family. The genus Scissurella d'Orbigny, 1824, which is unknown in the eastern

Pacific, differs in having the selenizone positioned on the shoulder, a lower spire, and a protoconch with axial

I sculpture.

Anatoma erispata (Fleming, 1832)

I Figure 1.2A

Scissurella crispata Fleming, 1832:385.--Fretter and Graham, 1976:2, figs. 1, 2.

I Scissurella (Anatoma) crispata: McLean, 1967:405, pl. 56, figs. 1-4.

Scissurella (Schizotrochus) kelseyi Dall, 1905:124.

I ScissurelIa chiricova Dall, 1919: 370.

Material Examined. California, Santa Maria Basin, Phase I: sta. 36, 492 m (1); sta. 108,492 m (2).

I Other material: 30 lots in the LACM collection.

Description. Globose, delicate, translucent white, whorls 4, edges of selenizone sharp, raised. Axial

sculpture off'me raised ridges, crossed by fine spiral ridges of equal strength. Outer lip thin, inner tip slightly

I reflected; umbilicus deep. Height 3-6 mm.

Type Locality and Type Specimens. Noss Island, Shetland Islands, Scotland; type material not

located. Scissurella kelseyi: off Point Loma, San Diego County, California, 640 fathoms; holotype: USNM

I 181820, specimen missing. Scissurella chiricova, Island, Alaska, fathoms; holotype:

SE of Chirikov 695

USNM 206509.

Distribution. Circumboreal, Arctic Ocean to Isla Cedros, Baja California (28°N); also south to the

I Mediterranean, the Azores, New England, and Japan.

Habitat. Soft bottoms from sublittoral to

I abyssal depths, becoming deeply submergent at lower latitudes. In the eastern Pacific it occurs as

shallow as 55 m off the Queen Charlotte Islands,

I BCritish Columbia, and at 400-1800 m off alifornia

Remarks. This species, described

ooriginally f additionalfromsyntheonyms NorthfroAtlant m local ic,ities has oather number than

the eastern Pacific, as indicated by McLean (1967).

I The excepsyn tionally onymlarge S. kelse specimen, yi Dall6 mmwasinbased height.onOnean

other species of the genus, Anatoma lyra (Berry,

I southern 1947), occurs California in rocky (McLean, sublittoral 1967). habitats in

Figure 1.2. Scissurellidae.A. Anatoma crispata (Fleming, 1832): 511-530 m, NW of TannerBank, California

I (LACM 77-250.23); height 4.0 ram.

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Superfamily FissureHoidea I

Family Fissurellidae Fleming, 1822 I

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Diagnosis. Shell conical, of limpet form, with anterior slit, notch, or dorsal foramen marking position

of exhalent siphon; operculum lacking in adult. Shell composed of lamellar aragonite (some with exterior ill

calcitic layer); shell interior not nacreous. Sculpture of radial ribs and concentric ribs of lesser strength; I!

muscle scar horseshoe-shaped. Heart diotocardian, ctenidia paired, of equal size, with afferent and efferent

n

membranes. Left kidney reduced, nearly vestigial. Epipodial tentacles stubby, in single row along sides of m

foot. Mantle folds capable of expansion to envelop shell, head, and foot. Radula rhipidoglossate, strongly i

asymmetric, the rachidian and 4 laterals weakly cusped, pluricuspid (outermost fifth lateral) large and strongly

cusped; marginals numerous. BII

Biology. Members of the geologically oldest genera live from shallow to moderate depths and are n

known to graze upon encrusting invertebrates, particularly sponges and compound ascidians. The youngest

genus, Fissurella Brugui_re, 1789, feeds upon intertidal algae. Fissurellids are broadcast spawners; the !

planktonic veliger stage is short (Fretter and Graham, 1976). I

Remarks. The fissurellid limpets are known as the slit limpets or the keyhole limpets, owing to the

slit or fissure in the shell that serves as the excurrent opening from the mantle cavity. This is the only limpet II

group with paired gills. n

The subfamily classification followed here is modified from that of Thiele (1929). Two subfamilies,

the Emarginulinae and the Fissurellinae, are recognized, based primarily on radular differences (McLean, I_

1984, both references). Characters of the radula and the ontogeny of the slit or foramen provide the basis for II

the principal dichotomy. In the older Emarginulinae, the radula has a rhomboidal rachidian tooth, the outer

lateral tooth is usually bicuspid, there are varying expressions of the slit or foramen, and a selenizone is ll

represented, if onty in early shell ontogeny. In the Fissurellinae, the rachidian tooth has a narrow neck and is 1

similar in shape to the adjacent laterals; the outer lateral tooth is usually quadricuspid. The excurrent opening

in the shell is always art apical foramen and there is no indication of a selenizone in early ontogeny. I

Key to Species of Fissurellidae i!

1A. Shell with anterior notch at margin ........................................................................ Scelidotoma bella

lB. Shell with interior septum; foramen on anterior slope ..................................................................... 2 !

ill

2A. Mature shell length < 4 mm ..................................................................... Puncturella punctocostata !

2B. Mature shell length > 6 mm ............................................................................................................. 3

3A. Radial ribs sllong, projecting at margin ............................................................. Cranopsis cucullata U

3B. Radial ribs low, not projecting at margin ........................................................... Puncturella cooped

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I Subfamily Emarginulinae Gray, 1834

I selenizonDiagnosis. e, at least inRaearly chidiangrowth toothstages. rhomboidal, outer lateral tooth bicuspid; slit or foramen preceded by

Remarks, All eastern Pacific fissurellids other than FissurelIa are assigned to the Emarginulinae.

I Genus Scelidotoma McLean, 1966

I SceIidotoma McLean, 1966. Type species (OD): EmarginuIa bella Gabb, 1865. Northeastern Pacific.

Diagnosis. Shell large, white, with coarse radial ribs. Apex nearly central; selenizone marked by

i inte edgerioofr channel shell. andarched exteriorly. Juvenile shells finely punctate and deeply notched, apex close to posterior

Remarks. Scelidotoma differs from other emarginuline genera in undergoing an ontogenetic change--

I tshells. he juvenile shells are deeply notched as inEmarginula, the notch becoming sinuous and marginal in mature

I had previouly thought that the genusArginula Palmer, 1937, type species Emarginula arata Conrad,

I 1833, from the Eocene of Alabama, was appropriate for the species bella, as used by Kozloff (1987:200), but no longer hold that view.

There are two species in the northwestern Pacific: SceIidotoma gigas (Von Martens, 1881), and S.

I (Yokoyama, 1922).

vadosinuata

I Scelidotoma bella (Gabb, 1865)

Figure 1.3A

I Emarginula bella Gabb, 1865:188.

Hemitoma bella: Oldroyd, 1927:840.--Abbott, 1974:21 (listed only).

I Scelidotoma beIla: McLean, 1966:1, figs 1-2. Subemarginula yatesii Dall, 1901:125.--Da11, 1902:555, pl. 38, figs. 1, 3.

I hard bottoms MatermialtheExanorth mined. and Not south.repre Other s ntedmateinrithe al: Santa 21 lotsMaria in theBas LACMin vouccollection. her collection, but occurring on

Description. Shell relatively large, yellow-white; sculpture of about 24 strong primary ribs alternating

I with 1-3 secondary ribs. Margin crenulate; interior glossy white. Mantle yellow-white, foot slightly larger than shell. Length 40-75 mrn.

Type Locality and Type Specimens. Monterey, California; "dredged." Holotype: UCMP 12552.

I Subemarginula yatesii: Monterey, holotype:

California; USNM 162062.

Distribution. Forrester Island, southeastern Alaska (55°N), to Cabo San Martin, Baja California

(30°N).

I Habitat. Uncommon throughout its range, it has been found intertidally only in Alaska; in California

it occurs on rocky bottoms below the thermocline at 20-1 t0 m.

I Remarks. The lack of an illustration and the smaller size (13.8 mm) of the original specimen of Scelidotoma bella accounted for the description of Dall's synonym Subemarginula yatesii.

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Figure 1.3. Fissurellidae. A. Scelidotoma bella (Gabb, 1865): 30 m, Monterey Bay, California (USNM 162062,

holotype of Subemarginula yatesii); length 51 ram, 3 views. B. Cranopsis cucuUata (Gould, 1846): I

12-38 m, Carmel, Monterey County, California 0_,ACM 60-24.71); length 31.8 ram, 3 views. C.

Puncturella cooped Carpenter, 1864:86-117 m, Santa Catalina Island, California (LACM 40-

155.17); length 7.2 ram, 3 views. D. Puncturella punctocostata Berry, 1947:39 m, Carmel, Monterey

County, California (LACM 60-24.72); length 3.1 mm, 3 views. I

Genus Cranopsis A. Adams, 1860 I

Cranopsis A. Adams, 1860. Type species (M): Cranopsis pelex A. Adams, 1860. Japan.

Diagnosis. Anterior slope with interior and exterior groove or seam that extends from constricted I

lower end of foramen to shell margin. Groove located exteriorly between 2 closely adjacent ribs that form

doubled, split. Septumanteriormost arched convexly, primary rib; not doubled supported ribbydeflected props. slightly toward right. Mantle skirt correspondingly I

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I Remarks. Earlier authors placed the Cranopis species in PunctureIla Lowe, 1827, but the split

mantle skirt of Cranopsis (described by Cowan and McLean, 1968) is unlike that ofPuncturella Lowe, 1827,

i in which there is neither a split mantle nor a seam on the anterior slope of the shell, a distinction sufficient for the separate generic ranking of the two taxa.

Species of Cranopsis occur in both the northern and southern hemispheres, in shallow water at high

I latitudes and at greater depths at low latitudes.

Cranopsis cucullata (Gould, 1846)

I Figure 1.3B

I Rimula cuculIata Gould, 1846: 159. Puncturella cucullata: Oldroyd, 1927:843, pl. 93, fig. 4.

Puncturella (Cranopsis) cucullata: Abbott, 1974:22, fig. 59.

I Material Examined. Santa Mafia Basin, Phase I: sta. 6, 109 m (1); sta. 20, 396 m (1). Other

material: 85 lots in LACM collection.

I Description. Shell relatively large, high conical, primary narrow, raised, usually

outline fibs 16,

anteriormost lower than others and doubled; interspaces broad, secondary fibbing faint; apex anterior to

center; foramen just anterior to apex, short, diamond-shaped; septum simple, convex. Length 30-42 mm.

I Type Locality and Type Specimens. Puget Sound, Washington. Lectotype (selected by Johnson,

1964: 63): USNM 4325.

I Distribution. Kodiak Island, Alaska (57°N), to Cabo San Quintin, Baja California (30°N). Habitat. Occurring on vertical rock surfaces where it feeds on sponges. It occurs in progressively

deeper water toward the southern part of its range: intertidal in the Gulf of Alaska, at 5 m in Southeastern

i Alaska, below 20 m in central California, and below 100 m in southern California.

Remarks. Other northeastern Pacific species of Cranopsis are C. major (Dall, 1891), C. multistriata

(Dall, 1914), and C. decorata Cowan and McLean, 1968. The latter two species are also known from deep

I water in southern California. Cranopsis cucuIlata differs from each in having fewer radial fibs.

I Genus Puncmrella Lowe, 1827 PunctureUa Lowe, 1827. Type species (monotypy): Patella noachina Linnaeus, 1771. Circumboreal.

I apex Diagnosis. Shell conical, white; young shells with fine white punctations. Foremen on anterior slope, inclined toward fight. Interior with forward-sloping septum separateing mantle cavity from visceral

region; septum often flattened, supported by props; septum continuous with depressed selenizone originating

I just anterior to apex. Remarks. Puncturella lacks the anterior seam on the shell and the split mantle roof that characterizes

Cranopsis. The shells are also somewhat smaller than those of Cranopsis. lake Cranopsis, Puncturella

species occur at high latitudes or in deeper waters atlow latitudes in both the northern and southern hemispheres.

!

!
!

Puncturella cooperi Carpenter, 1864 I

Figure 1.3C

! Puncturella cooperi Carpenter, 1864:651.--Oldroyd, 1927:842.--Palmer, 1958:120, pl. 18, figs. 16, 17.

Puncturella eyerdami Dall, 1924:133.

II

Material Examined. Not represented in the voucher collection from the Santa Mafia Basin, but

occurring to the north and south. Other material: 64 lots in LACM collection.

Description. Relatively small, high-conical; outline oval. Septum simple, extending straight across, I

lacking props. Sculpture of numerous low, unequal ribs of varying strength, interspaces usually with f'me

granular punctations. Length 6-9 mm, height 6-10 mm. II

Type Locality and Type Specimens. Off Santa Catalina Island, California, 20-40 fathoms. Holotype: II

USNM 11848. Puncturella eyerdami: Knight Island, Prince William Sound, Alaska; lectotype (here

designated): USNM 360136. •

Distribution. Kenai Peninsula, Alaska (59°N), to Punta Rompiente, Baja California (28°N). There

is also a record (LACM 69-59.2) from the Cerralvo Channel, Gulf of California.

Habitat. On rock bottoms at 20-130 m throughout its distribution, not uncommon. I

Remarks. Most specimens have the sculpture eroded. The lectotype of P. eyerdami is an uneroded

specimen that shows time punctations, which led Dall to believe that what he described as that species was

different. I

Puncturella punctocostata Berry, 1947 I

It

Figure 1.3D

Puncturella punctocostata Berry, 1947:11, pl. 1, figs. 7-9. i

Puncturella ralphi Berry, 1947:13, pl. 1, figs. 4-6.

Puncturella (Puncturella)punctocostata: McLean in Keen, 1971:313, fig. 11. llw

Material Examined. There are no specimens represented in the voucher collection from the Santa

Mafia Basin, but the species is common offshore to the north and south. Other material: 37 lots in the LACM

collection. I

Description. Small, apex relatively large, posterior to center. Septum thin, extending straight across

and lacking props. Ribs beaded and varying in number, strength, and prominence of the beading; rib interspaces I_

f'mely punctate. Outline variable, nearly circular to elongate-oval. Length 2.0-3.5 mm. I

Type Locality and Type Specimens. Middle Pleistocene, Lomita Marl, San Pedro, California.

Holotype: S. S. Berry Collection no. 11998. Puncmrella ralphi: Same locality; holotype: Berry no. 11999; 1

paratypes: SBMNH 34530. Holotypes for both taxa are missing; they have not been located in Berry material R

at SBMNH or CAS.

Distribution. Cordell Bank, off Point Reyes, Matin County, California (38°N), to Cortes Bank, 1

California (32°N) and Isla Guadalupe, Baja California (29°N). There is a single record from the Gulf of

California: 55 m at Isla Carmen (26°N) (SDNHM). II

Habitat. On rocky bottoms in 20-1 O0 m, particularly atthe offshore banks and islands where it may I

be collected by divers.

!
!

I Remarks. This species and its synonym were described originally from the Pleistocene of southern

California, but has subsequently been found living. Puncturella ralphi was based on specimens with more

I numerous ribs, but such specimens are well within the range of variation observed in Recent specimens.

Superfamily Trochoidea

i Family Turbinidae Rafmesque, 1815

Diagnosis. Shell small to large, interior usually nacreous, umbiticate or non-umbilicate; aperture

i oblique, sculpture varied. Operculum with paucispiml pattern on corneous inner surface; outer surface with thick calcareous deposit, with pattern of sculpture produced by envelopment by foot. Protoconch with pointed

tip. Ctenidium bipectinate, having free tip and either long or very short dorsal afferent membrane. Radula

i rhipidoglossate; Biology. rachid Turbinids ian tgene oothrallarger ly livethanon halaterals; rd bottoms, marginals grazingnumero on algaus.l films. They arebroadcast spawners,

with short veliger stages.

I by, the foot, Remexcept arks. Turb wheninids the animal are characterized is disturbedbyanda calcareous the foot withdrawn operculum,intowhich the isshell. secreted Hickman by, andandenMcLean veloped

(1990) divided the family into several subfamilies, including Liotiinae, Colloniinae, Turbininae, and Tricoliinae.

I The muchrhipidoglossate radula has a similar pattern throughout the family, unlike the Trochidae in which there is greater radular diversity. The relatively unspecialized mdula led Hickman and McLean (1990) to regard

the Turbinidae as more primitive than the Trochidae.

I

Key to Species of Turbinidae

!

1A. Opercutum with calcareous beads on external surface ............................... Macrarenefarallonensis

1 lB. Operculurn with fully calcified external surface .............................................................................. 2

2A. Axial sculpture present .................................................................................... Homalopo_a drape ri

I sculpture lacking ....................................................................................................................

2B. Axial 3

I 3A. 3B. Three More than spiral3cords spiral per cordswhper orl ................................................................ whorl .................................................................................................. HomaIopoma paucicostatum 4

I 4A. Four to 5 spiral cords per whorl .............................................................. Homalopoma cordellensis

4B. More than 6 spiral cords per whorl .................................................................... Homalopoma berryi

I

I

I
!

Subfamily Liotiinae H. and A. Adams, 1854 I

Diagnosis. Shell small to medium-sized, deeply umbilicate, interior nacreous. Aperture circular, i

peritreme complete, suture descending on final whorl; final lip usually thickened. Axial sculpture of sharp II

lamellar increments and more broadly spaced axial ribs, both kinds of axial sculpture especially prominent

deep within umbilicus; spiral sculpture of prominent cords. Operculum multispiral, comeous on inner surface,

outer surface with calcareous granules or beads deposited on each volution and aligned in radiating pattern;

chitinous fringe produced at outer edge of each volution. Left ctenidium lacking dorsal afferent membrane

along most of length. Cephalic lappets lacking, neck lobes simple. Rachidian tooth broad, cutting edge poorly I

developed; lateral teeth 5, with overhanging tips, bulging laterally and overlapping adjacent lateral teeth; II

marginals numerous, inner marginals like those that follow.

Remarks. The scaly sculpture, thickened final lip, and beaded operculum are characteristic features

of this subfamily. U

Genus Macrarene Hertlein and Strong, 1951 I

Macrarene Hertlein and Strong, 1951. Type species (OD): Liotia caIifornica Dall, 1908. Southern California.

Diagnosis. Relatively large for family, white, depressed turbiniform; umbilicus broad; aperture circular, i

inflated, suture descending on final whorl. Sculpture off'me axial fimbriations and coarse spiral cords; periphery

strongly keeled, made stellate by axial ribs on upper half of whorl. Final lip lacking varix; axial sculpture i

more closely spaced preceding final lip. Operculum markedly concave, each whorl with thin calcareous ridge i

and dense tufts of chitinous material projecting at margin.

Liotia. Remarks. Macrarenes are the largest liotiids. SmaU specimens have the appearance of the genus I

One other species occurs in California, the larger M. cookeana (Dall, 1918), which lives at depths

accessible to divers as far north as Anacapa Island (see McLean, 1978:23, fig. 9.8). i

Macrarenefarallonensis (A. G. Smith, 1952) II

Figure 1.4A i

Liotiafarallonensis A. G. Smith, 1952:385, pl. 20, figs. 2-4. i

g

Macrarenefarallonensis: McLean in Keen, 1971:348, fig. 137.

Arene (Macrarene) farallonensis: Abbott, 1974:55. i

II

Material Examined. Not represented in the voucher material from the Santa Mafia Basin, but occurring

to the north and south. Other material: 4 LACM lots. I

Description. Shell white, periphery stellate to spinose. Spiral sculpture of 3 major cords: peripheral I

cord, another at level of suture, and strong basal cord. Axial sculpture strongest across base. Height 9 mm,

diameter 12 mm.

Type LoealRy and Type Specimens. Southeast Farallon Island, California. Holotype: CAS 064382.

Distribution. Cordell Bank, California (38°N), to San Jaime Bank, off Cabo San Lucas, Baja California

Sur (23°N). i

Habitat. Rocky bottoms, 60-140 m, below depths accessible to divers.

i

24 i
!

!

!

!

!

!

!

!

I

!

!

!

I Figure 1.4. Turbinidae. A. Macrarenefarallonensis (A. G. Smith, 1952):64-66 m, Santa RosaIsland,California

(LACM 41-197.2); height 8.6 mm, 2 views. B. Homalopoma draperi McLean, 1984:20-30 m,

i Isthmus Cove, Santa Catalina Island, California (LACM 2001, holotype); height4 mm, 2 views. C. Homalopoma paucicostatum (Dall, 1871): 9-15 m, N side Santa Cruz Island, California (LACM

63-5.30); height 4.4 ram, 2 views. D. Homalopoma berryi McLean, 1964:21 m, N sideSan Nicolas

Island, California (LACM 72-101.33); height 3.7 ram, 2 views. E. Homalopoma cordellensis

I McL he ean,new species: 37-55 m, offPoint StarM, ontereyCounty,California (LACM2779,holotype); ight 7.2 nun, 2 views.

I species usually occur together: one developed sculpture

Remarks. Two forms of this has well axial

(as in the holotype figured by A. G. Smith) and the other has a more stellate peripheral cord and weaker axial

sculpture (figure 1.4A). Such variation would suggest the presence of two separate species if intermediate

I forms not known from most localities at which the species has been collected.

were

The species was first discovered in bird guano at Southeast Farallon Island, the original specimen

I having evidently fn-st been swallowed by a bottom-feeding fish.

i 25
!

Subfamily Colloniinae Cossmann, 1916 I

Diagnosis. Shell small to medium in size, sculpture chiefly spiral, columella usually with 1 or 2 low •

denticles, mature lip often thickened. Operculum calcareous on outer surface. il

Biology. Little is known of the biology of colloniine species. They occur on rock or gravel bottoms.

Remarks, Prior to the revision of Hickman and McLean (1990), this group was known as the I

Homalopomatinae Keen, 1960. Hickman and McLean used the older name Colloniinae Cossmann, 1916, a

,u

name that had originally been intended for non-nacreous genera. However, the presence or absence of interior

nacre is not significant in the subfamily, even at the generic level. The radula is similar to that of the Liotiinae, •

unlike the more modified radula of the Turbininae. II1

Genus HomaIopoma Carpenter, 1864 i

HomaIopoma Carpenter, 1864. Type species (M): Turbo sanguineus Linnaeus, 1758. Mediterranean.

I

Diagnosis. Shell relatively small, solid, spire depressed; juvenile shell umbilicate, adult shell usually I

imperforate. Suture shallow, descending on final whorl; lip thickened in mature shells. Sculpture chiefly

spiral, some species with axial fibs, columella with a moderately strong denticle at base, and another, smaller

denticle a little lower on basal lip. Aperture nacreous white, thin glaze of callus extending on base in advance 1

of aperture. Inner side of operculum corneous, with a multispiral pattern; external surface concave, showing

multispiral pattern at first, final whorl more rapidly expanding and thickened close to columella.

Cephali II c lappets lacking, left and fight neck lobes simple, mouth a vertical slit, lip not slit midventrally. Rachidian tooth of radula uncusped or weakly cusped, lateral teeth 5 pairs, their bases overlapping

those of adjacent laterals; intermediate uncusped tooth (lateromarginal plate) present between laterals and i

marginals; marginals numerous.

Remarks. Homalopoma species are abundantly represented in the northeastern Pacific, particularly

in shallow water (McLean, 1978). Another offshore species in southern California is Homalopoma grippi II

(Dall, 1911), treated by McLean in Keen (1971:350). g

Homalopoma draperi McLean, 1984 i

Figure 1.4B

Homalopoma draperi McLean, 1984:238, fig. 8. _l

Material Examined. California, Santa Maria Basin, Phase I: sta. 6, 109 m (1); sta. 20, 396 m (1). I

Other material: 45 lots in LACM collection.

Description. Medium sized, color usually pink with white flecks on cords; suture deeply impressed.

Early cords fine and numerous, mature cords irregular in strength and spacing; basal sculpture variable, some 1

specimens with 2-3 strong, well spaced cords, others with fine regular basal cords. Parietal callus covering

umbilical chink and forming projecting inner lip; callus extending in advance of aperture on base, forming

distinct glazed area. Height 3-5 mm. I

Type Locality and Type Specimens. Isthmus Cove, Santa Catalina Island, California, 20-30 m.

Holotype: LACM 2001. ill

Distribution. Pacific Grove, Monterey County (37°N), to Cortez Bank, California (33°N). I
I

Habitat. Living on gravel bottoms at 20-100 m, particularly at the Channel Islands off southern

California.

I on matureRemar whorls. ks.ItThis is highly speciesvariisable, charparicteorritozeitsd bdescription y the fme ear having ly cords beenandconfused the irregular with thestrecommon ngth of theshallow- cords

water species Homalopoma luridum (Dall, 1885), which differs in having cords of regular strength and

spacing. zone. This species occurs away from rocky bottoms; it replaces H. luridum on gravel slopes below the kelp

Homalopoma paucicostatum (Daft, 1871)

Figure 1.4C

Leptothyra paucicostatum Dall, 1871: 131, pl. 15, fig. 10.---Oldroyd, 1927:770.

HomaIopoma paucicostatum: McLean, 1978:24, fig. 10.1.

Material Examined. California, Santa Mafia Basin, Phase I: sta. 6, 109 m (1). Other material: 47

lots in LACM collection.

Description. Medium sized, sculpture of 4 strongly elevated spiral cords per whorl, base with 3

additional spiral cords; interspaces broader than cords; axial sculpture wanting; mature lip greatly thickened,

columellar denticle prominent; color usually tan with pink cords; red and white color forms also known.

Height 4.5 mm.

Type Locality and Type Specimens. Monterey, California. Lectotype (here selected): USNM 193796.

Distribution. Salt Point, Sonoma County, California (38°N), to Cabo San Quintin, Baja California

(30°N).

Habitat. Not uncommon on gravel bottoms under kelp; unknown from the intertidal zone.

i Remarks. This species lacks all traces of axial sculpture. It is one of the few species from hard bottoms in the Santa Mafia Basin that can also be collected at sublittoral depths accessible to divers.

Homalopoma berryi McLean, 1964

Figure 1.4D

I Homalopoma berryi McLean, 1964: 132, pl. 24, figs. 7, 8.

collection.Material Examined. Not recorded from Santa Mafia Basin. Other material: 24 lots in the LACM

Description. Shell relatively small, suture moderately impressed. Sculpture of t-me evenly spaced

spiral cords on whorls andbase, suture descending on final whorl,lip not greatly thickened in mature specimens. Color usually dark pink, white in area adjacent to columetla; gray, brown, or white forms also known. Height

3.5 mm.

and Middle Pleistocene, Timms Point Silt, San Pedro, California. TypeLocality TypeSpecimens.

Holotype: LACM 1138.

Distribution. Cordell Bank, Marin County, California (38°N), to Isla Todos Santos, Baja California

I (32ON)"

Itabitat. Rocky bottoms, usually below depths accessible to divers, 20-I 10 m.

1

1
!

Remarks. The shell is smaller and has much finer and more regular spiral cords than the common I

shallow-water species Homalopoma luridum (Dall, 1885). Homalopoma berryi was described from Lower

Pleistocene formations atSan Pedro and Santa Barbara, California; however, living specimens are now known a

from moderately deep water at Cordell Bank and near most of the Channel Islands in southern California. |

Homalopoma cordellensis McLean, new species I

Figure 1.4E

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material I

examined in addition to type lot: LACM 81-50.35, Cordell Bank, off Point Reyes, Matin County, California,

ul

37-52 m, on granite pinnacles; leg. R. W. Schmieder, 1981-1984 (604 shells). LACM 69-35.30, Tanner

Bank, California, 30-37 m; J. H. McLean, 5 October 1969 (11, bryozoan encrusted). LACM 49-141.5; !

Osborn Bank, California, 64 m (1 shell). W

Description. Shell of medium size for genus, spire moderately high; whorls 3.5; periphery rounded; I

suture not deeply impressed, descending on f'mal whorl. Color brick red, strongest on crests of spiral cords, I

interspaces lighter in color. Protoconch diameter 0.3 mm; early whorls rounded, at first with 4 cords, then 5

cords on final whorl opposite aperture. Mature cords on final whorl 11, including cord bordering columella, n

cords of even strength and spacing throughout; interspaces slightly broader than cords. Immature specimens

==

narrowly umbilicate. Mature aperture oblique; parietal area in advance of aperture glazed, red. Columella

with 1prominent tubercle and 1 lesser tubercle below; outer lip weakly lirate within. Nacre of aperture thin, mm

overall

appearing white. Operculum typical for genus, opaque white, thicker where adjacent to columella.

Height 7.2 mm (holotype with operculum in place); height 6.2 mm (live-collected paratype with operculum in

place). Ill

Type Locality and Type Specimens. Unnamed rocky bank, 3 miles WSW of Point Sur, Monterey I

County, California (36°17'N, 121°58'W), 37-55 m, collected by R. W.Schmieder and other divers of Cordell

Expeditions, September 1988 and October 1989; 386 mature specimens and numerous juveniles, of which ill

two mature specimens have the operculum in place. Holotype LACM 2779; 370 paratypes LACM 2780; 5 il

paratypes USNM 887573; 5 paratypes SBMNH 142894; 5 paratypes CAS 102982.

Distribution. Cordell Bank (38°N) to Tanner Bank, Califomia (32.5°N).

Habitat. Rocky bottoms of offshore banks, 30-55 m. I

Remarks. This species differs from the common shallow-water species Homalopoma luridum (Dall,

1885) in having a consistent red color and fewer and more broadly spaced spiral cords (11 cords rather than

13-14 cords). Populations of H. luridum that occur in southern California have smaller shells and color U

variations that include white and red to gray-brown forms, although more northern populations tend to be

larger and more consistently reddish. I

==

I had initially considered that the new species could represent a form ofHomalopoma luridum with

fewer cords that occurs at greater depths. The two species seldom occur at the same station. However, one lot m

from Cordell Bank, collected by Schmieder (LACM 82-50.34, 17 specimens), is identified asH. Iuridum and

suggests that where the two occur together, H. luridum is mature at a much smaller size and has more

m

numerous spiral cords. This is consistent with the observation that other lots identified as H. luridum from

offshore stations throughout the distribution of the species are generally of smaller size than those occurring 1

in shallow water. I

Homalopoma draperi McLean, 1984, differs in its lower profile and more numerous early cords; it t

also replaces H. luridum at depths below the thermocline in southern California. Homalopoma draperi is

more commonly represented offshore in southern California than isH. cordeUensis, which is better represented

from material from central and northern California. I
!

I The name derives from the Cordell Expeditions (organized by Robert Schmieder) to that bank and

other offshore banks. Numerous empty shells were collected at Cordell Bank during four years diving expeditions

I during Sur the fall months of 1981-84, but it was not until the 1988-89 expeditions to the unnamed bank off Point that two live-collected specimens were obtained.

I Family Trochidae Rafinesque, 1815

i oblique, Diagn peritreme osis. complete Shell size ormedium incomplete, to large, final interior lip rarely usuallythickened; nacreous;sculpture umbilicatevaried. or non-umbilicate, Opereulum corneous, aperture

pattern multispirnl on both surfaces, never enveloped by foot. Protoconch with pointed tip. Ctenidium bipectinate,

i with membrane. free tipEpipodium and either variously long or very developed; short dorsal cephaafferent membrane, or monopectinate and lacking afferent lic lappets present or lacking; neck lobes simple, digitiform,

or rolled to form siphons. Radula rhipidoglossate, lateral teeth typically 5 pairs, usually similar to rachidian

i tooth, laterornarginal Biology. Life plate modesweltandeveloped, feeding habits marginalsthenumerous. Trochidae are diverse and differ among the numerous

subfamilies. Some groups live on hard bottoms and feed by grazing on algal films, others live on soft bottoms

I and feeding, ingesthaving mud, while highlyothers modified have become gills forinfaunal the purpose. and derive Somepart grouofpstheir havenutrition becomefromcarnivorous, deposit or suspension feeding on

cnidarians. Differing feeding specializations are reflected in the diverse radular patterns known in the family.

I Remarks. The Trochidae, commonly known as the top shells, are extremely diverse. Numerous genera and species inhabit all oceans, living at all depths, on both hard and soft bottoms. Like the turbinids, the

innermost shell layer is nacreous. They differ from the turbinids in lacking the exterior calcified layer on the

I opereulum. Hickman and McLean (1990) divided this large family into 10 subfamilies, based on characters of the

radula, gill, and external morphology of the epipodium. Unlike the turbinids in which the radular plan is

conservative, the trochid radula has specialized features in each of the subfamilies. Modifications of the epipodium and snout are diverse in the trochids; these features are useful as characters for subfamily

classification. Shell characters axe often convergent between subfamilies and there are many examples of

I similar shell morphology in unrelated groups having widely differing radular and epipodial characters.

I Key to Species of Trochidae

i 1A. Interior not nacreous ......................................................................................... Halistylus pupoideus 1B. Interior nacreous .............................................................................................................................. 2

I 2A. Umbilicus lacking in mature shell 3

2B. Umbilicus present in mature shell .................................................................................................... 7

I 3A. Sculpture coarsely clathrate ............................................................................................................. 4

3B. Sculpture of finely beaded spiral cords ............................................................................................ 5

!

4A. Mature shell height > 40 mm ............................................................................. Bathybembix bairdii

"l 4B. Mature shell height < 30 mm ................................................................................... Cidarina cidaris
!

5A. Surface gray, spiral cords weak to absent ....................................................... CaIliostoma platinum I

5B. Surface not gray, spiral cords strong ............................................................................................... 6

i

6A. Whorls rounded ............................................................................................... Calliostoma turbinum

6B. Whorls nearly flat-sided .............................................................................. Calliostoma variegatum I

7A. Body whorl with strong spiral sculpture ........................................................ SoIarieIla peramabilis

7B. Body whorl smooth ................................................................................................... Solariella nuda I

l

Subfamily Calliotropinae Hickman and McLean, 1990

Diagnosis. Umbilicus open or closed, shell sculpture coarsely cancellate, often with prominent nodes, m

Outer lip simple, columella lacking denticles. Gill with short afferent membrane; snout tip expanded. Base of l

rachidian tooth broad, excavated at sides to interlock with adjacent marginals, lateral teeth often reduced to 3

pairs. m

Remarks. Hickman and McLean (1990) recognized this group at the level of a tribe in the subfamily i

Eucyclinae Koken, 1897. I have elected to follow Waren and Bouchet (1993:11 ) in treating it at the subfamily

level in order to avoid the ambiguity of including an extinct fossil group for which the affinity remains n

speculative. II

Genera of Calliotropinae occur offshore on soft bottoms. The expanded snout tip assists in feeding by

mud-ingestion. I

Genus Bathybembix Crosse, 1893 ill

Bathybembix Crosse, 1893. New name forBemb/x Watson, 1879, not DeKoninck, 1844. Type species (OD): I

Bembix aeola Watson, 1879. Japan.

Diagnosis. Shell moderately large, thin, suture deeply impressed; umbilicus closed in mature shell. I

Sculpture of well-spaced axial and spiral cords, nodose at intersections. Color white under a gray to greenish,

waxy-brown periostracum. Rachidian tooth broadly flanged, bearing numerous fine serrations along sides; Im

lateral teeth 3 pairs; marginal teeth numerous. !m

Remarks. All species occur offshore on soft bottoms, some at abyssal depths. In addition to the

following species, the genus is represented in the eastern Pacific by Bathybembix macdonaldi (Dall, 1890) l

and B. humboldti Rehder, 1971, both of which occur off the western coast of South America. Several species II

occur off Japan.

Bathybembix bairdii (Dan, 1889) I

Figure 1.5A

l

Turcicula bairdii Dall, 1889: 378.--Da11, 1890:346, pl. 7, fig. 3--Oldroyd, 1927:793. (U

Lischkeia bairdii: Abbott, 1974:39, fig. 263. ,,

Ba ,| thybembix bairdii: McLean in Keen, 1971:331, fig. 62.--Hickman and McLean, 1990:82, fig. 44B.

Solariella oxybasis Dall, 1890:352.

|
I

I

I

I

I

I

I

I

I

I

I

I

Figure 1.5. Trochidae.A.Bathybembixbairdii(Dall, 1889): 757 m, offSan Clemente Island, Calffomia(USNM

94865, lectotype); height 50 ram, 2 views. B. Cidarina cidaris (Carpenter, 1864): 265-274 m, SE of

I Santa turbinumCatDall alin,a 189 Islan6:1d,83Cmaliforni

, off Point a (LAConc CMept40-ion, 146.1Santa ); height Barbara23County .3 nun,

,Cal 2 iviews forni.a (USNM C. Callios 1225toma78,

holotype); height 12 mm, 2 views. D. Calliostoma variegatum Carpenter, 1864:73 m, off Long

I Point, Santa Catalina Island, California (LACM 53-69.1), height 22.9 nun, 2 views.

I

Material Examined. Not represented in the voucher collection from the Santa Mafia Basin. Other

i material: D51escriplotstionin. theSuture LACMstrongly collection. channeled, early whorls with 3 nodular carinations connected by fine

axial ridges, middle carination sometimes obsolete; final whorl with prominent peripheral nodes; base with

I fmely beaded lirations; interior nacreous white. Height 40-50 mm. Type Locality and Type Sl_eiraens. OffSan Clemente Island, California, 414 fathoms. Lectotype

(designated by McLean in Keen, 1971:944): USNM 94865. SoIariella oxybasis: 414 fathoms off the Santa

1 Barbara Islands; holotype: USNM 87587.
!

Distribution. Queen Charlotte Islands, British Columbia (54°N), to the Gulf of Tehuantepec, Mexico i

(15°N).

Habitat. Mud bottoms, 350-1400 m. Merriman (1967) reported the gut of Bathybembix to be filled

with mud, as expected in a deposit feeder, although Hickman (1981) described selection for small particles. II

Remarks. The synonym SolarieUa oxybasis was based on a juvenile specimen, i

m

Genus Cidarina DaU, 1908

Cidarina Dall, 1908. Type species (OD): Margarita cidaris Carpenter, 1864. Northeastern Pacific. i

i

Remarks. The genus is monotypic; generic and specific accounts are combined.

1

C/dar/na c/dar/s (Carpenter, 1864) I

Figure 1.5B i

i

Margarita cidaris Carpenter, 1864:653 [cited as "A. Adams, n. s."].

Cidarina cidaris: Oldroyd, 1927:795, pl. 91, fig. 7--Palmer, 1958:137, pl. 17,fig. 13 (holotype).--I-Iickman I

and McLean, 1990:82, fig. 44G. m

Lischkeia (Cidarina) cidaris: Abbott, 1974:39, fig. 264. I

Material Examined. California, Santa Maria Basin, Phase I: sta. 14, 299 m (1 juvenile). Other

material examined: 127 lots in LACM collection.

Description. Suture deeply channeled, early whorls with 3 strongly noded cords, intercalary cords I

appearing on later whorls; whorls fiat-sided at first, mature whorls evenly rounded; base with evenly spaced

spiral cords, strongly beaded near umbilical chink. Color white under a closely adherent, gray, dull-surfaced li

periostracum. Lateral teeth of radula 3 pairs. Height 25-35 ram. i

Type Locality and Type Specimens. Neah Bay, Clallum County, Washington. Holotype: USNM

15600. i

Distribution. Prince William Sound, Alaska (60°N), through the Gulf of Alaska, south to Isla Cedros, i

Baja California (28°N).

Habitat. This species occurs as shallow as 35 m in the northern part of its range, but is usually found i

deeper than 100 m in California, extending to depths of 300 m. The feeding biology of this species has not Ill

been investigated, but may prove to be similar to that of the closely related Bathybembix bairdii.

Remarks. Shells are smaller, thicker, and more sharply sculptured than those ofBathybembix. Cidarina !

i

occurs at shallower depths than Bathybembix. Specimens from California tend to have more closely spaced m

nodes than those from British Columbia.

I

Subfamily Calliostomatinae Thiele, 1924

I

Diagnosis. Shell morphology and sculpture varied. Afferent ctenidial membrane present; cephalic i

lappets present. Left and right neck lobes with simple margins or rolled into siphons. Snout with pseudoproboscis,

a gutter-shaped, tubular extension of mid-ventral lip directed to right. Foot expanding over base of shell.

Rachidian and lateral teeth thin, with long tapered overhanging tips, finely serrate on each side; marginal teeth II

numerous; first marginal strong, with a number of inwardly directed cusps; second marginal of similar shape

but thinner and longer; outer marginals long. I
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I Biology. The diet of Calliostoma, unlike other trochids, has been reported to include hydroids,

gorgonians, and anemones (Perron and Turner, 1978). The large first rnarginals of the calliostomatine radula

I delicate are probably appearance the mosthat t important they probably in cutting; are usedthesolely rachidian, in sweeping. the lateral, and the marginal teeth have such a

Remarks. The subfamily Calliostomatinae has two features that readily distinguish it from all other

I trochid Calliostoma subfamilies: is readilytherecognized enlarged fibyrstitsmarginal beaded sculpture, of the radulbuta there and thearepseudoproboscis. other members ofThe the subfamily familiar genus with

smooth shells--the south boreal group that includes Photinula and related genera. These genera are included

I pseudoproboscis. in the subfamily because they have both the characteristic radula with the enlarged first marginal and the

The function of the pseudoproboscis is not well understood, although Fretter and Graham (1962)

I have suggested that it may assist in grasping the food.

Genus Calliostoma Swainson, 1840

I Calliostoma Swainson, 1840. Type species (SD, Hermannsen, 1846): Trochus conulus Linnaeus, 1758.

Northeastern Atlantic.

I Diagnosis. Moderately high-spired, whorls flat-sided or rounded; f'mallip not thickened. Peristome

incomplete, columella lacking folds or denticles, although somewhat thickened and truncate toward base;

I bbeading ase usually eitherwithpersisting umbilicalor lost depressio in latern growth or openstages. umbilicus. Protoconch Early scwith ulpture reticulate of beaded sculpture spiraland cords, rounded cords pits. and

Color patterns often variegated; interior iridescent, exterior surface often showing metallic luster.

I AlthoughRemtherearksare. many All Call generic-level iostoma have taxaaavailable reticulateaspsubgenera, rotoconch anefforts d earlyto arrange sculpturetheofworldwide three spirmembers al cords.

of the group into a meaningful scheme have not been successful.

I CaUiostoma turbinum Ball, 1896

I Figure 1.5C

Calliostoma turbinum Dall, 1896: 8.----Oldroyd, 1927:792, pl. 100, fig. 1.

I Material Examined. California, Santa Mafia Basin: Phase I, sta. 79, 98 m (1). Phase II, sta. R-8, 90

m, 1juvenile specimen. Other material examined: 14 LACM lots.

I cords peDr ewhorl; scription. cordMs enadium-siz rrow, with ed, ma broadtureinte whorsrpac ls roues,ndupped,ersutur moste chcoarnneled. ds beaded; Spirinalters sculppactuesre with of 4 prin coppciery pal

iridescence, major cords darker brown in radial pattern; basal cords numerous, finely beaded. Height 15-21

! - Type Locality and Type $1_mem. Off Point Conception, Santa Barbara County, California, 100

fathoms. Holotype: USNM 122578.

I Distribution. Point Conception, Santa Barbara County, California (34°N), to Islas San Benito, Baja California (28°N).

i Ha Rembiatart.ks.Rock Theycopp bottery oms,irid70-14 escence 0 m.of the shell surface is the most striking feature of this species.

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Calliostoma variegatum Carpenter, 1864 I

Figure 1.5D

I

Calliostoma variegatum Carpenter, 1864: 652--Oldroyd, 1927: 787, pl. 100, fig. 10---Palmer, I958:141, pl.

17, figs. 9, 10---Abbott, 1974:47, fig. 357.

II

Material Examined. California, Santa Maria Basin, Phase I, sta. 6, 109 mm (1 juvenile). Other II

material examined: 37 LACM lots.

Description. Flat-sided andrelatively high-spired, base ofjuvenile shell def'med by double carination, I

periphery of mature shells tending to become rounded. Sculpture of evenly beaded spiral cords. Color yellowish m

with brown flecks on cords, interspaces on early whorls with greenish or bronze iridescence. Height 18-27 !

Type Locality and Type Specimens. Puget Sound, Washington. Holotype: IJSNM 4201.

Distribution. Forrester Island, southeast Alaska (55°N), to Isla Cedros, Baja California (28°N). I

Habitat. Rocky bottoms 10-60 m; occurring below 60 m in southern California. II

Remarks. Calliostoma variegatum has sculpture rather similar to that of C. annulatum Lightfoot,

1786, a species occurring in shallower water, but does not have the striking color pattern of purple and gold I[

bands of that species. IB

Calliostomaplatinum Dall, 1890 I

Figure 1.6A

II

Calliostoma platinum Dall, 1890:343, pl. 7, fig. 2.--Oldroyd, 1927:788.--Abbott, 1974:48, fig. 362. i

Material Examined. Although not recorded from the Santa Mafia Basin, the area is within the II

distribution of this offshore species. There are 18 lots in the LACM collection from throughout the distribution. HI

Description. Relatively large, thin shelled; early whorls with 2 carinations that become obsolete on I

later whorls; periphery defined by single projecting thread upon which succeeding suture is laid. Spiral sculpture I[

consisting of f'me striae, most strongly developed on base. Height 25-32 mm.

Type Locality and Type Specimens. "Near Santa Barbara Islands, California," 414 fathoms. In

Holotype: USNM 96558. |

Distribution. Moresby Island, Queen Charlotte Islands, British Columbia (53°N), to San Diego,

CaliforniaHabi (33°N). I tat. Soft bottoms, 180-700 m. Feeding biology is unknown, but would be of unusual interest, as

this species is unlike most members of the genus in its soft rather than hardbottom habitat.

Remarks. A related species is Calliostoma titanium McLean, 1984, which has a sturdier shell and I

finely beaded spiral cords; it is known only from the original material from the vicinity of Santa Catalina i

Island and Cortez Bank. Calliostoma chilena Rehder, 1971, from Chile, is very similar to C. platinum.

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I Figure 1.6. Trochidae. A. CaUiostoma platinum DaU, 1890:752 m, off Santa Barbara Island, California (USNM

96558, holotype); height 32 ram, 2 views. B. Solariella peranutbilis Carpenter, 1864:88-93 m,

Cortes Bank, California (LACM 41-112.2); height 16.3 ram, 2 views. C. Solariella nuda Dall,

I 1896:421-695 E of Point, Santa Catalina Island, California (LACM 40-130.5); height

m, Long

11.3 mm, 2 views. D. Halistylus pupoideus (Carpenter, 1864): 22 m, off Point Vicente, Los Angeles

County, California (LACM 19731); height 5.7 ram.

I

I Subfamily Solariellinae Powell, 1951

Diagnosis. Shell umbilicate, whorls rounded, peristome complete; aperture not strongly oblique.

I Afferent ctenidial membrane Cephalic lappets lacking, left and right neck lobes much reduced. Snout

present.

with long papillae around mouth; mouth broad, opening a longitudinal silt; cephalic tentacles expanded at

base. Foot with drawn-out anterior-lateral comers and elongate, tapered posterior end, capable of expanding

I over base of shell. Rachidian, lateral, and marginal teeth of radula with overhanging and pointed tips; rachidian with broad base but not flanged; lateral teeth 2-3 pairs, with denticles only on outer sides; marginal teeth

llumerous.

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Biology. The Solariellinae live offshore on soft bottoms. The external features of the animal, particularly I

the morphology of the snout and the minimal development of neck lobes, are unlike those of other trochids,

modified for a particular kind of food collecting on soft bottoms, as first noted by Fretter and Graham (1977) Im

and treated further by Herbert (1987) and Hickman and McLean (1990). The gill condition, in which the !

afferent membrane is present, is that of advanced trochids; the reduced neck lobes and absence of the cephalic

lappets may therefore be a secondary loss in this group. I

Remarks. All genera have an open umbilicus, an impressed suture and a complete peritreme with I

little slant to the apertural margin. The radula is unique to the subfamily.

The subfamily is widely distributed in all seas and has an extensive fossil record since the Upper

Cretaceous. Herbert (1987) reviewed the numerous southern African members of Solariellinae. I

Genus Solariella Wood, 1842 I

Solariella Wood, 1842. Type species (M): Solariella maculata Wood, 1842. Pliocene, England.

Diagnosis. Shell relatively small, thin, broadly umbilicate; aperture not oblique, nearly circular, I

peritreme nearly complete; umbilicus bordered by beaded cord. Opercular volutions with fine projecting

fringe. Epipodium and radula as described for subfamily.

Remarks. The genus is broadly distributed in world oceans. m

SolarieUaperamabilis Carpenter, 1864 I

Figure 1.6B

II

Solariella peramabilis Carpenter, 1864:653.--Da11, 1921:177, pl. 17, fig. 8.--Oldroyd, 1927:797, pl. 91, I

fig. 8, pl. 101, fig. 7.--Palmer, 1958:138, pl. 17, figs. 3, 4 (Iectotype).--Abbott, 1974:41, fig. 296.

Solariella rhyssa Dall, 1919:360. m

m

Material Examined. California, Santa Mafia Basin: included on soft-bottom faunal check list, but

no voucher examined. Other material: 271 lots in the LACM collection.

II

Description. Shell moderately large for genus; suture impressed, whorls shouldered below suture.

Sculpture of rounded cords on body whorl, base, and within umbilicus; intercalary cords of lesser strength i

arising on final whorl; cords bordering umbilicus strongly beaded; incremental axial sculpture predominating 1

on early whorls. Lateral teeth 3 pairs, marginal teeth relatively few. Color tartwith axial flecks of brown. Shell u

height 10-20 mm.

1

Type Locality and Type Specimens. Santa Catalina Island, California, 30-120 fathoms. Lectotype I

(Palmer, 1958:138): USNM 16281. SolarieUa rhyssa: Catalina Channel, Califomia; holotype: USNM 173803.

Distribution. Forrester Island, southeastern Alaska (55°N), to Baja California, the central and southern m

Gulf of California, to Isla Clari6n, Islas Revillagigedos Islands, Mexico (18°N) l

Habitat. Common on soft bottoms, 50-350 m. Hickman and McLean (1990) illustrated the external

anatomy of this species. Long tentacular processes near the mouth sweep the substrate for detrital feeding on

soft bottoms. l

Remarks. This species is placed in the subgenus Minolia A. Adams, 1860 (type species Minolia

punctata A. Adams, 1860, offshore waters of Japan), because of its very prominent spiral cords. The synonym I

SolarieUa rhyssa was based on a juvenile specimen of 1.7 mm in height on which the axial sculpture is II

prominent, a characteristic feature of juvenile shells.

I

36 m
I

I Solariella nuda Dall, 1896

Figure 1.6C

I Solariella nuda Datl, 1896: 9.--Oldroyd, 1927:799, pl. 91, figs. 2, 5.--McLean in Keen, 1971:331, fig. 70.

I lots in LACM Matericolle al Exction. amined. California, Santa Maria Basin, Phase I, sta. 46, 597 m (1). Other material: 12

Description. Moderately large for genus, uniformly gray-white. Low-spired, with broadly inflated

I and rounded whorls; suture distinct, umbilicus bordered by sharp angulation. Shell smooth, polished except

for fine reticulate sculpture on first 2 whorls. Lateral teeth 3 pairs. Height 9-15 m.

Type Locality and Type Specimens. Off Isla Clari6n, Islas Revillagigedos, Mexico, 460 fathoms.

I Lectotype (designated by McLean in Keen, 1971:944): USNM 122580.

Distribution. Tofmo, British Columbia (49°N), to Isla Clarion, Islas Revillagigedos, Mexico (18°N).

I Habitat. Soft bottoms, at continental slope to abyssal depths, 360-2800 m. Remarks. The smooth sculpture of mature specimens is unlike that of most members of the genus,

but the strongly sculptured juvenile shell is quite typical of the genus.

I

Subfamily Halistylinae Keen, 1958

I Diagnosis. Shell small, thick, lacking interior nacre, sculpture smooth to spiral. Ctenidium lacking

free tip, monopectinate, with enlarged bursicles; eyes on dorsal surface of eyestalk. Radula lacking cusps on

I rachidian andlateral teeth.

Remarks. Hickman and McLean (1990:122) discussed the affinity of the group, which has links to

the subfamilies Lirulariinae and Umboniinae, particularly in its radular characters, but does not have the

I extreme modification of gill that the Lirulariinae

characterizes and Umboniinae.

I Genus Halistylus Dall, 1890

Halistylus Dall, 1890. Type species (OD): Halistylus columna Dall, 1890. Brazil.

I Diagnosis. Shell small, high-spired, whorls rounded, sculpture spiral. Remarks. An additional species, HaIistylus genecoani McLean, 1984, has been described from

central and southern Baja California.

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Halistylus pupoideus (Carpenter, 1864)

I Figure 1.6D

Fenella pupoidea Carpenter, 1864: 656;

I Halistyluspupoideus: Palmer, 1958:142, pl. 19, fig. 4 [syntype].--Abbott, 1974:52, fig. 399 [aspupoides].--

Hickman and McLean, 1990:120, figs. 76A, D, E, 77, 78A.

I Fenella subpupoidea Tryon, 1887:394 [unnecessary new name for Fenella pupoidea Carpenter]. Halistylus subpupoideus: Dall, 1921:174.--Oldroyd, 1927:775.

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Material Examined. Not represented in the Santa Maria Basin voucher collection, but common I

offshore to the north and south. There are 92 lots in the LACM collection.

toward base. Description. Spiral sculpture Shell smofall,raised high-spired, threads whorls of irregular rounded, width, butinterspaces more flattened narrow. nearAperture suture andround, bulginglip I

thickened posteriorly by callus deposition. Color yellowish tan, gray, or pink, some specimens with irregular

gray markings; T early whorls colorless. Juvenile shell with open umbilical area. Height 5.0-6.6 mrn. I ype Locality and Type Specimens. Monterey, California, 20 fathoms. Holotype: USNM 14824.

Distribution. Forrester Island, southeastern Alaska (55°N), to Bahia San Bartolom6 (Bahia Tortuga),

Baja California (28°N). I

Habitat. 20-90 m, common on sand and gravel bottoms.

pupoidesRema A. Adams, rks. Tryon's 1860.replacement name was unnecessary; Carpenter's name is not a homonym of Fenella I

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I Superfamily Cerithioidea

I Family Cerithiidae Fleming, 1822

Diagnosis. Shell tall-spired, usually with flared outer lip and reflexed siphonal canal. Operculum

I paucispiral. Males aphallate; females with open pallial gonoducts.

Biology. Cerithiids are generally understood to be detritus browsers and grazers. As in other groups

that feed at low trophic levels, the species are often abundant in their habitat.

I Remarks. Anatomy for the family was summarized by Houbrick (1988:116), who indicated about 25

genera and hundreds of species, most of which occur in tropical and subtropical waters.

I Key to Species of Cerithiidae

I 1A. Axial sculpture lacking on body whorl ......................................................... Lirobittium paganicum

lB. Axial sculpture present on body whorl ............................................................................................ 2

I

2A. Three primary spiral cords per whorl .............................................................................................. 3

I 2B. Four primary spiral cords per whorl ................................................................................................ 4

3A. Terminal whorl with intercalary spiral cords ..................................................... Lirobittium rugatum

I 3B. Terminal whorl lacking intercalary spiral cords ................................................. Lirobittiumfetellum

I 4A. Whorl profile rounded ...................................................................................... Lirobittium lomaense

4B. Whorl prof'de flat-sided, projecting over suture below ........................................... Lirobittium larum

I Subfamily Bittiinae Cossmann, 1906

I Shell small, with short anterior canal. Diagnosis.

Remarks. Houbrick (1993) reviewed the subfamily andprovided anatomical definitions for the group

i and for six genera that he recognized worldwide. The subfamily is represented in the shallow-water and offshore fauna of southern California by the genus Lirobittium. Houbrick also used Stylidium Dall, 1907, for

the shallow-water species Stylidium eschrichtii (Middendorff, 1849)

I Genus Lirobittium Bartsch, 1911

Lirobittium Bartsch, 1911. Type species (OD): Bittium (Lirobittium) catalinensis Bartsch, 1911; = Bittium

I 1864. Northeastern Pacific. purpureum Carpenter,

Diagnosis. Shell small, tall, whorls moderately inflated; sculpture axial and spiral,usually reticulate;

I sculpture of base spiral; anterior canal very shallow, anal sinus weak, lip thin, smooth. Protoconch of two and one-half whorls, becoming bicarinate early to late on second protoconch whorl.

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Biology. Direct development has been noted for the intertidal species Stylidium eschrichtii I

(Middendorff, 1849)by Strathmann (1987:241). Reproduction in other northeastern Pacific species has not

been studied, and there are no papers that treat the biology of any species in the group.

Remarks. Northeastern Pacific species are unlike those of typicalBittium of the northeastern Atlantic I

and Mediterranean in having sculpture of spiral carinations on the second and third whorls of the protoconch.

Anatomical differences were found by Houbrick (1993), who assigned the species other than Stylidium _l

eschrichtii to the genus Lirobittium Bartsch, 1911. I

Eastern Pacific species were last reviewed by Bartsch (1911), who proposed an excessive number of

names, of which many are now relegated to synonymy. The numerous synonyms are an indication of the high ll

variability among the species. Shallow-water species were reviewed by McLean (1978). The synonymies

given here are based on my study of type material of all eastern Pacific members of the group.

Lirobittium rugatum (Carpenter, 1864)

Figure 1.7A I

Bittium rugatum Carpenter, 1864:539. 1

(Semibittium) rugatum: Bartsch, 1911:397, pl. 56, figs, 4, 5.--Oldroyd, 1927:625.--Palmer, 1

Bittium

1958:179.--Abbott, 1974:106, fig. 1016.

Bittium subpIanatum Bartsch, 1911:395, pl. 57, fig. 5.--Oldroyd, 1927:625, pl. 77, fig. 5. •

Alabina calena Dall, 1919:345.----Oldroyd, 1927:614. l

Material Examined. California: Santa Mafia Basin, Phase I, sta. 40, 392 m (1); Phase II, sta. R-2, •

161 m (1); sta. R-6, 410 m (1). Other material: 247 lots in LACM collection. I

Description. Axial ribs strong to weak, retractive, extending suture to suture; 3 major spiral cords

per whorl, incremental cords added in final whorl. Intersections of cords producing large beads that may 1

project with high relief; whorl sides rounded. Height 7-11 mm.

Type Locality and Type Specimens. Middle Pleistocene, Santa Barbara Formation, Santa Barbara,

California. Holotype: USNM 7154. Bittium subplanatum: Santa Catalina Island, California; holotype: USNM D

160076. Alabina calena: San Luis Obispo Bay, 252 fathoms; holotype: USNM 271070.

Distribution. Off Point Pifios, Monterey County, California (37°N), to Islas San Benito, Baja California

(28°N). I

Habitat. Common on soft bottoms, 30-500 m.

Remarks. Shells vary in the number of axial ribs per whorl and relative strength of the axial ribs. •

Carpenter's name was based on Pleistocene material from Santa Barbara. Hertz (1981 ) illustrated additional l

fossil specimens. The synonym "Alabina'" calena represents a form occurring at depths of 200-300 m, in

which axial sculpture is not developed on the mature whorls. •

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Figure 1.7. Cerithiidae, Turdtellidae. A. Limbittium rugatum (Carpenter, 1864): 95-101 m, off East Point,

I Santa Rosa Island, California (LACM 41-168.8); height 10.9 mm. B. Lirobittium lomaense (Bartsch, 1911): 165-185 m, off Long Point, Santa Catalina Island, California (LACM 39-104.13); height

10.3 mm. C° Limbittiumfetellum (Bartsch, 1911): 64 m, off Santa Barbara Point, Santa Barbara

I County off Whites , Calif Point, orniaLos(LACM Angeles5-2County, 0.8); height California 8.9 mm(LACM . D. Lirobi 48-14.2); ttium height larum 10.5 (Bartsch, ram. F19_,Lirobi .11): 59ttiumm,

paganicum (Dall, 1919): 410 m, off Purisima Point, Santa Barbara County, California (SBMNH

i 142800, Santa Maria Basin, sta. R-6); height 7.8 nun. F. Turritefla cooperi Carpenter, 1864:46-51 m, S of Fraser Point, Santa Cruz Island, California (LACM 41-06.3); height 35.8 ram.

I Lirobittium lomaense (Bartsch, 1911) Figure 1.7B

I Bittium (Lirobittium) lomaense Bartsch, 1911:505, pl. 56, f. 1.---Oldroyd, 1927:636.

asperum

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material

I examined: 11 LACM lots.

Description. Whorl profile rounded, but slightly indented below suture. Axial ribs strong except

below suture, ribs broadly spaced; 4 spiral cords per whorl, subsutural cord weakest. Initial whorl ofpmtoconch

I round. Shell flesh colored under thin periostracum.

Type Locality and Type Specimens. Off Point Loma, San Diego County, California, 71-75 fathoms.

i LectotypeDistribution. (here designatSedan)ta: Cruz USNMIslan19513 d (34°N) 0. , to San Diego, California (33°N).

Habitat. Soft bottoms, 30-100 m, much less abundant that Lirobittium rugatum.

I Remarks. This differs from Lirobittium rugatum in its pink coloration, more broadly spaced axial ribs and lack of incremental spiral cords. This was described initially as a riving form of the extinct Neogene

species Lirobittium asperum (Gabb, 1861), but does not have the even, more broadly spaced axial ribs and

I lacks the strongly carinate protoconch of that species. It is here first recognized as a full species.

I 41
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LirobittiumfeteUum (Bartsch, 1911) I

Figure 1.7C

BittiumfetelIum Bartsch, 1911:409, pl. 51, fig. 4.--Oldroyd, 1927:638, pl. 75, fig. 4.--Abbott, 1974:107. i

Material Examined. California: Santa Maria Basin, Phase I, sta. 85, 113 m (13); Phase II, sta. PJ- i

1,146 m (2); sta. PJ-10, 147 m (2); sta. PJ-18 (3). Other material examined: 58 LACM lots. m

Description. Slender, with rounded whorls; three strong spiral cords intersect with sharply defined I

axial ribs to produce square cancellations. Height 7-9 ram. _l

I

Type Locality and Type Speelraens. Off Santa Catalina Island, California, 16 fathoms. Lectotype

(here designated): USNM 198617.

Distribution. Point Conception, Santa Barbara County (35°N), to Huntington Beach, Orange County, 1

California (34°N).

Habitat. Common on mud bottoms along the mainland coast, 50-180 m, often occurring with l

Lirobittium rugatum and with L. Larum.

Remarks. Differs from Lirobittium rugatum in having more rounded whorls and a more slender

prof'de and completely lacking the spiral threading of the latter species, i

Lirobi_'um larum (Bartseh, 1911) i

II Figure 1.7D

Bittium (Semibittium) larum Bartsch, 1911:407, pl. 57, fig. 4. Ri

Bittium (Lirobittium) Iarum: Oldmyd, 1927:636. Abbott, 1974:107, fig. 1024.

Material Examined. Not represented in Santa Mafia Basin voucher collection, although this species i

is common to the south of Point Conception. Material examined: 136 LACM lots.

u

Description. Relatively large, color medium brown, especially showing at aperture; whorls fiat-sided I

and bulging toward base; 4 major spiral cords per whorl, with minor cords between; axial ribs broadly l

spaced, producing beads andelongate, rectangular cancellations where intersecting with spiral cords. Height

10-14 mm.

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Type Locality and Type Specimens. Off San Pedro, Los Angeles County, California, 4 fathoms. D

Lectotype (here designated): USNM 195156.

Distribution. Gaviota, Santa Barbara County, California (34°N), to south of Bahia Magdalena, Baja I

California Sur (24°N). Usually not occurring near offshore islands.

Habitat. Common on mudbottoms, 30-100 m.

I

Remarks, The elongate cancellations produced by this species in the early whorls are distinctive, i

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I Lirobittium paganicum (Dall, 1919)

i Figure 1.7E

Stylidium paganicum Dall, 1919:345.

I Bittium (Stylidium) paganicum: Oldroyd, 1927:622.

Material Examined. California: Santa Maria Basin, Phase II, sta. R-3, 409 m (1); sta. R-6, 410 m

i (6). Other material examined: type material and LACM 65-38.1, offSan Pedro, Los Angeles County, 640 m; LACM 54-53.1, off Dana Point, Orange County, 366 m.

Description. White, ribs obsolete on final whorl; sculpture of incised spiral lirae. Protoconch worn

I on all speTcypimee nLs.ocalit Height y and8.0Tmm. ype Specimens. Off Point Pifios, Monterey County, California, 292-356 fathoms.

Lectotype (here designated): USNM 271078.

I Distribution. Off Point Pifios, Monterey County (37°N), to Dana Point, Orange County, California (33°N).

Habitat. 360-650 m on mud bottoms.

I Remarks. Dall's type material has not been figured and the identity of this species was uncertain until

the recognition of more recently collected LACM material from southern California. It remains arare species

i known the lectotype), from relatively or, the depo fewsspecimens. its may be lacking Specimens or minimal, may be cobutatedpartially with rusty fillingappearing the spiralmistriations, neral deposgiving its (asthein

appearance of narrow, reddish brown colored bands.

I Family Turritellidae Ltven, 1847

I Diagnosis. Tall shells with numerous whorls; aperture round, lacking anterior or posterior canals;

operculum multispiral. Radula taeniogIossate.

I are furtherBiomodifications logy. Anatomyforissuspen typicallysioncerithi feedingoidean(Houbrick, with aphall 1988:118). ate males and open pallial gonoducts. There

Remarks. Genera of Turritellidae were reviewed by Marwick (1957).

I Genus Turritella Lamarck, 1799

I Turritella Lamarck, 1799. Type species (M): Turbo terebra Linnaeus, 1758. East Indies.

Diagnosis. Outer lip and basal lip forming broad sinus.

I species Turrit Biology. elIaGra commun ham(is1938Riss ) ano,d1Yong 826, ean(194 d it6)hasgavbeee dnetails assuofmethed tfunct hat other ionalspeci morpesholofitgthy isofpathettern. European They

showed that T. communis lies partially buried on soft bottoms and has a ciliary feeding mechanism, using a

I modified to keep large ctenidi particles um to out collecoftthefoodmantle particlcavity. es. Modifications of the mantle form a tentacular screen that serves

Current knowledge of biology and distribution of Turritella species has been reviewed by AUmon

I (1988).

Remarks. Turritella species have notbeenreviewed worldwide and amodem subgeneric classification

remains to be done, although many names are available. Subgeneric characters areprovided by morphology

I of the protoconch and sculpture of the early whorls.
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There are about 15 species in the Panamic Province and a single species living offshore in southern •

California. The Cretaceous and Tertiary formations of the West Coast are rich in species and are useful for

stratigraphic correlation. The fossil species have been given monographic treatment by Merriam (1941) and •

Saul (1983). l

The type species of Turritella is relatively large for the family. Nothing is known of its biology.

I

Turritella cooped Carpenter, 1864 i

Figure 1.7F i

Turritella cooped Carpenter, 1864:655.----Oldroyd, 1927:657.--Palmer, 1958:168, pl. 20, fig. 7.

Turritella jewetti Carpenter, 1864:539.--Oldroyd, 1927:656.--Palmer, 1958:169.--Valentine, 1962:3. R

Turritella orthosymmetra Berry, 1953:412, pl. 28, fig. 5.

Material Examined. Although this species is not represented in the voucher collection from the i

Santa Maria Basin, it is a common species on sand bottoms in southern California. The LACM collections

contains 117 lots from California as well as additional lots from Baja California.

Description. Shell small for genus, thin; spiral sculpture of 2 prominent carinations per whorl and •

fine spiral striae; basal carination projecting and similar in strength to principal carinations; suture laid below I

basal angulation, base smooth except for fine spiral striae. Early whorls with lowermost carination developing

first. Tan with brown axial markings. Height 35-55 ram. H

i

Type Locality and Type Specimens. Santa Barbara, California, middle Pleistocene. According to

Palmer (1958), the type material ofTurritella cooperi has not been located. Turritellajewetti: middle Pleistocene, i

Santa Barbara; type material not located by Palmer (1958:169). Turritella orthosymmetra: Catalina Island, •

50 fathoms; holotype: SBMNH 34534. W

Distribution. Point Pifios, Monterey County, California (37°N), to southern Baja California south of i

Bahia Magdalena (24°N), and the western side of the Gulf of California north to San Felipe. Its presence in i

m

the Gulf of California has not previously been reported.

Habitat. Living populations have not been studied. Common (except in Gulf of California) on sand •

bottoms at 25-100 m. n

Remarks.The two strong carinae may project or be nearly flush with the profile of the shell (as in the

synonym T. orthosymmetra). Merriam (1941) recognized three "varieties" of Pliocene age. Turritellajewetti i

is a probable synonym; it was discussed by Woodring (1946:69) and Valentine (1962).

Turritella cooped is similar in size and occurrence to the North Atlantic species T. communis Risso,

1826, which has been extensively studied, i

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I Superfamily Rissoidea

I Family Rissoidae Gray, 1847

i Subfamily Rissoinae Gray, 1847

Diagnosis. Small to minute mesogastropods with moderately elevated spires; sculpture smooth or

i with various Biolokinds gy. Species of axialofandmostspiral generacords. feedRadula on diatoms taenioglo or micmalgal ssate. films. Most species have a planktonic

larval stage, but direct development is also known in some genera.

I particularlyRemarks. This is a large family, with numerous genera and species, often occurring in large numbers, in shallow water. Ponder (1985) has reviewed genera worldwide and has revised the classification

based on anatomical characters.

I 1865. Two subfamilies were recognized by Ponder (1985): Rissoinae Gray, 1847 and Rissoininae Stimpson,

I Key to Species of Rissoidae

I 1A. Shell height about 2.0 mm ......................................................................................... AIvania rosana

lB. Shell height about 1.2 mm ......................................................................................... Alvania tumida

I Genus Alvania Risso, 1826

I Alvania Risso, 1758]. 1826. Mediterranea Type speci n. es (SD, Nevill, 1885):AIvania europea Risso, 1826 [= Turbo cimex Linnaeus,

Diagnosis. Shell minute, moderately elevated, whorls rounded, suture deep; sculpture clathrate, with

I axial and spiral elements; aperture simple, rounded.

Biology. Like other rissoids, species of Alvania feed on diatoms and microalgae. Eastern Pacific

I species have Remark a planktonic s. The genus larvalisstage. worldwide with numerous species. Ponder (1985:36) treated anatomical

characters and recognized five subgenera worldwide. Eastern Pacific species were placed in the typical subgenus.

I The two species treated here are the only species occurring offshore in southern California.

Alvania rosana Bartsch, 1911

I Figure 1.8A

i Alvania rosana Bartsch, 1911:349, pl. 31, fig. 6.----Oldroyd, 1927:609, pl. 82, fig. 6.--Abbott, 1974:72, fig. 609.

Alvania burrardensis Bartsch, 1921:38.--Bartsch, 1927:30, pl. 3, fig. 5.--Oldroyd, 1927:740.--Abbott,

I 1974:74 (listed only).

Material Examined. California, Santa Mafia Basin, Phase I sta. 17, 654 m (2); Phase I, sta. 73, 98

i m (6). Phase II, sta. PJ-12, 145 m, 34 specimens. Other material: 98 lots in LACM collection.

I 45
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Description. Shell small (maximum length 2.6 mm), whorls rounded, suture deeply impressed, aperture •

slightly less than half length of sheU; protoconch whorls 2.5, smooth. Aperture oval, columella with umbilical

chink. Axial sculpture of narrow, sharply raised ribs and regular spiral cords of lesser strength, producing •

rectangular clathrations; base with spiral sculpture only. II

Type Locality and Type Specimens. Off Santa Rosa Island, California, 48 fathoms. Lectotype

(here designated): USNM 213688. Alvania burrardensis: Burrard Inlet, British Columbia (no depth •

mentioned); holotype: USNM 340938.

Distribution. Kachemak Bay, Alaska (59°N), to Punta San Pablo, Baja California Sur (27°N).

Habitat. Soft bottoms offshore, 20-300 m. I

Remarks. This species is common throughout its distribution. It resembles Alvania compacta

(Carpenter, 1864), which occurs in shallow water (see McLean, 1978:27, fig. 14.4), but has a deeper suture •

and more rounded whorls. |

Alvania tumida Carpenter, 1857 I

III

Figure 1.8B

Alvania tumida Carpenter, 1857:360.--Bartsch, 1911:36 l, pl. 32, fig. 2.--Keen, 1971:370. I

Material Examined. California, Santa Mafia Basin, Phase I, sta. 73, 98 m (7). Phase II, sta. R-4, 92 am

m (1); sta. R-8, 90 m (2). Other material: 98 tots in the LACM collection, i

Description. Shell small for genus (maximum length 1.2 mm), whorls rounded, suture deeply impressed,

aperture length one-third shell length. Protoconch whorls 2, rounded, faintly sculptured. Axial and spiral •

sculpture of fine cords of equal strength and spacing, producing square cancellations. Aperture oval, inner lip II

strongly set off from umbilicus.

Type Locality and Type Speeimens. Cabo San Lucas, Baja California Sur. Holotype: USNM 16206. •

Distn'bution. Santa Mafia Basin, California (34.5°N), to Panama. In the LACM collection this

species is common near Isla Cedros, Baja California, and to the south. There is one specimen in the LACM

collection from near Santa Catalina Island. The material from the Santa Mafia Basin is a major extension to

the north for this typically tropical species. Keen (1971) gave the distribution as Mazatlfin, Mexico, to Panama.

Habitat. Soft bottoms offshore throughout the range, 20-90 m.

Remarks. This is the smallest eastern Pacific species of Alvania, easily recognized by its small size I

and fine clathrate sculpture.

I

Family Vitrinellidae Bush, 1897 I

Diagnosis. Shell small to minute, spire depressed, variously ornamented with axial and spiral sculpture. I

Remarks. This is a large family, with numerous genera, particularly in tropical regions, but very

little is known of the anatomy and biology. Few species are common and most are known only from empty •

shells. Pilsbry and Olsson (1945, 1952) reviewed the genera and species of the tropical eastern Pacific, but

work on the radula and anatomy remains to be done.

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I Figure 1.8. Rissoidae, Vitrinellidae,Caecidae.A.Alvania rosana Battsch, 1911:64-183 m,offNewport Beach,

OrangeCounty, California (LACM 65-63.4); height 2.1 min. B. Alvania tumida Carpenter, 1857:

I 27-34 eschnaurae m, NEBaenxtsch, dIsla1907: Cedros San, BPedro, ajaCali LosfoAngeles rnia (LACMCounty, 71-California 156.21); height (USNM1.2127559, ram. C.holotype); Vitrinella

diameter 2.3 nun, 3 views. D. Caecum crebricinctum Carpenter, 1864: San Diego, California

i CUSNM14930, lectotype);length 6.2 ram.

i Genus Vitrinella C. B. Adams, 1852

I V_trinella C. B. Adams, 1852. Type species (SD, Bush, 1897): VitrineIla helicoidea C. B. Adams, 1852.

Pacific Panama.

I Diagnosis. Shell with low, nearly fiat spire; umbilicus broadly open, exposing whorl below; surface

smooth; axial and spiral sculpture lacking.

I Remarks. Eastern Pacific species are in need of review; SEM work has not been done on shells or radulae. They are sufficiently uncommon to suggest that they may have specific habitat requirements that are

yet unknown.

I V'arinella eschnaurae Bartsch, 1907

I Figure 1.8C

Vitrinella eschnauri Bartsch, 1907:168, fig. 2.---Oldroyd, 1927:819, pl. 106, figs. 5,7,9.

I V_trinella eschnaurae: Abbott, 1974:83 (listed only).

Teinostoma (Pseudorotella)salvania Dall, 1919:369.---Oldroyd, 1927:828.--Abbott, 1974:89 (listed only).

I Material Examined. California, Santa Maria Basin, Phase I, sta. 21, 49 m (1). Phase II, sta. R-8, 90

m (2). Other material: 20 lots in the LACM collection. The voucher material is not mature and can only

tentatively be identified as this species.

I Description. Shell small, spire low, whorls 3, rounded, smooth; aperture circular; protoeonch whorls

1.5, smooth; sculpture lacking, except for growth lines. Maximum diameter 2.3 mm.

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Type Locality and Type Specimens. San Pedro, California. Holotype: USNM 127559. Dali's species •

Teinostoma salvania seems to be based on a smaller specimen of Vitrinella eschnaurae; it came from Isla

Coronado, Baja California (Holotype: USNM 225190). •

Distribution. Monterey, California (37°N), to Isla Asunci6n, Baja California Sur (27°N).

Habitat. Specimens in the LACM collection were taken by divers on rock bottoms at depths of 10-20

m, although not in abundance. The Phase ITspecimens were taken on hard bottoms at 90 m. n

I

Remarks.This species differs from other species of VitrineIla in having a higher shell profile. Other

species in southern California have more nearly flat spires. The name was emended by Abbott (1974) to the am

feminine ending because Bartsch mentioned that the original specimens came from Mrs. Eschnaur's collection. •

Family Caecidae Gray, 1850 I

Diagnosis. Adult shell small to minute, tubular, slender, curved. Protoconch spiral, deciduous. Apical

area sealed with a septum or plug, often with a mucro, or pointed tip. With growth, new septa are formed n

within and earlier shell portions of lesser diameter are shed. Operculum multispiral. Eyes at bases of tentacles,

foot short. Radula taenioglossate.

Biology. Caecids are often extremely abundant in shallow water. The food consists of detritai particles I

and diatoms. Caecids have a planktonic larval stage. Fretter and Graham (1978) described the life habits of

British caecids. m

Remarks. Although there are systematic papers that treat regional faunas, the Caecidae are in major I

need of a worldwide revision. A dozen genera are available but these have not been reviewed and there is little

agreement as to their application. Moore (1962) showed that the family has rissoidean affmities. The shallow- II

water caecid species of southern California were illustrated by McLean (1978). |

Genus Caecum Fleming, 1833 n

iii

Caecum Fleming, 1833. Type species (SD, Gray, 1847): DentaIium trachea Montague, 1803. Northeastern

Atlantic.

Micranellum Bartsch, 1920. Type species (OD): Caecum crebricinctum Carpenter, 1864. Northeastern I

Pacific.

Diagnosis. Shell sculpture offings or annulations that correspond to the round aperture; longitudinal n

sculpture lacking. Mucro of septum projecting, pointed.

w

Remarks. The genus Micranellum was proposed for the relatively large species treated here. Although •

the type species has more numerous annulations, this seems to be a specific level character only. |

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I Caecum crebrieinetum Carpenter, 1864

Figure 1.8D

I Caecum crebricinctum Carpenter, 1864:655.--Abbott, 1974:91, fig. 873.--McLean, 1978:30, fig. 14.1.

i MicraneIIum Caecum magnumcrebricinc "Steams", tum; Bar Tryon, tsch 1920:568.---Oldroyd, 1886, vol. 8:219, pl. 67, 1927:647.--Palmer, f. 83. 1958:174, pl. 20, figs. 10-11.

Micranellum pedroense Bartsch, 1920:569.

I Micranellum catalinense Bartsch, 1920:569.

Micranellum profundicolum Bartsch, 1920:569.

I Micranellum barkleyense Bartsch, 1920:569. oregonense

Micranellum rosanum Bartsch, 1920:569.

I Material Examined. Santa Maria Basin, Phase I, sta. 50, 591 m (1). Phase II, sta. R-l, 91 m (1).

Other material examined: 298 lots in the LACM collection.

I Description. Relatively large for with numerous annulations; mucro sharply projecting. Shell

genus,

cream colored, often with light brown mottling. Length 5-6 mm.

i

14930. Bar TypetschLoca desc

lityribedandsixTypothee

rSpecim taxa froens. m nort SanheDastiegerno,Pacif Califi

o

c

rnia localit. L

ieecs,totasyplist e (her ed inetdesignated) he synonymy: UaSbNMove.

These have not been figured, but I have examined and photographed the type material andall fall within the

range of variation for Caecum crebricinctum. All types are in the USNM.

I Distribution. Kachemak Bay, Alaska (59°N), to Punta Abreojos, Baja California Sur, Mexico (26°N).

Habitat. 10-200 rn, on many different kinds of subsU'ates, including gravel near rocks and f'me

I particles indeeper water. Remarks. This species is the largest species known in the family. It is the only member of the family

occurring in relatively deep water in the northeastern Pacific. Draper (1982) described the living animal of

I Caecum crebricinctum.

I Superfamfly Vanikoroidea

I Family Vanikoridae Gray, 1840

Diagnosis. Shell small to medium in size, white, turbiniform, umbilicate, with large f'mal whorl;

I paucispiral. sculpture axial Radula and spiral. taenioglossate. Protoconch conical, whorls 2-4. Foot with large fleshy epipodial folds. Operculum

Biology. Little is known about the biology of vanikorids. The multispiral protoconch is indicative of

I planktotrophic development.

Remarks. This is a poorly known family with few genera. Wartn and Bouchet (1988:94) expanded

i the concept of the family, adding small-shelled genera previously placed in the Fossaridae. They gave a more

complete diagnosis of the family, with a detailed description of radular characters.

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Key to Species of Vanikoridae I

am

1A. Base lacking broad channel .................................................................. Megalomphalus californicus II

m

lB. Base with broad charmel ........................................................................ Megalomphalus schmiederi

Genus Megalomphalus Brusina, 1871 I

J

Megalomphalus Brusina, 1871. Type species (M): Stomatia azonea Brusina, 1864. Adriatic Sea. 1

Macromphalina Cossmann, 1888. Type species (OD): Sigaretus problematicus Deshayes, 1864. Eocene, 1

Paris Basin.

Chonebasis Pilsbry and Olsson, 1945. Type species (OD): Chonebasisperuviana Pilsbry and Olsson, 1945. •

Peru. II

Diagnosis. Shell small to minute, turbiniform, with small spire and large final whorl; umbilicus 1

broad, funnel-shaped. Protoconch small, elevated, with two and one-half whorls. Foot with projecting epipodial 1

folds (War6n and Bouchet, 1988, fig. 7).

Remarks. Although Dall (1921) had correctly treated the species that follows as Megalomphalus in •

the Vanikoridae, Keen (1971:453) used the genusMacromphalina in the Fossaridae. Gougerot and Le Renard !1

(1981) returned MacromphaIina to the synonymy of Megalomphalus, retaining it in the Vanikoridae. The

type species of Megalomphalus, as figured by War6n and Bouchet (1988, figs. 30-32) is very similar to •

Megalomphalus californicus. II

m

Megalomphalus californicus (Dall, 1903) II

III

Figure 1.9A

Ill

Mac romphalina califo mica Dall, 1903:175.--Abbott, 1974:90, fig. 858.--McLean, 1978:34, fig. 17.5. I

MegaIomphalus californicus: Dall, 1921:166, pl. 14, fig. 7.--Oldroyd, 1927:737, pl. 97, fig. 7. Ill

Material Examined. Not represented in voucher material from Santa Mafia Basin. Other material: 1

20 lots in LACM collection.

Description. Shell small, cream colored, with large final whorl; umbilicus broad, exposing broad II

umbilical wal/; sculpture of numerous, narrow, raised axial ribs, cresting on base and umbilical rim, and f'mer

spiral cords; spiral sculpture anastomosing at later growth stages; protoconch whorls brown. Height 5.5 ram.

Type Locality and Type Specimens. Off Avalon, SantaCatalina Island, California. Holotype: USNM 1

I

109307. aa

Distribution. Santa Cruz Island, California (34°N), to Isla Asunci6n, Baja California Sur (27°b0. •

Habitat. Rocky bottoms, 10-70 m, not common. |

Remarks. This species is here returned to MegalomphaIus, in which it had been placed by Dall

(1921). I

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I Megalomphalus schmiederi McLean, new species

Figure 1.9B

I Material Examined. California: Santa Maria Basin Phase I, sta. 6, 109 m (1); sta. 16, 591 m (1).

LACM materialin addition to the type lot: LACM 40-223.3, W of Santa Barbara Island, California, 274 m

I (1); LACM 41-288.6, Cortes Bank, California, 91 m (1).

Description. Shell small, teleoconch dull white, whorls rounded, slightly tabulate below suture,

i suture deeply impressed. Protoconch projecting, brown, second stage (protoconch ]I) of 1.3 whorls, with spiral ribs apparent on last quarter turn. Aperture oblique; axial ribs raised, strong, wavy, colabral, with

slightly broader interspaces except below suture and toward basal terminus, interspaces compressed at suture

and basal termination. Spiral sculpture weak, producing fine crenulations on back side of axial ribs. Peristome

I complete, columellar lip slightly thickened, straight, narrow, open; nearly smooth,

Umbilicus umbilical wall

umbilicus bordered by broad cord separated from base of shell by channel of same width as cord bordering

umbilicus; axial ribs extending across both channel and cord bordering umbilicus. Height 2.3 ram, diameter

I mm (holotype).

2.0

Type Locality and Type Specimens. Unnamed rocky bank, 3 miles WSW of Point Sur, Monterey

i County, Expeditions, California (36°17'N, 121°58%V),37-55 m, collected by R. W. Schmieder and other divers of Cordell

September 1988 and October 1989.91 specimens recovered from sediment samples. Only one

specimen was live-collected and retains the operculum; it is a LACM paratype rather than the holotype

i because the lip is broken. Holotype LACM 2777; 84 paratypes LACM 2778; 2 paratypes USNM 887574; 2

paratypes SBMNH 142895; 2 paratypes CAS 102983.

Distribution, Point Sur, Monterey County (36°N), to Cortez Bank California (32.5°N).

I be in errHabitat. or. 40-270 m, soft bottoms. The record cited above for the voucher specimen from 591 m may

Remarks. This species differs from Megalomphalus californicus, the only other member of the

I genus known from California, in its smaller size (half the height), less broad profile, more numerous and more closely spaced axial ribs, and in having the deep channel on the base of the shell.

The name honors Robert Schmieder, leader of Cordell Expeditions, whose team conducted the diving

I at type locality.

the

I Superfamily Calyptraeoidea

I Family Calyptraeidae Lamarck, 1809

Diagnosis. Shell of limpet form; outline irregular; with septum or deck formed by columellar border

I aperture. Early traces spiral sculpture strongly or weakly represented or lacking.

of teleoconch with of

Operculum lacking. Head and neck long, flattened, bearing cephalic lappets; tentacles short, eyes at bases.

Mantle cavity deep, filaments of ctenidium modified for filter feeding, tips of filaments overlying food groove

I leading to mouth.

Biology. Calyptmeids are sedentary limpets with unique modifications of the ctenidium and mantle

i cavity for filter feeding. The shell conforms to the area of attachment. All are protandrous hermaphrodites, i starting as male and changing to female as larger sizes are attained.

Remarks. Genera are distinguished by placement of the septum and amount of coiling retained.

I Genera that retain the most coiling in the teleoconch are considered the most primitive in the family.
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Key to Species of Calyptraeidae I

ml

1A. Deck attached on both sides .......................................................................... Crepidula glottidiarum 1B

lB. Deck not attached on left side .......................................................................................................... 2

2A. Exterior lacking coarse ribs .............................................................................. Crepipatella dorsata I

1

2B. Exterior with coarse ribs .............................................................................. CrepipateIla orbiculata

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Genus Crepidula Lamarck, 1799

Crepidula Lamarck, I799. Type species (M): PatelIafornicata Linnaeus, 1758. Northeastern Atlantic. I

Diagnosis. Shell oval, apex at or near posterior margin, coiled stage limited to protoconch; deck

extending across full width of aperture, attached on both sides. 1

Remarks. Species are distinguished by position of the apex, whether atthe margin or raised above it,

and by the curvature of the edge of the deck and whether it projects forward at either edge. Subgenera are

based on the form and position of the deck or septum, and are useful for the placement of fossil forms, n

although they were not utilized by Hoagland (1977). Applicable subgenera for this and the following genus

are mentioned under the remarks for each species.

Crepidula glottidiarum Dall, 1905 U

Figure 1.9C I

llr

Crepidula nivea var. glottidiarum Dall, 1905:26.--Oldroyd, 1927:723.--Hoagland, 1977:377 [under

Crepidula nummaria Gould, 1846]. I

Material Examined. Santa Maria Basin, Phase I, sta. 64, 59 m (2). Other material examined: 12 lots

in LACM collection. •

Description. Shell elongate, about twice aslong as broad, moderately inflated, white; apex at posterior

margin, slightly deflected to right; septum extending half length of shell, undulating on left side; periostracum

light brown, thin, closely adherent. Length 15-23 mm. n

Type Locality and Type Specimens. San Pedro, Los Angeles County, California. Lectotype (here

designated): USNM 183455. aa

Distribution. Point Conception (34°N), to Laguna Beach, Orange County, California (33°N). 1

Habitat. Living on the sides of the burrowing brachiopod GIottidia alba Hinds, 20-70 m. It assumes

a convex form to match that of the outer surface of the Glottidia, but does not grow larger than its host. •

Remarks.This species has usually been dismissed as a form or variety of some other species since it

was first described by Dall. Hoagland (1977:393) placed it under Crepidula nummaria Gould, 1846 [for

which see Abbott, 1974:142, fig. 1571]. I tentatively recognize it here to call attention to the fact that this •

question needs to be investigated by means of life history studies and biochemical genetics. As Hoagland notes

(1977:393), all of the white-shelled, planar species of CrepiduIa from the eastern Pacific are in need of

attention. Typical specimens of Crepidula nummaria have a thick periostracum and are more oval at all •

growth stages. The fact that very small specimens ofC. glottidiarum are relatively elongate suggests that the

species may be more than a situs form of C. nummaria.

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Figure 1.9. Vanikoridae, Calyptraeidae. A. Megalomphalus californicus (Dall, 1903): 8-21 m, Isla Asuncfon,

i Baja California Sur (LACM 67-66.49); height 4.6 ram. B. Megalomphalus schmiederi McLean, new species: 37-55 m, off Point Sur, Monterey County, California (LACM 2777, holotype); height

2.3 ram. C. Crepidula glottidiarum Dall, 1905: San Pedro, California (USNM 183455, syntype);

length 20.8 mm, 2 views. D. CrepipatelIa dorsata (Broperip, 1834): 109 m, off Point Buchon, San

I Luis Obispo County, California (SBMNH 142801, Santa Maria Basin, sta. 6); length 14.9 mm, 2

views. E. CrepipateUa orbiculata (Dalt, 1919): 591 m, off Point Buchon (SBMNH 142802, Santa

Maria Basin, sta. 16); length t3.2 mm, 2 views.

i

i and C. perforans The planar(Valenciennes, species of Crepidula, 1846), areincluding allocated thetomore the subgenus typically lanacus shallow M6rch, water species 1852, for C. nummar/a which the

type species is the Mediterranean C. unguiformis Lamarck, 1767.

I Genus Crepipatella Lesson, 1830

i CrepipatelIa Lesson, 1830. Type species (OD): Calyptraea (Crepipatella) adolphi Lesson, 1830 [=Crepidula dilatata Lamarck, 1822]. Chile.

Diagnosis. Shell broadly oval, apex at posterior margin; coiled stage limited to early juvenile, deck

I attached to shell at left side and posteriorly; fight side of shell interior marked by deep sinus separating deck.

I

I _
I

Remarks. The genus differs from Crepidula in having the deck attached only on the left side. Hoagland I

(1977) placed it in the synonymy of Crepidula. However, following the tradition of previous authors,

Crepipatella is here used as a genus because the deck condition is a clear character that can be recognized in •

fossil forms. 1

Crepipatella dorsata (Broderip, 1834) I

Figure 1.9D

Calyptraea dorsata Brodefip, 1834:38. I

CrepipateIIa dorsata: Keen, 1971:461, fig. 819.

Crepidula dorsata: Hoagland, 1977:373 [discussed but not illustrated], l

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Crepidula lingulata Gould, 1846:160.

Crepipatella lingulata: Keen, 1971:461, fig. 82(P--Abbott, 1974:141, fig. 1554.--McLean, 1978:37, fig. •

18.7. 1

Crepidula biIobata "Gray", Reeve, 1859, pl. 5, fig. 29.

Crepidulafissurata Sowerby, 1883:67, pl. 9, fig. 151. I

I

Material Examined. Santa Maria Basin, Phase I, sta. 6, 109 m (1); sta. 25, 390 m (1). Other

material examined: 830 lots in LACM collection. 1

Description. Shell broadly oval, but irregular in outline, moderately inflated; surface smooth or with

undulating radial fibs; apex at posterior margin; coiled stage limited to early juvenile, deck with raised medial

fold, attached at left side and posteriorly; color mottled brown and white. Length 12-25 mm. !

Type Locality and Type Specimens. Santa Etena, Ecuador. Type material unknown. The species is

better known by its junior synonym Crepidula lingulata Gould, described from Puget Sound, for which the 1

holotype is USNM 5871. 1

I

Distribution. Bering Sea, Alaska, to Bahia Independencia, Peru (14°S).

Habitat. Abundant on rocky substrates from the intertidal zone to 20 m, less common in deeper water •

to 100 m. This is one of the most abundant of eastern Pacific gastropods. 1

Remarks. The name Crepipatella dorsata has previously been applied to forms occurring in the

tropical eastern Pacific, whereas Crepipatella lingulata has been used for the northern forms, but there is no •

evidence that there is a distinction between a tropical and a more northern species. Hoagland (1977:373) 1

placed the taxa C. dorsata and C. linguIata in synonymy, a course of action followed here.

I

CrepipateUa orbiculata (Dall, 1919) 1

Figure 1.9E •

Crepidula orbiculata Dall, 1919:351.

Verticumbo charybdis Berry, 1940:8, pl. 1,fig. 6-10. 1

Crepipatella I (Verticumbo) charybdis: Abbott, 1974:141 (listed only).

Material Examined. Santa Mafia Basin, Phase I, sta. 13, 197 m (5); sta. 16, 591 m (1). Other •

material examined: 3 lots in LACM collection. 1

I

I
i

I Description. Shell of moderate size, smooth or with sculpture of weak to strong divaricating fibs.

Apex close to posterior margin, protoconch with tip immersed, early teleoconcb with f'me spiral sculpture

through rugose sculpture shell diameter of someof 1.3 specimens ram, atthis emerging diametater later theregrowth is a resting stage.stage; Decksubsequent attached half growth lengthsmooth. on rightCoarse side.

Length 10-26 ram.

I Holotype:TyUSNM pe Locali 31100. ty andVerticumbo Type Specharybdis cimens. Victoria, was descriVancouver bed from theIsland, middleBritish Pleistocene Columbia, Timms60Point fathoms. Silt,

San Pedro, California. Paratypes are at the SBMNH, but the location of the holotype is unknown.

I Distribution. Bering Sea, Alaska (Dall, 1921), to San Diego, California (33°N).

Habitat. 100-2100 m, attached to shells or other hard substrata. This is the only calyptraeid in the

northeastern Pacific known from abyssal depths [LACM 64-109, from the Cascadia Abyssal Plain, Oregon,

I 2086 m]. It is an uncommon species in collections, owing to its deep-water habitat.

Remarks. This differs from Crepipatella dorsata in the presence of rugose sculpture (although not in

all specimens) and in having a larger spiral phase of the early teleoconch and a less deeply detached deck on

I the fight side.

Dali's holotype ofC. orbiculata was smooth-shelled and the apical area is worn. Berry (1940) noted

I sculptured (1946) reported and smooth Berry's forms speciesintothebe living Pleistocene off southern speciesCalifornia he described and compared as Verticumbo the morphology charybdis. ofWoodring the deck

to that of Dall's species. Here I synonymize the two taxa and note that the Recent material also varies from

I smooth toThisstrongly sculptured. species is the type species and only known member of the subgenus Verticumbo Berry, 1940,

which is distinguished by its relatively large early teleoconch with fine, sharp spiral cords through a shell

I diameter of 1.3mm_Hoagland (1977:374) did not have sufficient material to determine the status of Verticumbo.

I Superfamily Cypraeoidea

I Family Ovulidae Fleming, 1822

Diagnosis. Shell usually slender, attenuate anteriorly and posteriorly, lip broadly expanded, coveting

I the entire spire. Protoconch cancellate, concealed in mature sheU. Radula taenioglossate. Biology. All species are parasitic on cnidarians, including gorgonians, corals, hydrocorals, and sea

pens. In living animals, the mantle expands over the surface of the shell and often has a color pattern and

I pattern of tubercles and papillae resembling that of the host gorgonian./ndividuals may leave the host in search of uncolonized hosts, but eggs are laid directly on the host. Liltved (1989) provided a useful overview

of anatomy and biology in the family.

1 Remarks. Ovulids are similar to cowries in having the involute spire, but are more elongate and lack ll the denticles on the outer and inner lips.

Cate (1973) reviewed the family worldwide, proposing a number of genera for eastern Pacific species

I previously placed in Neosimnia. However, he did not distinguish his genera from others nor did he provide

any remarks to justify his new taxa. Most of the genera proposed by Care seem to be based on differences in

1 shell characters that are significant only at the species level. Until the Care genera can be re-evaluated, the

I1 species are retained in Neosimnia.

1

I
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Key to Species of Ovulidae I

!

1A. Final whorl inflated; lip edge slightly thickened ......................................... Neosimnia loebbeckeana

lB. Final whorl not greatly inflated; lip edge thick ............................................. Neosirnnia barbarensis

Genus Neosimnia Fischer, 1884 I

Neosimia Fischer, 1884. Type species (M): BuUa spelta Linnaeus, 1758. Mediterranean. 1

Diagnosis. Shell small to moderate in size for family, attenuate anteriorly and posteriorly; inner lip ll

defined by projecting ridge, outer lip with simple thickening.

m

Remarks. Species in California in addition to the two species treated here are Neosimnia aequalis II

m

(Sowerby, 1832), which occurs in relatively shallow water [see McLean 1978:40, asSimnia vidleri (Sowerby,

1881)], and Neosimnia bellamaris Berry, 1946 [see Cate, 1973:72, as Subsimnia beUamaris], which has Im

been reported only from San Diego County, California. 1

Neosimnia loebbeekeana (Weinkauff, 1881) I

Figure 1.10A

Ovula loebbeckeana Weinkauaff, 1881:197, pl. 50, figs. 6, 7. I

Spiculata loebbeckeana: Cate, 1973:82, figs. 182,182a

Simnia loebbeckeana: Abbott, 1974:152 [described, no figure]. I

1

Material Examined. California: Santa Maria Basin, Phase 11,sta. R-5, 154 m (1). Other material:

25 lots in the LACM collection. II

Description. Shell size small to moderate; ivory white, smooth except for fine growth lines and fine U

spiral sculpture near tips. Mature lip thickened, convex; body whorl bulging at midpoint, slightly concave at

either end opposite mature lip; columella weakly plicate and concave anteriorly; spire tip with weak funicle. •

Length 15-22 ram. II

Type Locality and Type Specimens. "Vancouver Island," evidently in error (Cate, 1973:82). Type

material is said by Cate to be in the Museum Loebbeckeana, Dusseldorf. !

Im

Distribution. Monterey, California (37°N) to Bahia Magdalena, Baja California Sur, (25°N).

Habitat. On gorgonians, 40-140 n_ I

Remarks. This is the type species ofSpiculata Cate, 1973. It differs fromNeosimnia barbarensis in II

its smaller size and concave profile at either end.

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I Figure 1.10. Ovulidae, Triviidae.A. Neosimnia loebbeckeana (Weinkattff, 1881): 41 m, off Huntington Beach,

Orange County, California (LACM 49-135.7); height 14.4 mm. B. Neosimnia barbarensis (Dall,

I catalinensis 1892):91 m,);off height Aval23on,mm. SantCaC. aTrivia talinariIstteland, ri Raymond, California1903:74 (SBMNHm,3449 off7W, hol end, otypeAnacapa of NeosIsla imnniad,

California (LACM 50-17.4); height 11.2 mm.

I Neosimnia barbarensis (Dall, 1892)

l Figure 1.10B

Ovula (Simnia) deflexa var. barbarense Dall, in Williamson, 1892:206, pl. 21, fig. 1.

I Simnia (Neosimnia) barbarensis: Oldroyd, 1927:534, pl. 98, fig. 4.

Spiculata barbarensis: Cate, 1973:83, figs. 183 [copy Dall figure], 183a [neotype].

Neosiminia catalinensis Berry, 1916:21, pl. 5, fig. 2.

Ill Simnia (Neosimnia) catalinensis: Oldroyd, 1927:533.

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Material examined:

I 18 lots in the LACM collection.

Description. Shell relatively large, thin, translucent white, broadly inflated, upper portion of whorl

i convex; smooth except for fine growth lines andfine spiral sculpture at the tips; columella concave anteriorly; spire tip with weak funicle. Length 25-30 mm.

Type Locality and Type Specimens. San Pedro, Los Angeles, California. Neotype designated by

I Care Berry(1973:83 Collection ): CA1224 S 32[not 437.nowNeosatimnia SBMNHcataIinensis or CAS]. : Santa Catalina Island, California, 50 fathoms; holotype:

Distribution. Fort Bragg, Mendocino County, California [LACM 71-131] (40°N), to Isla Cedros,

I Baja California [LACM 71-157] (28°N). Habitat. 40-90 m, on gorgonians andthe sea plume Ptilosarcus.

Remarks. This differs from Neosinmia loebbeckeana in its larger size andbroaderbody whorl. The

I synonymy was by (1973).

established Cate

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Superfamily Lamellarioidea I

Family Triviidae Troschel, 1863 I

Diagnosis. Spire low, apertureelongate, inner andouter lips bearing dcnticles.

Biology. Feeding is upon mnicates. Lilt-red (1989) provided a detailed treatment of anatomy and I

biology of the family.

Remarks. There are two subfamilies, Triviinae and Eratoinae. Worldwide species of Eratoinae were

reviewed by Cate (1977) and those of Triviinae by Cate (1979). I

II

Subfamily Triviinae Troschel, 1863 I

Genus Trivia Broderip, 1837

Trivia Broderip, 1837.Type species (SD Gray, 1847): Cypraea europaea Montagu, 1808; northeastern Atlantic.

Diagnosis. Spire concealed, outer lip folded in; sculpture of transverse ridges extending within inner m

and outer lips. il

Biology. As described for family.

Remarks. Cate (1979) proposed a number of genera in addition to those already available. However, 1

Gosliner and Liltved (1987) and Liltved (1989) argued that genera have been based on superficial shell m

characters and recommended that the single genus Trivia be used until anatomical characters have been

evaluated. !

Trivia ritteri Raymond, 1903

Figure 1.10C _l

Trivia ritteri Raymond, 1903:85.---Oldroyd, 1927:536.---Grant and Gale, 1931:754.--Bertsch and Myers, I

1982:96, figs. 2-8. l

Trivia (Dolichupis) ritteri: Abbott, 1974:149, fig. 1637. I!

Decoriatrivia ritteri: Cate, 1979:98, fig. 94. i

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

examined: 39 lots in the LACM collection. 1

Description. Shell of medium size for genus, white, sculpture of about 20 sharp ridges extending

across dorsum to curved outer lip and into aperture on columella, several ribs terminating before reaching 'l

columella; ribs not interrupted by dorsal furrow on dorsum. Length 10-12 mm. !1

Type Locality and Type Specimens. Santa Catalina Island, California, 60 fathoms. Lectotype (selected

by Care, 1979:98): CAS 066533. Im

Distribution. Cordell Bank, California [LACM 81-50.2] (38°N), to Punta San Pablo, Baja California

Sur [LACM 71-176.14] (27°N).

Habitat. 50-150 m, rocky bottoms. This is a characteristic species of rocky banks offshore, usually !

I

in depths below those accessible to divers. g

I
!

I Remarks. This species has usually been allocated to the subgenus Dolichupis Iredale, 1930,

characterized by its weak dorsal furrow, in contrast to the subgenus Pusula Jousseaume, 1884, used for most

i placed of the shallow water eastern Pacific species, in which there is a strong dorsal furrow. However, Cate (1979) it inDecoriatrivia Cate, 1979. Bertsch and Myers (1982) illustrated the radula and treated variation in

size and number of ridges.

I

i Superfamily Naticoidea

Family Naticidae Forbes, 1838

I Diagnosis. SpLrcshort, body whorl large; suturenot deeply impressed; apertureoval to semicircular,

umbilicus broad, often filled with callus, operculum paucispiral. Body expanding to envelop shell; radula

i taeniogloBio ssate. logy. Naticids are well adapted for burrowing in soft sediments and feeding on bivalves, which

they drill with the radula, forming round holes. Eggs are deposited in "sand collars," formed by cementing

I sand grains R with mucus. emarks. Marincovich (1977) monographed the fossil and Recent species of naticids from the

northeastern Pacific. Authors have traditionally been conservative in the usage of such broadly defined genera

asmorePolinices Monffort, 1810. However, I follow the more recent generic revision of Majima (1989), in which taxa are recognized at the generic level. Kabat (1991) reviewed all generic level names in the family; in

many cases he corrected erroneous citations of type designations.

I Nati Four subfamilies are recognized by recent authors: Ampullospirinae, Polinicinae, Sininae, and cinae.

Key to Species of Naticidae

l 1A. Umbilicus sealed with callus pad ....................................................................... Cryptonatica affinis

lB. Umbilicus narrowto broad .............................................................................................................. 2

I 2A. Umbilicus narrow ..................................................................................................... Euspira pallida

i 2B. Umbilicus broad .............................................................................................. CaIinaticina oldroydii

i Subfamily Polinicinae Finlay and Marwick, 1937

Diagnosis. Shell globose; opereulum chitinous, filling aperture.

I (1977:244). Remarks. This is a large diverse subfamily ofnaticids. For a more detailed diagnosis, see Marincovich

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Figure 1.11. Naticidae. A. Euspira pallida (Broderip and Sowerby, 1829): 256 m, SE of Santa Catalina Island,

California (LACM 39-131.6); height 10.8 man. B. Calinaticina oldroydi (Dall, 1897): Off Santa l

Catalina Island, California (CAS 064315holotype); height 35 mm.C. Cryptonatica affinis (Gmelin,

1791): 256 m, SE of Santa Catalina Island, California (LACM 39-131.5); height 9.5 mm. i1

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Genus Euspira Agassiz, 1837 m

Euspira Agassiz, in J. Sowerby, 1837. Type species (SD, Bucquoy, Dautzenberg and Dollfus, 1883): Natica

glaucinoides J. Sowerby, 1812 [non Deshayes, 1832], = Natica labelIata Lamarck, 1804. i

I

Lunatia Gray, 1847. Type species (OD): Natica ampullaria Lamarck, 1822. Tertiary, Europe.

Diagnosis. Shell globose, spire moderately elevated; umbilicus open, umbilical callus weakly developed, li

only partially blocking umbilicus. g

Remarks. Majima (1989:33) and Bouchet and Wartn (1993:767) have treated Euspira as a full

genus, placing Lunatia in synonymy. Species were previously placed in Polinices, but that genus is best left i

for tropical species with glossy shells. Kabat (1991:429) corrected errors in type designation citations of II

previous authors.

It

Euspirapallida (Broderip and Sowerby, 1829) i

Figure 1.11A i

Natica pallida Broderip and Sowerby, 1829:372.

,1

Polinices (Euspira)pallidus: Oldroyd, 1927:728, pl. 97, fig. 9.--Marincovich, 1977:278, pl. 25, figs. 1-6, 8. 1

Lunatiapallida: Abbott, 1974:156, fig. 1693.

Euspira paIlida: Majima, 1989:37, pl. 3, figs. 7-13. i

Material Examined. Not represented in the voucher collection, but common on soft bottoms at

moderate depths. Material examined: 220 lots in the LACM collection, i

60 l
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I Description. Shell of medium size, moderately elevated, umbilicus narrow; whorls flattened below

suture; spiral sculpture of fine striae, axial sculpture of free growth lines; parietal callus white, only partially

I filling umbilicus; Type Locaoperculum corneous, paucispiral, filling aperture. Length 30-40 mm. lity and Type Spedmens. Icy Cape, Alaska, Arctic Ocean. Type material unknown

(Majima, 1989:37).

I Distribution. Arctic Ocean, south to Mexican border; also Japan (Marincovich, 1977:277).

Habitat. Common on soft bottoms; at depths of 100-500 m in southern California. Occurring in

shallower water in the northern partof its distribution; it may occur in the intertidal zone in Alaska.

I Remarks.This is the most commonly occurring member of the family at continental shelf and slope

depths in the northeastern Pacific. Additional synonyms were cited by Marincovich (1977:277).

I Genus Calinaticina Burch and Campbell, 1963

Calinaticina Burch and Campbell, 1963. Type species (OD): Sigaretus oMroydii Dall, 1897. Northeastern

I Pacific.

Diagnosis. Shell large, thin, umbilicus open, partially obstructed by thin parietal wall.

I Remarks. The genus is monotypic. The shell form is intermediate between that of Euspira and

Sinum. A more detailed diagnosis was provided by Marincovich (1977:333).

I Calinaticina oldroydii (Dall, 1897)

Figure 1.11B

I Sigaretus oIdroydii Dall, 1897: 85.--Dall, 1899:85.

Eunaticina oldroydii: Dall, 1921:165, pl. 14, figs. 1, 3.----Oldroyd, 1927:734, pl. 92, fig. 11.

I CaIinaticina oldroydii: Burch and Campbell, 1963:221, fig. 2 [radula].--Abbott, 1974:155, fig. 1686.-

Marincovich, 1977:334, pl. 31, fig. 3.

i Material Examined. Not representedin the Santa Maria Basin voucher collection. Other material: 32 lots in the LACM collection.

Description. Shell relatively large, thin; whorls 4; spire elevated; umbilicus narrow; parietal wall

I thin, edge straight, to partially umbilicus; spiral sculpture striae; sculpture

reflected obstruct of fine axial of

growth increments; light brown under thin brown periostracum; periostracum shaggy within umbilicus;

operculum comeous, paucispiral, filling aperture. Length 40-70 mm.

I Type Locality and Type Specimens. Santa Catalina Island, California, "in deep water." Holotype:

CAS 064315.

I Baja CalifDistribution. ornia [LACMYa3ch9-a114.9 ts, Linc ] (2ol8°nNC).ounty, Oregon [LACM 61-47.2] (44.5°N), to Islas San Benito,

! Habitat. Soft bottoms, 30-400 m.

I Remarks. Specimens inthe LACM collection arewell represented throughout the of the species. range

The inner and outer lips are sharp and thin in growing specimens, becoming thicker and stronger in mature

specimens.

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Subfamily Naticinae Forbes, 1838 i

Diagnosis. Shell globose, mostly smooth, spire low to moderately elevated, umbilicate, parietal callus

well developed; operculum with external calcareous layer.

Remarks. This is primarily a tropical group; the genus Cryptonatica is an exception.

i

Genus Cryptonatica Dall, 1892

Cryptonatica Dall, 1892. Type species (SD, Cossmarm, 1896):Natica (Cryptonatica)floridana Dall, 1892. I

Tertiary, southeastern United States. II

Sulconatica Golikov and Kussakin, 1974. Type species (OD): Natica janthostoma Deshayes, 1839.

Northwestern Pacific. I

Boreonatica Gotikov and Kussakin, 1974. Type species (OD): Natica clausa Broderip and Sowerby. Northern

Pacific.

Diagnosis. Shell globose, smooth, umbilicus filled by thick smooth, semicircular callus; operculum i

smooth, concave, outer edge marked by narrow furrow. i

Remarks. Dali's original proposal of the genus was not accompanied by a valid type designation.

Although Dall's (1909) type designation ofNatica clausa has been cited by recent authors, Kabat (1991:428)

noted that an earlier designation by Cossmann (1896) takes precedence. The concept of the genus does not n

change, however. Bouchet and War6n (1993:758 ) provided the most recent and complete account of the genus |

and treated three species. A single species is represented in the northeastern Pacific.

Cryptonatica affinis (Gmelin, 1791) I

dim

Figure 1.11C i

m

Natica affinis Gmelin, 1791: 3675.

Cryptonatica affinis: Bouchet and War, n, 1993:763, figs 1789-1792, 1810-1812, 1817-1821, 1840, 1846, /

1877, 1891, 1905. II

Natica clausa Broderip and Sowerby, 1829: 372. m

Natica (Cryptonatica) clausa: Dall, 1921:163, pl. 14,fig. 1.---Oldroyd, 1927:724, pl. 97, fig. 2.--Marincovich, B

1977:410, pl. 41, figs. 7-10; pl. 42, figs. 1-6.

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material: B

232 lots in the LACM collection. D

Description. Shell small to medium-sized, white under tan periostracum; spire moderately elevated, In

umbilicus f'flled by white callus pad, operculum with thin white calcareous layer, marked by groove along l

outer edge. Length 10-60 mm.

Type Locality and Type Specimens.Iceland, type material unknown (Bouchet and War6n, 1993:763). ll

Natica clausa: type locality and type material unknown. II

Distribution. Arctic Ocean, circumpolar; south in the eastern Pacific to San Diego (33°N); south in

the North Atlantic to the Bay of Biscay, France, and North Carolina (Bouchet and War6n, 1993). I

Habitat. Soft bottoms, from 0-200 m in the Arctic; at greater depths near southern range limits.

There are 68 lots from southern California in the LACM collection, occurring at depths of 80-400 m.

l
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I Remarks. Natica affinis is a common member of the offshore assemblage in southern California.

Specimens are small at maturity (length to 15 mm), compared to those from northern localities, which may

i approachThis a length of 60 mm. species is better known to authors treating the eastern Pacific under the name Natica clausa

Broderip and Sowerby, 1829, from an unknown type locality, but Bouchet and War6n (1993:763) showed that

i usage Mrincovich of the older (1977:410), name iswhounavoidable. cited numerous It hasynonyms s been 1Teated from intypemorelocalities detail inbytheBouchet Arctic, and NorthWar6n Atlanticandandby

North Pacific.

Superfamily Janthinoidea

I Family Epitoniidae Berry, 1910

i Diagnosis. Shell usually white, with high spire, deep suture and rounded aperture. Sculpture predominantly axial; spiral elements if present of lesser strength. Radula ptenoglossate, consisting of numerous

similar teeth in the row.

I tropical species Biology.occur All omemn corals. bers Living of the fspecimen amily ares may parasitic occasionally on crtidarian be collected s, chieflyawayanemfromones,thealt hosthough anemones some

when they are searching for new hosts. The proboscis is acrembolic and they are hermaphroditic and aphallic,

I prod the rasping ucing spermat and cutting ozeugmata of the cnidarian or sperm prey. packets. The numerous fanglike teeth of the radula are designed for

Remarks. Epitoniids are commonly known as wentletrap shells. Northeastern Pacific species were

t reviewed by DuShane (1979). Bouchet and War6n (1986) discussed classification at the generic level in their review of the North Atlantic species.

I Key to Species of Epitoniidae

I 1A. Axial thick NodiscaIa

costae spongiosa

1B. Axial costae thin .............................................................................................................................. 2

I 2A. Axial costae rounded in profile ........................................................................................................ 3

2B. Axial costae projecting at shoulder .................................................................................................. 4

!

3A. Axial costae > 25 per whorl .................................................................................... Epitoni_ berryi

I 3B. Axial costae < 20 per whorl ........................................................................... Epitonium indianorum

i 4AB.. Axial i costae back projecting, -foldednot .......................................................................................... back-folded .......................................................................................... Epitonium lowei5

I 5A. Axial costae 8-14 per whorl ................................................................................... Epitonium hindsii

5B. Axial costae 14-21 per whorl ............................................................................... Epitonium sawinae

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Genus Epitonium Rrding, 1798 I

Epitonium Rrding, 1798. Type species (SD, Suter, 1913): Turbo scalaris Linnaeus, 1758. Indo-Pacific. lit

Diagnosis. Shell with high spire and deep suture, lacking a basal disk. Axial sculpture of thin, projecting 1

costae, spiral sculpture lacking or of fine striae.

Remarks. DuShane (1979) placed species with rounded whorls and no spiral sculpture inNitidscala i

DeBoury, 1909 (type species Scalaria unifasciata G. B. Sowerby IT,West Indies), which differs from the type Ill

species of Epitonium only in its more slender profile. Those with fine spiral sculpture were placed inAsperiscala

DeBoury, 1909 (type species Scalaria bellastriata Carpenter, 1864, California). Authors prior to DuShane II

(1979) had used these taxa at the subgeneric level. In treating the North Atlantic species, Bouchet and Warrn II

(1986) ignored all subgenera and placed allspecies with rounded whorls, varying expression of spiral sculpture

and lacking a basal disk inEpitonium. Here I return the northeastern Pacific species to Epitonium, mentioning

the appropriate subgenus in the remarks under each species. II

This treatment of the offshore species of Epitonium includes allthat are known fxom southern California.

DuShane (1979) did a monumental job of reducing the excessive number of unillustrated taxa described by 1

Dall in his declining years. Revisions to the synonymy of two of the species treated by DuShane (E. berryi and II

E. sawinae) are made here.

I

Epitonium berryi (Dall, 1907)

Figure 1.12A I

Scala berryi DaU, 1907:127.

Epitonium (Nitidiscala) berryi: Oldroyd, 1927:361. 1

II

Epitonium (Epitonium) berryi: Abbott, 1974:123 [listed only].

Scala rectilaminata Dall, 1907:127. I

"'Epitonium (Nitidiscala) inconspicua G. B. Sowerby II, 1847".---of Dall, 1921:116. U

Epitonium ["group" Nitidoscala sic] sawinae vat.? catalinense Dall, 1917:481.

Nitidiscala catalinense: DuShane, 1979:110, fig. 28 [holotype]. I

Material Examined. Santa Maria Basin, Phase I, sta. 20, 396 m (1). Other material: 38 lots in the

LACM collection. I

Description. Shell white, up to 10 whorls; costae thin, low, numerous, not spinose, up to 30 per

whorl; suture deep, base rounded. Length 10-22 mm. all

Type Locality and Type Specimens. San Pedro Bay, Los Angeles County, California, 200 fathoms. I

Holotype: USNM 107724. Scala rectilaminata: Monterey Bay, California, 12 fathoms; lectotype (here

designated): USNM 11048. Epitonium catalinense: Santa Catalina Island, California; lectotype (here II

designated): USNM 109502. U

Distribution. Vancouver Island, British Columbia (49°N), to Isla Todos Santos, Baja California

(32°N) (DuShane, 1979:111).

Habitat. Soft bottoms, 20-360 m (DuShane, 1979:111). II

Remarks. This species is characterized by its numerous low, closely spaced costae. ,Ira

The name of the species has been confused in the literature owing to the factthat the holotype of Scala !

berryi is a juvenile specimen, as is the holotype ofS. rectiIaminata. Mature specimens were not seen by DaU

u

until the description of Epitonium catalinense by Dall (1917). DuShane (1979:110) incorrectly used the i

junior "Nitidiscala" catalinense Dall, 1917, for this species. I

synonym
I

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I Figure 1.12. Epitoniidae. A. Epitonium berryi (Dall, 1907): Off Santa Catalina Island, California (USNM 109502,

holotype of Epitonium sawinae var. catalinense); height 13.5 mm. B. Epitonium hindsii (Carpenter,

I 1856): San Pedro, Los Angeles County, California (USNM 46222; height 18.2 mm. C. Epitonium indianorum (Carpenter, 1864): 12-38 m, Carmel, Monterey County, California (LACM 60-24.73);

height 37.8 mm. D. Epitonium sawinae (Dall, 1903): Off Newport, Orange County, California

(USNM 110638, holotype of Epitonium acrostephanus); height 20 ram. E. Epitonium lowei (Dall,

i t906): 210 m, off E1 Capitan, Santa Barbara County, California (LACM 118106); height 30.5 ram. FoNodiscala spongiosa (Carpenter, 1864): 46-49 m, E of Santa Barbara Island, California (LACM

39-79.9); height 9.8 nun.

i

I Epitonium hindsii (Carpenter, 1856)

Figure 1.12B

i Scalaria hindsii Carpenter, 1856:165.

Nitidiscala hindsii: DuShane, 1979:111, figs. 32-35.

I Epitonium ["group" Nitodoscala sic] persumrum Dall, 1917:478.

Epitonium contrerasi Jordan and Hertlein, 1926:446, pl. 30, fig. 4.

I Epitonium cooperi Strong, 1930:194, pl. 20, fig. 6-8.

Material Examined. Not represented in the voucher collection, but occurring offshore in southern

I California. Material examined: 73 lots in the LACM collection.

Description, Shell white, maximum size about 22 ram, up to 11 whorls; costae thin, low, 8-14 per

whorl, not spinose; suture deep, base rounded.

I

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Type Locality and Type Specimens. Panama. Lectotype (designated by DuShane, 1979:112): NHML n

1963.21. Epitonium persuturum: San Diego; holotype: USNM 211021. Epitonium contrerasi: Bahia Tormga;

Baja California; holotype: CAS 2121. Epitonium cooperi: San Pedro, Los Angeles County, California; holotype: 1

SDNHM 345. I

Distribution. Forrester Island, Alaska (55 °N), to Peru (14° S) (DuShane, 1979:112).

Habitat. Intertidal to 195 m, on soft bottoms (DuShane, 1979: 112). I

m

Remarks. This species is characterized by the low number of slightly spinose costae perwhorl. The

synonymy cited above follows that of DuShane (1979:111 ). /

II

Epitonium indianorum (Carpenter, 1864)

Figure 1.12C I

Scalaria indianorum Carpenter, 1864:660. 1

Epitonium (Nitidscala) indianorum: Oldroyd, 1927:355.--Palmer, 1958:186, pl. 20, figs. 23, 24. I

Epitonium (Epitonium) indianorum: Abbott, 1974:119, fig. 1233.

Nitidiscala indianorum: DuShane, 1979:113, figs. 36-38. I

Epitonium ["group" Nitidoscala sic] columbianum Dall, 1917:481.

Epitonium ["group" Nitidoscala sic] montereyensis Dall, 1917:481; not Scala (Cirsotrema ) montere yensis

Dall, 1907. II

Scalaria regiomontana Dall, in DeBoury, 1919:39 [new name for E. montereyensis Dall, 1917].

Material Examined. Not represented in the voucher collection, but occurring offshore in California. n

Other material: 84 lots in the LACM collection.

Description. Shell white, relatively large, up to 10 whorls; costae 10-17 per whorl, low, strong,

continuous whorl to whorl, usually lacking small spine at shoulder; suture deep, base rounded. Length 15-38

mllL

Type Locality and Type Specimens. Neah Bay, Clallam County, Washington. Lectotype (designated n

by Palmer, 1958:187): USNM 15521. Epitonium columbianum: Off Columbia River, Oregon, 27 fathoms; I

holotype: USNM 111211. Epitonium montereyense: Monterey Bay, California, 30 fathoms; holotype: USNM

111217. I

Distribution. Forrester Island, Alaska (55°N), to Bahia Todos Santos, Baja California (32° S)

(DuShane, 1979:113).

Habitat. Intertidal to 120 m, soft and rocky bottoms. Occurring below 30 m in southern California. I

Remarks. This species is characterized by its large size and moderate number of rounded costae

without spines. Itis the largest species of Epitonium occurring offshore in southern California. The synonymy n

indicated here follows that of DuShane (1979:113). r|

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I Epitonium sawinae (Dall, 1903)

Figure 1.12D

I Scala sawinae Doll, 1903:175.

i Epitonium Epitonium (Nitidiscala Epitonium)) sawinae sawinae:: Abbott, Dall, 1921:115, 1974:119,pl.fig. 6, fig. 1237.12.---Oldroyd, 1927:358, pI. 31, fig. 2.

Nitidiscala sawinae: DuShane, 1979:115, figs. 42-44 [not figs. 39, 40].

I Epitonium (Crisposcala) acrostephanus Dall, 1908:251.

Epitonium (Crisposcala) catalinae Dall, 1908:252.

I Epi Epitonium tonium ["group" Nitodoscala, sic] tabulatum D011, 1917:482. ["group" NitodoscaIa, sic] regime Dall, 1917:484.

I 91 m (3)Material ; sta. R-4,Examined. 92 m (1); sta.CaR-8, liforni90a:mSanta (1). Oth Mari eramaBastein, rial:Phase 340 Ilo,tsstain. 42the, 1LACM 00 m (1c);olPhlectio asenII. , sta. R-I,

Description. Shell white, medium to large, up to 10 whorls; costae 14-21 per whod, strong, with

I pron 15-2o4uncmm. ed shoulder spine, producing tabulate profile, costae often broad and fused with those behind. Length

Type Locality and Type Specimens. Santa Catalina Island, California, 16fathoms. Lectotype (here

I designated): USNM 109309. Epitonium acrostephanus: Newport, Orange County, Colifomia; lectotype (here designated): USNM 110638. Epitonium cataIinae: Santa Catalina Island, California, 16 fathoms;

holotype USNM 198628. Epitonium tabuIatum: Islas Coronados, Baja California, 16 fathoms; holotype:

USNM 109569. Epitonium Point Reyes, Matin County, California, 61 fathoms; holotype: USNM

regum:

206576.

Distribution. British Columbia (54°N), to Bahia Magdalena, Baja California Sur (24°N) (DuShane,

i 116).

1979:

Habitat. 18-360 m on soft bottoms (DuShane, 1979:116).

I costae that Remararke ofts.This en folded specback. ies is cThhaerasucteturirezeids bdyeeiper tsdethan ep sutture hat ofanEd.tabberryi ulate. aspect produced by the spinose

Changes are made here to the synonymy of this species given by DuShane (1979:115): Epitonium

I catal removed. inae Doll, 1908, is added to the synonymy, and E. berryi and E. rectilaminatum, both Doll, 1907, are

I Epitonium lowei (Dall, 1906)

Figure 1.12E

I Scala lowei Doll, 1906:44.

Epitonium (Asperiscala) lowei: Oldroyd, 1927:354, pl. 31, fig. 11.

I Aperiscala lowei: Abbott, 1974:123. fig. 1290.

Asperiscala lowei: DuShane, 1979:101, fig. 9.

I Material Examined. California: Santa Maria Basin, Phase lI, sta. R-2, 161 m (1); sta. R-4, 92 m

(1); sta. R-8, 90 m (1). Other material: 18 lots in the LACM collection.

I
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Description. Shell white, medium in size, up to 9 rapidly expanding whorls, umbilicus deep; costae I

25-32, slightly spinose at the shoulder, with some costae larger than the others; spiral sculpture of raised

spiral ridges; sulci between ridges twice width of ridges. Length 15-25 mm. It

Type Locality and Type Specimens. Off Avalon, Santa Catalina Island, California, 40-60 fathoms. II

Holotype: lost (DuShane, 1979); paratype USNM 191548.

Distribution. Santa Maria Basin, California (35°N) to Bahia Magdalena, Baja California Stir (24°N), E

and the Gulf of California. DuShane (1979) placed the northern limit at Anacapa Island, California, so the

present material constitutes a significant range extension. DuShane's (1979) record of the species from Panama

was based on a specimen of 5 mm in length and needs to be confirmed by specimens of larger size. I

U Habitat. Soft bottoms, 25-170 m (DuShane, 1979:103).

Remarks. This species resembles the Epitonium bellastriatum (Carpenter, 1864), which occurs in m

shallower water, but is larger and more slender. These two species are assigned to the subgenusAsperiscala,

|

characterized by the spiral sculpture.

Genus Nodiscala DeBou_, 1889 I

i

Nodiscala DeBou_, 1889. Type species (OD): Scalaria bicarinata Sowerby, 1844. Phi_ppines.

Diagnosis. Small, slender, with serrate sumxe defined by temfinus of a_al ribs; spiral sculpture E

freely punc_te, ape_ure senfilunate, basal disk poorly developed.

Remarks. Pdthough usually _-eated as a subgenus ofOpalia H. and A. Adams, 1853, in which there I

is a well-marked basal disk, Nodiscala is sufficiently d_erent in s_e and sculp_LretOquarry as a _U genus.

g

DuShane (1974) recognizedNodiscala as a subgenus, but she used only Opalia for the Nodiscala species in

her subsequent paper 0)uShane, 1979). I

Nodiscala spongiosa (Carpenter, 1864)

Figure 1.12F E

Opalia spongiosa Caqaenter, 1864:660.--DuShane, 1979:126, figs. 62, 63. I

Epitonium (Nodiscala) spongiosum: Ol_oyd, 1927:351. II

Opalia (Nodiscala) spongiosa: Palmer, 1958:193, pl. 22, figs. 1-3, 6-8.--DuShane, 1974:69, figs. 138, 139. m

OpaIia (Dentiscala) spongiosa: Abbot, 1974:116, fig. 12_. !

Opalia retiporosa Carpenter, 1864:660.

Opalia (Nodiscala) retiporosa. Ol_oyd, 1927:352. I

Material Exam_ed. Ca_fonfia: Santo Mafia Basin, Phase I, sin. 16, 591 m (1). Other material: 58

lots in the LACM collection. ,_

Description. Shell _ght brown, smaU; whorls 8-9; a_al ribs 10-15, s_ong, crenulafing suture, fibs I!

undulating, occasionM fibs larger, fomfing vafices; spiral sculp_re of rows of time puncmtions; Final Up

thickened. Len_h 7-13 mm.

Type Loeafity and Type Speehnens. Monterey, Calffonfia. Holo_pe: USNM 14830. Opalia

retiporosa: San_ Cata_na Island, CMffonfi_ 40 fathoms; holo_pe: USNM 11843.

D_Hbufion. Monterey, Cahfon_a (37 °N), to the Gulf of California, southern Mexico, and the

Galapagos Islands (DuShane, 1979:126).

w

Habitat. Rocky and gravel bottoms, 18-72 m (DuShane, 1979:126). Ill

Remarks. The depth record of 591 m for the voucher specimen may be erroneous. II

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I Superfamily Eulimoidea

I Family Eulimidae Troschel, 1853

Diagnosis. Shell white to brown, usually glossy and smooth; often showing scars from earlier positions

i of Radula outerlacking lip; aperture in most lacking genera. siphonal canal; protoconch smooth, of 2.5-4 whorls; operculum usually present.

Biology. Parasitic on echinoderms, attached by the snout or proboscis, by which they extract body

I fluids. Individuals found away from their hosts are thought to be resting between feeding or searching for new hosts. Development in most eulimids is planktotrophic. Genera are usually restricted to one class of echinoderms;

some species are known only from single host species.

I Remarks. Most species of eastern Pacific Eulimidae were treated by Bartsch (1917). Few changes

were made in the classification of the group until War6n (1984) published his generic review, which has

enabled a complete reassessment of the genera in use for the eastern Pacific species. I

i Key to Species of Eulimidae

1A. Whorls markedly inflated ................................................................................................................. 2

i lB. Whorls flat-sided or nearly so .......................................................................................................... 3

2A. 2B. Outer Outer liplip projecting not projecting....................................................................................... ....................................................................................... Pseudosabinella HaIiella abyssicola bakeri

I 3A. Shell axis nearly straight 4

3B. Shell axis flexed ............................................................................................................................... 7

I 4A. Shell with brown spiral lines .................................................................................. Eulima raymondi

4B. Shell white ....................................................................................................................................... 5

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5A. Whorl sides straight .............. '............................................................................. Polygireulima r_tila

I 5B. Whorls faintly inflated ..................................................................................................................... 6

6A. Breadth of shell about 4 mm ........................................................................................ Balcis micans

I 6B. Breadth of shell under 3 mm ................................................................................... Balcis oldroydae

I 7A. Mature shell < 3.5 mm in length .................................................................................. V_treolina yod

7B. Mature shell > 4.5 mm in length ...................................................................................................... 8

I 8A. Body whorl more strongly inflated than other whorls ..................................... Vitreolina columbiana

8B. All whorls very faintly inflated ............................................................................... _treolina macra

i

I 69
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Genus Eulima Risso, 1826 I

Strombiformis Da Costa, 1779 [rejected, ICZN Opinion 1718]. m

Eulima Risso, 1826. Type species (ICZN Opinion 1718): Turbo subulatus Donovan, 1803. Southwestern m

Europe.

J

Diagnosis. Shell tall, slender, with color pattem of brown spiral bands; whorls flat-sided, aperture 1

elongate; raduta ptenoglossate.

il

Biology: Waren (1984:45) noted that the European Eulima bilineata is parasitic on ophiuroids. roll

However, there are no records of hosts for eastern Pacific species, n11

Remarks. The tall shell with flat-sided whorls, elongate aperture and color pattern of brown lines

characterize this genus. The genus was also discussed by Bouchet and War6n (1986:318) and War6n (1992:179). I

,u

Eulima raymondi Rivers, 1904

Figure 1.13A I

Eulima raymondi Rivers, 1904:70, fig. 2 (Pleistocene). i

Strombiformis raymondi: Grant and Gale, 1931:864. 1

Strombiformis riversi Bartsch, 1917:339, pl. 45, fig. 3 (Pleistocene). ml

Strombiformis californica Bartsch, 1917:340, pl. 45, fig. 5.----Otdroyd, 1927:380, pl. 40, fig. 5.--Abbott, U

1974:127, fig. I395.

Strombiformis lapazana Bartsch, 1917:341, pl. 46, fig. 3. I/

Strombiformis townsendi Bartsch, 1917:340, pl. 46, fig. 4.

Material Examined. California: Santa Mafia Basin, Phase I, sta. 21, 49 m (1); sta. 64, 59 m (1); m

Phase II, sta. R-5, 154 m (1); sta. PJ-6, 148 m (I); sta. PJ-7, 123 m (1). Other material: 115 lots in the LACM II

collection.

Description. Shell tall andnarrow,length to 12 ram; whorls fiat-sided, aperture extremely elongate; I

light brown with 2 narrowdark brown bands on final whorl, lowermost covered by growing lip so that only n

single band shows on spire whorls.

Type Locality and Type Specimens. Santa Monica Canyon, Pleistocene. Holotype not located.

Strombiformis riversi: Santa Monica Canyon, Pleistocene; holotype: USNM 251390. Strombiformis I

californica: San Diego Bay; holotype: USNM 249619. Strombiformis lapazana: La Paz, Baja California

Sur; holotype: USNM 211388. Strombiformis townsendi: La Paz, Baja California Sur; holotype: USNM 1

252391.

Distribution. Monterey Bay, California [LACM 60-23] (37°N), to Bahia Guatulco, Oaxaca, Mexico

[LACM 38-9] (16°N). '!

'1i

Habitat. Soft bottoms, 40-100 m.

Remarks. This species is characterized by the two narrow dark brown bands on the final whorl, i

although color fades rapidly in dead specimens and is not apparent on fossil specimens. Bartsch (1917) II

overlooked the earliest name, which was based on fossil specimens, and proposed four additional names for

this species. The species is best known under the nameEuIima californica. The synonymy above isintroduced II

here. II

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Figure 1.13, Eu_imidae.A.Eu_imaraym_ndiR_vers__9_4_46-53m___Aba__neP_int_OrangeC_unty_Ca_if_r_ta

t (LACM 40-104.53); height 12.2 ram. B. Pseudosabinella bakeri (Bartsch, 1907):82-91 m, Isthmus Cove, Santa Catalina Island, California (LACM 70-15.7); height 5.6 mm. C. Balcis micans

(Carpenter, 1864): 66-86 N, Off Platt Point, Santa Cruz Island, California (LACM 41-83.22):

height 12.8mm. D.Balcis oMroydae(Bartsch, 1917): on sea cucumbers,off SanPedro,Los Angeles

I C131ounty, m, offCalifornia Santa Catalina (LACM 151631): height 8.6 mm. E. Polygireulima rutila (Carpenter, 1864): Island, California (LACM 54-76.9); height 7.0 mm.

I Genus Pseudosabinella McLean, 1995

Pseudosabinella McLean, 1995. Type species (OD): Sabinella bakeri Bartsch, 1917. California.

I Diagnosis. See species description below.

Biology. Echinoderm host unknown (Waren, 1992:189).

I Remarks. McLean (1995:39) proposed a genus for this monotypic species because the shell characters

are well marked and unique in the family. Earlier, War6n (1992:189) noted forSabinella baked: "This species

i probably is an eulimid despite having a rather fragile and irregular shell. I have examined a specimen with dried soft parts, and it has a ptenoglossate radula, similar to species of Eulima.'"

I Pseudosabinella bakeri (Bartseh, 1917)

Figure 1.13B

I Sabinella bakeri Bartsch, 1917:334, pl. 43, fig. 5.--Waren, 1992:189.

MelaneUa (SabineUa) bakeri: Oldroyd, 1927:376.--Abbott, 1974:126. fig. 1382.

I Pseudosabinella bakeri: McLean, 1995:39. Alaba catalinensis Bartsch, 1920:572.

i Alaba serrana Smith and Gordon, 1948:225, pl. 4, fig. 1,2.
!

Material Examined. California: Santa Maria Basin, Phase I, sta. 2, 200 m (1); sta. 20, 396 m (1); I

Phase II, sta. R-4, 92 m (1); sta. PJ-1,145 m (1); sta. PJ-10, 147 m (1); sta. PJ-22, 143 m (2). Other material:

19 lots in the LACM collection. II

Description. Shell thin, light brown; whorls rounded, suture deep, f'mal lip inflated and produced •

anteriorly, umbilical chink present, lip scars slightly raised, variable in position. Ptenoglossate radula present.

Length 3-6 ram. l

II Type Locality and Type Specimens. San Diego, California. Holotype: USNM 215786. Alaba

catalinensis: Santa Catalina Island, California, 40 fathoms; holotype: USNM 213369. Ataba serrana: Carmel

Bay, Monterey County, California, 25 fathoms; holotype: CAS 065531. I

I

Distribution. Carmel, Monterey County, California (36°N), to Isla Cedms, Baja California (28°N).

Habitat. Rocky bottoms, 10-150 m. Specimens have been recovered from gravel taken by divers in m

the shallow sublittoral in southern California. !!

Remarks. This species is characterized by its light brown shell with inflated whorls and greatly

expanded outer lip. There are significant size differences, which may be an indication of sexual dimorphism. /

It has been confused with the genusAlaba (family Litiopidae), from which it differs in shell characters by its il

glossy shell, growth scars, and expanded lip. The synonymy was first indicated by McLean (1995).

B

Genus Balcis Leach, 1847 U

Balcis Leach in Gray, 1847. Type species: Balcis montagui Leach in Gray, 1847 [= Strombiformis albus Da j

Costa, 1779]. Western and southern Europe. l

Diagnosis. Shell tall, white, straight-sided, aperture of moderate length.

Biology. The type species is parasitic on holothurians (Warrn, 1984:32). I

Remarks. On shell characters, the genus differs from Melanella in its straight-sided, rather than

inflated whorls. I

Balcis micans (Carpenter, 1864) ll

Figure 1.13C I

Eulima micans Carpenter, 1864:659. J

Melanella micans: Oldroyd, 1927:372. U

Balcis micans: Palmer, 1958:195. II

Melanella (Balcis) micans: Abbott, 1974:125, fig. 1338. I

Material Examined. Not represented in the voucher collection. Other material: 32 lots in LACM

collection. I

Description. Shell moderately large for family; diameter exceeding 3 ram; glossy white, up to 15

whorls; whorls nearly straight-sided; growth scars few, irregular in position. Length 10-13 mm. I

Type Locality and Type Specimens. San Pedro, Los Angeles County, California. Holotype: USNM m

14850.

Distribution. Kodiak Island, Alaska (Bartsch, 1917:308) (57°N), to Punta Abreojos, Baja California

Sur (Dall, 1921:118) (27°N). The LACM collection contains material ranging from Vancouver Island to II

southern California.

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I Habitat. The echinoderm host is unknown, although the species is presumed to be parasitic on

holothurians. Depth records are 30-100 m.

I species inRemsizarks e. Shell . This diame is one ter eofxcetheedslargest 3 mm ofandnomartheas y reach tern 4Pa.5 cmmific. eulimids, differing from the following

i Balcis oldroydae (Bartsch, 1917)

Figure 1.13D

Melanella (MeIanella) oldroydi Bartsch, 1917:309, pl. 36, figs. 5-7.----Oldroyd, 1927:374, pl. 41, figs. 5-

7.--Abbott, 1974:126, fig. 1378.

I Melanella (Melanella) micans borealis Bartsch, 1917:305, 35, 7. pl. fig.

Material Examined. California: Santa Maria Basin, Phase I, sta. 21, 49 m (1). Other material: 63

i lots in the LACM collection.

Description. Shell of moderate size for family, length to 10 mm, diameter not exceeding 2.6 mm;

glossy white, up to 14whorls; whorls nearly straight-sided; growth scars few, irregular in position. Length 8-

I 10mm.

Type Locality and Type Specimens. San Pedro, Los Angeles County, California. Holotype: USNM

i 109641. DMelane istribuUtiaonmicans . Kachemak borealisBa: y,ComAlaska ax, V[LACM ancouver80-74.8 Island,] (Bri 59°N) tishmCoIslalumbia; Cedros, holotype: Baja California USNM 150954. [LACM

71-158] (28°N).

I Habitat. Soft bottoms, 20-90 m. A single LACM lot from San Pedro, Los Angeles County, California, is annotated: "off sea cucumbers" (figure 1.13D).

Remarks.This is a smaller species thanBalcis micans, but is much larger than the following species.

I Bartsch's Melanella micans borealis is here added to the synonymy ofBalcis oldroydae. The name is emended to oIdroydae because Bartsch indicated that it was named after Mrs. Oldroyd.

I Genus Polygireulima Sacco, 1892

Polygireulima Sacco, 1892. Type species (OD): Eulima spina Grateloup, 1838. Miocene, France.

I Diagnosis. Shell tall, straight-sided, smooth, except for growth scars.

Biology. The host echinoderm of the European species is unknown (War6n, 1984:66), but the eastern

I Pacific species assigned to this genus is known to be parasitic on sea stars of many genera.

Remarks. The following species has traditionally been placed in Balcis, the species of which are

aim parasitic on holothurians. War6n (1984) precludes the placement of species from different classes of echinoderms

L" in the same genus. Because the following species is known to be associated with sea stars rather than

holothurians, it is here assigned to the genus Polygireulima, the type species of which is a close match for

shell characters.

I

I

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Polygireulima rutila (Carpenter, 1864) I

Figure 1.13E

II

Eulima rutila Carpenter, 1864:659. 1

Melanella (Melanella) rutila: Bartsch, 1917:306, pl. 35, fig. 2, 3, 6. i

MelaneUa rutila: Oldroyd, 1927:373, pl. 46, figs. 2,3,6. |

Balcis rutiIa: Palmer, 1958:195.

MelaneIla (Balcis) rutila: Abbott, 1974:125, fig. 1340. i

Material Examined. California: Santa Maria Basin, Phase I, sta. 23, 195 m (2); sta. 84, 394 m (1);

sta. 101,357 m (1); Phase IT,sta. PJ-12, 145 m (1); sta. PJ-14, 134 m (1); sta. PJ-15, 155 m (1); sta. PJ-17, I

126 m (2). Other material: 140 lots in the LACM collection. i

Description. Small, slender, straight; up to 13 whorls, fiat-sided, smooth; growth scars few; aperture

moderately elongate. Length 6-8 mm. i

i

Type Locality and Type Specimens. Monterey, California. Holotype: USNM 14928.

Distribution. Monterey Peninsula, California (37°N) to Bahia Magdalena, Baja California Sur (Dall, i

1921:118) (25 °N). Occurring on hard and soft bottoms, 10-400 m. 1

Habitat. I have observed this species on asteroids of various species in the vicinity of Monterey Bay,

California. It occurs in depths accessible to divers and offshore to 200 m or more. 1

U

Remarks. Although the shell characters of this species resemble those ofBalcis oldroydae, it differs

in its smaller size and its more slender profile. The species is here first assigned to Polygireulima.

I

Genus Vitreolina Monterosato, 1884

_treolina Monterosato, 1884. Type species (SD, Bucquoy, Dautzenberg and Dollfus, 1883):Eulima incurva i

Bucquoy, Dautzenberg and Dollfus, t 883. Europe. l

Diagnosis. Shell slightly to strongly curved, oval in cross section, transparent; suture making marked 1

dip at position of growth scar on shell. II

Biology. The echinoderm host of the type species in unknown. European species for which hosts are

known occur on ophiuroids, echinoids, and holothurians, i

Ul Remarks. Vitreolina is an exception among eulimid genera in having host echioderms of different

classes. The other possibility admitted by War6n (1984) is that the genus is being used too broadly. However, i

the genus is well defined by its curved profile. •

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I V'm'eolina columbiana (Bartsch, 1917)

I Figure 1.14A

Melanella (BaIcis) columbiana Bartsch, 1917:324, pl. 41, fig. 5.--Abbott, 1974:126 [listed only].

i Melan Melanella ella columbiana (BaIcis) grippi : Oldroyd, Bartsch, 1927:369, 1917:326, pl. pl.47, 42,fig. fig.5.5.

Balcis (Vitreolina) titubans Berry, 1956:154, fig. 7, 8.

I Material Examined. Santa Maria Basin, Phase I, sta. 4, 393 m (1). Other material: 53 lots in the

LACM collection.

I Description. Relatively large, transparent, extremely flexed; whorls 15, body whorl slightly inflated; aperture long. Length 6-9.5 mm.

Type Locality and Type Specimens. Departure Bay, British Columbia. Holotype: USNM 207771.

I Melanalla grippi: Newport, Orange County, California; holotype: USNM 203665. Balcis titubans: Anacapa

Island, California; holotype: SBMNH 34554.

Distribution. Attu, Aleutian Islands, Alaska [I.,ACM 86-312} (53°N), to Isla Cedros, Baja California

I ILACM 71-156] (28°N).

Habitat. One LACM record (LACM 151103) indicates association with a large holothurian Cucumaria

I sp. distMost ribution, recowhere rds in the it mayLACMoccurcolle in the ctionintertidal indicatezone. a depth of 50 m or less. Common inthe northern part of the

Remarks. This species has the most extreme curvature of the northeastern Pacific species, differing

I als examination o from other of type species material. in its relatively broad f'mal whorl. The synonymy given above is a based on my

I

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! Figure 1.14. Eulimidae. A. Vitreolina columbiana (Bartseh, 1917): 27 m, off San Pedro, Los Angeles County,

California (LACM 65-35.22); height 6.8 ram. B. _qtreolinamacra: (Bartseh, 1917): 82 m, off Ship

I Rock, Santa Catalina Island, California (LACM 65-6.22); height 4.9 mm. C. Vitreolina yod (Carpenter, 1857):366-375 m, off SE sideSan Nicolas Island, California 0.,ACM 77-132.8); height

3.2 ram.D.Haliella abyssicola Bartsch, 1917:392 m, offPointSal,Santa BarbaraCounty,California

I (SBMNH 142803, Santa Maria Basin, sta. 40); height 10.2 ram.
!

Vitreolina macra (Bartsch, 1917) I

Figure 1.14B

In

Melanella (Balcis) macra Bartsch, 1917:326, pl. 41, fig. 6.--Abbott, 1974:126 [listed only]. II

Melanella macra: Oldroyd, 1927:370, pl. 47, fig. 6. Ill

Melanella (Balcis) catalinensis Bartsch, 1917:329, pl. 40, fig. 7. l

Melanella (Balcis)prefaIcata Bartsch, 1917:327, pl. 42, fig. 4 (Pleistocene).

BaIcis (Vitreolina) obstipa Berry, 1954:8, pl. 1, fig. 5,6 (Pleistocene). U

Material Examined. Santa Mafia Basin, Phase I, sta. 1, 98 m (2). Other material: 44 lots in the

LACM collection. I

Description. Shell of medium size, transparent, moderately flexed; whorls 13, nearly flat-sided, aperture

relatively long. Length 6-7.5 mm. n

Type Localities and Type Specimens. Departure Bay, British Columbia. Holotype: USNM 207772. I

Melanella catalinensis: Catalina Channel, California; holotype: USNM 173802. MeIanella prefalcata: San

Pedro, Los Angeles County, California, Pleistocene; holotype: USNM 215774. Balcis obstipa: San Pedro, I!

Los Angeles County, California, Pleistocene; holotype: SBMNH 34546. i

Distribution. Kachemak Bay, Alaska [LACM 80-76.36] (59°), to Islas Coronados, Baja California

[LACM 79-106] (32°N). 1_

Habitat. Hard and soft bottoms, 20-100 m; host unknown. II

Remarks. This species is readily recognized by its longer aperture, more slender and less flexed

pmf'de than that of _qtreolina columbiana. The above synonymy is based on my examination of type material. I

_treolina yod (Carpenter, 1857) 1

Figure 1.14C

Leiostraca yod Carpenter, 1857:441.--Brann, 1966, pl. 48, fig. 55b.--Keen, 1968:407, fig. 35. I

U

Melanella (Balcis) yod: Bartsch, 1917:330, pl. 40, fig. 9.

Melanella (Balcis) taravali Bartsch, 1917:328, pl. 42, fig. 2. B

U

Material Examined. Santa Maria Basin, Phase I, sta. 4, 393 m (1); sta. 2, 200 m (1). Other material:

27 lots in the LACM collection. I

Description. Small, transparent, substantially flexed; whorls 8, slightly inflated; aperture moderately 1

long. Length 2-3 mm.

Type Locality and Type Specimens. Mazatlfin, Sinaloa. Syntype: NHML Carpenter tablet 2027. 1

MeIaneIla taravalli: Punta Abreojos, Baja California Sur; holotype: USNM 215779. I

Distribution. Santa Maria Basin [voucher specimen] (35°N) ] to Mazatlfin, Sinaloa [type locality]

(23°N). Although this species has a Panamic type locality and is common in the Gulf of California, there are II

13 tots in the LACM collection from southern California. U

Habitat. Soft bottoms, 20-400 m, host unknown. n

Remarks. The distribution is here extended north from the Gulf of California; Keen (1971:446) I

indicated a distribution from Guaymas, Sonora, to Mazatlhn, Sinaloa. This species is readily distinguished

from other Vitriolina by its small size. The synonym Melanella taravali Bartsch is based on an immature •

specimen. i

76 !
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I Genus Haliella Monterosato, 1878

I Haliella Monterosato, 1878. Type species (M): Eulima stenostoma Jeffreys, 1858. North Atlantic.

Diagnosis. Shell tall and slender, with rounded whorls, aperture elongate, columella indented.

I althoughBWargn iology. Species occur in relatively deep water at depths below 400 m. Echinoderm host unknown, (1984:47) suggests that it may prove to be an ophiuroid. War6n has studied the anatomy of

the type species and notes that it is a protandric hermaphrodite.

I Remarks. War6n (1984:47) indicated that there may be about six species worldwide.

I Haliella abyssicola Figure Bartseh, 1917 1.14D

I Haliella abyssicola Bartsch, 1917:336, pl. 43, fig. 8.--Abbott, 1974:128, fig. 1406.

Material Examined. California: Santa Maria Basin, Phase I, sta. 40, 392 m (1). Other material: two

I specimens at CAS [ex Stanford 51315].

Description. Shell tall, moderately large, opaque white; whorls 10, slightly rounded; growth scars

irregular in position; columella with indentation; aperture long. Length 11 ram.

I Type Locality and Type Specimens. Off San Diego, 822 fathoms. Holotype: USNM 251266.

Distribution. Off Columbia River, Oregon [CAS 102991 ] (46°N), to Punta San Pablo, Baja California

I Sur [USNMHabita266887] (27°N). t. Soft bottoms, 400-1500 m; echinoderm host unknown.

Remarks. This species can be confused with no other. Its opaque shell with inflated whorls is

I characteristic.

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Superfamily Muricoidea m

Family Muricidae Rafinesque, 1815 I

mini

Diagnosis, Shell with moderately high spire and short to long anterior canal. Sculpture both axial and

spiral, often producing elaborate spines on regularly spaced varices. Operculum oval, with apical or lateral m

nucleus. Radula rachiglossate; rachidian multicuspid; lateral unicuspid.

Biology. Species are carnivorous, capable of boring into barnacle and bivalve shells. Egg capsules

are attached to rocks or other shells, m

Remarks. The most recent comprehensive treatment of worldwide muricid shells was that Radwin

and D'Attilio (1976), although many groups were omitted. Muricid classification is in a state of flux, as m

recently discussed in two papers by Kool (1993). m

Two subfamilies are utilized for the offshore species included here: Ocenebrinae and Trophoninae.

m

Key to Species of Muricidae m

l

1A. Canal of mature shell sealed ............................................................................................................ 2 U

1B. Canal of mature shell open .............................................................................................................. 4

l

2A. Whorls rounded ..................................................................................................... Ocinebrina lurida

2B. Whorls shouldered ........................................................................................................................... 3 N

3A. Shoulder spinose .......................................................................................... Ocinebrina barbarensis m

3B. Shoulder aspinose ............................................................................................ Ocenotrophon painei m

m

4A. Sculpture axial, or axial with faint spiral striae ............................................................................... 5 mI

4B. Sculpture axial, with well-developed spiral cords .......................................................................... 15

5A. Shell length > 40 mm .............................................................................. Austrotrophon catalinensis U

5B. Shell length < 30 mm ....................................................................................................................... 6

l

6A. Shoulder aspinose ............................................................................................................................ 7

6B. Shoulder spinose ............................................................................................................................ 10 I

7A. Axial lamellae under 20 on final whorl ............................................................................................ 8

n

7B. Axial lamellae over 20 on final whorl .............................................................................................. 9 mm

8A. Canal as long as aperture length ................................................................ Boreotrophon eucymatus m

8B. Canal shorter than aperture length ............................................................... Boreotrophon apolyonis

I

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I 9A. Canal as long as aperture length ....................................................................... Boreotrophon kabati

9B. Canal shorter than aperture length ................................................................ Boreotrophon tolomius

I

10A. Spines laterally projecting .............................................................................................................. 11

I 10B. Spines upturned ............................................................................................................................. 13

11A. Profile tabulate, shoulder area narrow ......................................................... Boreotrophon raymondi

I 11B. Profile not tabulate, shoulder area broad ....................................................................................... 12

I 12A. Profile broad, length about 1.5 times breadth ......................................... Boreotrophon triangulatus 12B. Profiles narrow, length about 1.9 times breadth ....................................... Boreotrophon avalonensis

I 13A. Prof'lle broad, length about twice breadth .............................................. Boreotrophon multicostams

13B. Profile narrow, length about 3 times breadth ................................................................................. 14

! 14A. Spines long, lamellae fading across base ........................................................ Boreotrophon bentleyi

i 14B. Spines short, lamellae strong across base ........................................................... Boreotrophon keepi

15A. Axial sculpture dominant in early teleoconch ................................................................................ 16

I 15B. Spiral sculpture of two lamellar cords in early teleoconch ............................................................ 18

I 16A. Shoulder with upturned varices ......................................................................... Boreotrophon stuarti 16B. Shoulder lacking upturned varices ................................................................................................. 17

I 17A. Whorl profile angulate ............................................................................... Boreotrophon pedroanus

17B. Whorl profile rounded ..................................................................................... Boreotrophon hazardi

I 18A. Whorls rounded ................................................................................................ Scabrotrophon lasius

i 18B. Whorls shouldered ......................................................................................................................... 19

19A. Shoulder lacking strong spines ....................................................................................................... 20

I 19B. Shoulder with spines 21

strong

I 20A. Shoulder area with axial and spiral sculpture .............................................. Scabrotrophon maltzani 20B. Shoulder area with axial sculpture only .................................................... Scabrotrophon cerritensis

I 21A. Spines laterally projecting .............................................................................. Scabrotrophon grovesi

2lB. Spines bent toward shell axis ........................................................................... Scabrotrophon clarki

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Subfamily Ocenebrinae Cossmann, 1903 I

Diagnosis. Shell small to medium-sized, sculpture usually of scaly spiral cords, outer lip usually with •

denticles; operculum with lateral nucleus.

Remarks. Subfamilial classification of the Ocenebrinae was discussed in both papers by Kool (1993).

II

Genus Ocinebrina Jousseaume, 1880 •

Ocinebrina Jousseaume, 1880. Type species (OD): Murex corallinus Scacchi, 1836; = Murex aciculatus

Lamarck, 1822. North Atlantic and Mediterranean. U

Diagnosis. Shell small to medium in size; canal relatively short, sealed at maturity; mature lip thickened

and projecting, with labial denticles; sculpture of f'melyimbricate spiral cords overriding axial ribs; protoconch !

paucispiral; operculum with lateral nucleus.

Biology. The paucispiral protoconch is indicative of direct development. Life histories of several m

species from Puget Sound were studied by Spight et aI. (1974).

Remarks. The northeastern Pacific species formerly treated in Ocenebra Gray, 1847 [type species

Murex erinaceus Linnaeus, 1758], are here placed in Ocinebrina on the recommendation ofE. Vokes (personal II

communication). In the type species ofOcenebra, a thick lip with labial denticles and a sealed canal is formed II

repeatedly during resting stages of shell growth, whereas in Ocinebrina the siphonal canal remains open until

final maturity is reached and the lip expands and produces labial denticles. Radwin and D'Attilio ( 1976:125, In

pl. 21, fig. 8) treated the European type species of Ocinebrina, which better resembles the eastern Pacific |

species than does the European type species of Ocenebra (Radwin and D'Attilio, 1976:119, pl. 18, figs. 4, 5).

Oeinebrina iurida (Middendorff, 1848) I

Figure 1.15A N

Tritonium luridum Middendorff, 1848:244.--Middendorff, 1848:479, pl. 4, figs. 4,5.

Tritonalia lurida: Oldroyd, 1927:314. m

!1 Urosalpinx lurida: Abbott, 1974:179, fig. 1886.

Ocenebra lurida: Radwin and D'Attilio, 1976:123, pl. 20, figs. 4,5.--McLean, 1978:43, fig. 22.7. mi

Vitularia aspera Baird, 1863:66 [unfigured]. U

Ocinebra lurida var. munda Carpenter, 1864:663.

Ocenebra lurida var. munda: Dall, in Williamson, 1892:215, pl. 20, fig. 3.--Palmer, 1958:204. '1n

Material Examined. California: Santa Maria Basin, Phase I, sta. 13,197 m (1 juvenile); sta. 27, 611

m (1). Other material: 215 lots in the LACM collection. II

Description. Shell small, whorls rounded, suture moderately impressed, profile varying from slender U

to robust. Sculpture of low axial ribs that are overriden by spiral cords; axial ribs reduced anteriorly.

Periostracum thick; shell color dark yellow to brownish orange, some specimens with darker brown bands or

uniformly dark brown. Length 13-28 ram. U

Type Locality and Type Specimens. Sitka, Alaska. Type material apparently lost; not in Zoological

Institute, St. Petersburg. _tularia aspera: Esquimalt Harbor, Vancouver Island, British Columbia; type material

in the Natural History Museum, London. Ocenebra Iurida var. munda: type material lost (Palmer, 1958:205);

Palmer considered the name to have been validated by Dall in Williamson, and designated a lectotype from

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I Figure 1.15. Muricidae.A. Ocinebrinalurida (Middendorff,1848):12-38m,Carmel, MontereyCounty,California

(LACM 60-24.66); height t4.1 mm. B. Ocinebrina barbarensis (Gabb, 1865): 86 m, S of Long

I Point, catalinensis Santa(Oldroyd, Catalina Island, 1927): 46California m, off San(LACM Pedro,41-179.8); Los Angelesheight County, 20.1Califor mm. nC.iaAustrotrophon (CAS 063306,

holotype); height 100 mm.

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Monterey, California (USNM 46708), the locality taken from the specimen label. However, Carpenter's

description is hardly less detailed than that of others of Carpenter and the name is here considered validated by

I Carpenter.

Distribution. Kenai Peninsula, Alaska (59°N, 151°W), to Puerto Santo Tormis, Baja California

i (31.5°N).Habitat. Rocky bottoms, intertidal zone to 200 m.

Remarks. Ocinebrina lurida varies in overall length and in breadth of the shell. Shells from deeper

I water, as in the offshore form illustrated here, axe generally more slender. Ocinebrina sclera (Dall, 1919), which was figured by Radwin and D'Attilio (1976, pl. 20, fig. 5), is a separate species of more northern

occurrence; it is not a synonym of O. lurida, under which it was placed by Radwin and D'Attilio.

I Ocinebrina barbarensis (Gabb, 1865)

I Figure 1.15B

Murex (Muricidea ) barbarensis Gabb, 1865:183.

I Dall, 1921:108, pl. 6, fig. 5, pl. fig.

Tritonalia barbarensis: 15, 11.

Ocenebra barbarensis: Abbott, 1974:183, fig. 1915.--McLean, 1978:42, fig. 22.2.

I Ocenebra crispatissima Berry, 1953:414, pl. 28, fig. 6.

Material Examined. Santa Mafia Basin, Phase I, sta. 1, 98 m (1 immature). Other material: 52 lots

i in the LACM collection.

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Description. Shell slender, with canal longer than aperttu'e. Shoulder strongly angulate, usually with I

upturned spines, but aspinose specimens known. Single upturned spine of terminal lip larger and more outwardly

projecting than earlier formed spines. Length 15-20 mm. •

Type Locality and Type Specimens. Santa Catalina Island, California, 40 fathoms. Lectotype (here

designated) UCBP 10611 [syntypes were reported by Coan and Bogan, I988:277].

Distribution. Monterey Bay, California (37°N), to Isla Cedros, Baja California (28°N). Ocenebra B

crispatissima: Isthmus Cove, Santa Catalina Island, California, 33 fathoms; holotype: CAS 064468. I

Habitat. Rocky bottoms, 10-100 m. BI

Remarks. The species concept is based on figures of Dall (1921 ), not of type material, which has not I

been illuslxated. Ocenebra crispatissima Berry was based on an aspinose specimen. Although both Ocinebrina

barbarensis and O. crispatissima were treated by Radwin and D'Attillio (1976:120) as synonyms of Ocinebrina I

foveolata (Hinds, 1844), Ocinebrina barbarensis is a well-differentiated species, having the longest canal of I

the northeastern Pacific species.

Genus Austrotrophon Dall, 1902 I

Austrotrophon Dall, 1902. Type species (SD, Grant and Gale, 1931): Trophon cerrosensis Dall, 1891. Tropical

eastern Pacific. I

Diagnosis. Shell moderately large; axial sculpture of broad winglike varices producing long spines at m

shoulder; color light brown, surface not chalky. Operculum with lateral nucleus. _l

Remarks. This genus is known from southern California and the Panamic Province. The type species,

figured by Keen (1971:537, fig. 1044), has prominent spiral cords on the body whorl.

Although the shell sculpture of broad varices resembles that expected in the subfamily Trophoninae, l

the nucleus of the operculum is lateral rather than ternYmal.Austrotropon has been placed indifferent subfamilies

by previous authors. Radwin and D'Attillio (1976:176) did not treat it in their book on Muricidae because •

they considered that the radula placed it in the family Thaididae (now the muricid subfamily Rapaninae). The 1

operculum is similar to that ofForreria belcheri (Hinds. 1844), which was placed by Kool (1993: 227) in the

Ocenebrinae on the basis of anatomical characters. I therefore groupAustrotrophon in the Ocenebrinae. I

Austrotrophon eatalinensis (Oidroyd, 1927)

Figure 1.15C I

Trophon (Austrotrophon) catalinensis Oldroyd, 1927:327, pl. 34, f. 1-5. l_

Forreria (Austrotrophon) catalinensis: Grant and Gale, 1931:726. 1

Trophon (Austrotrophon) cerrosensis catalinensis: Abbott, 1974:191, pl. 9, fig. 2008.

I

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material U

examined: 43 LACM lots.

Description. Shell light brown, often with darker brownaxial markings andfaintspiralbands; sculpture I

of 7 sharpvarices per whorl, upturned andprojecting diagonally, forming spines open in front; aperture white, g

glossy; parietal wall broad, inner lip raised; siphonal canal long, open, forming a fasciole, nearly straight or

twisted to left. Length 80-100 mm. l

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I Type Locality and Type Specimens. Off San Pedro, Los Angeles County, California, 25 fathoms.

Holotype: CAS 063306.

I Baja California DistributSur ion. Off Point Conception, Santa Barbara County, California (34.5°N) to Punta Abreojos, (26°N).

Habitat. 40-160 m on soft bottoms.

I Remarks. In her description this species, Oldroyd (1927:327) species "has

of noted that this been

called Trophon Iriangulams Carpenter," evidently referring to illustrations of Dall (I 891, pl. 5, figs. l, 6).

However, in my opinion, Dali's illustrations represents a variant of the type species Trophon cerrosensis,

I which Dall described in the Abbott 1974:191 catalinensis

sanle paper. ( ) placedAustrotrophon as a

subspecies

of underA, cerrosensis, but that species, the type species of Austrotrophon, differs in having spiral sculpture

and more laterally directed shoulder spines. Austrotrophon pinnatus (DalI, 1902), which has a more southern

I distribution in Baja California, is a similar species. It differs in its higher spire, more twisted fasciole and

more extensive development of a broad flaring varix below the aperture.

I Subfamily Trophoninae Cossmann, 1903

I Diagnosis. Shell fusiform, usually white; sculpture of axial larnellae not forming varices of alternating

strength, with spiral sculpture usually weak or lacking. Opereulum with apical nucleus.

i Remarks. Genera occur in cool water, usually at high latitudes, or deeply submergent at tropical latitudes. The genera are poorly known and in need of review. Generic characters emphasized here are provided

by the early teleoconch sculpture.

I genus Trophon Kool (1993:55) Montfort, noted 1910,that is atypical Trophonof geversianaus species usually(Pallas, included1774), in Trophoninae, the type species and suggested of the nominate that the

group may not be monophyletic.

I Genus Boreotrophon Fischer, 1884

i Boreotrophon Fischer, 1884. Type species (M): Murex clathratus Linnaeus, 1767. Arctic Ocean.

Diagnosis. Shell fusiform, with long anterior canal; protoconch paucispiral, of 1.5 rounded, wellseparated whorls; axial sculpture of strong lamellae, early teleoconch sculpture axial; spiral sculpture if

I not crossing or raised above axial lamellae.

present

Remarks. Many species of Boreotrophon lack spiral sculpture altogether. Spiral sculpture in the

I species that have it is of lesser strength than the axial sculpture and does not override the axial lamellae. The earliest teleoconch sculpture is always dominated by axial elements.

This is a genus of the Northern Hemisphere, with numerous species in the Arctic Ocean, the North

I Atlantic Egorov and the North Pacific. (1992, 1993) reviewed the species of the northwestern Pacific. Radwin and D'Attilio (1976:179)

treated the type species, but did not treat other species of the genus. All species known from southern California

I are treated here.

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Figure 1.16. Muricidae. A. Boreotrophon triangulatus (Carpenter, 1864): 55 m, off White's Landing, Santa w

Catalina Island, California (LACM 1028, holotype of Trophon albospinosus), height 21 mm. B.

Boreotrophon mutticostams (Eschscholtz, 1829): 29 m, off Santa Catalina Island, California (USNM I

109165, holotype of Boreotrophon peregrinus); height 23 mm. C. Boreotrophon bentleyi Dall,

1908:37 m, off San Diego Harbor, California (USNM 110648, holotype); height 19.5 mm. D.

Boreotrophon avalonensis Dalt, 1902:146 m, off Avalon, Santa Catalina Island, California (USNM •

I09109); height 16.5 mm. E. Boreotrophon avalonensis DalI, 1902:40 m, Cortez Bank, California |

(CAS 102992); height 19 ram. F. Boreotrophon eucymatus Dall, 1902:227 m, off San Diego,

California (USNM 109089, holotype); height 21 mm [not 27 mm]. .,

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Boreotrophon triangulatus (Carpenter, 1864) i

ill

Figure 1.16A

Trophon triangulatus Carpenter, 1864:653.--Da11, 1891:181, pl. 5, fig. 3. i

Trophon (Austrotrophon) triangulatus: Oldroyd, 1927:326, pl. 34, figs. 6, 7.

Trophonopsis triangulatus: Palmer, 1958:207. I

IJ

Trophon (Boreotrophon) albospinosus Willett, 1931:66, pl. 4, fig. 3.

Material Examined. No material is present in the voucher collection. Other material: 26 LACM lots. 1

IB

Description. Shell white, relatively small, profile broad; whorls markedly angulate, anterior canal

short; axial lamellae about 6-8 per whorl, lamellae spinose, laterally projecting at periphery; spiral sculpture 1

lacking. Length 15-21 mm. 1

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Type _ and Type Spee_ens. Santo C_a Island, C_o_a_ _m_ (sel_ by P_er,

1958:_8): UC_ 12572. Trophon albospinosus: _te's _ng, San_ Ca_lina Island, 30 fathoms; 1

holo_e: LACM 1_8. I

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I (33°N). Distribution. Point Conception, Santa Barbara County (34.5°N), to Santa Catalina Island, California

Habitat. Rocky bottoms, 20-90 m.

I Remarks. This is broad, with the whorl and base It resembles specie_ body having a

triangular profile.

Boreotrophon avalonensis, but is broader, with a shorter canal. It also resembles the more northern species B.

multicostatus (Eschscholtz, 1829), which reaches larger sizes, has a more cylindrical profile, and has spines

I that are upturned.

I Boreotrophon multicostatus (Eschscholtz, 1829) Figure 1.16B

I Murex multicostatus Eschscholtz, 1829 829-31 2:1 4. [1 ],part 1, pl. 4, fig.

Trophon (Neptunea) multicostatus: Oldroyd, 1927:331, pl. 33, fig. 11.

I Trophon (Boreotrophon) multicostatus: Grant and Gale, 1931:722. Boreotrophon peregrinus Dall, 1902:543.--Da11, 1921:110, pl. 8, fig. 5.

I 78 LACMMatlots, eriaincluding l Examine13d. lots Notfrom represented south ofinPoint the Santa Conception. Maria voucher collection. Other material examined:

Description. Shell white, aperture usually brown within; small to medium size, profile broad; anterior

I canal moderately long, less than length of aperture; axial lameUae 6-10 per whorl, spinose at shoulder, spiral sculpture of fine striae. Length 20-30 mm.

Type Locality and Type Specimens. Sitka, Alaska; type material not located. Boreotrophon

I Catalina Harbor, Santa Catalina Island, California, 16 fathoms; USNM 109165.

peregrinus: holotype:

Distribution. Kenai Peninsula, Alaska (59°N, 152°W), to Laguna Beach, Orange County, California

i (33.5ON)"Habitat. Rocky and gravel bottoms, intertidal to 100 m. Specimens from southern California are

submergent, at depths greater than 40 m.

I Boreo Remarks. Dali's taxon Boreotrophon peregrinus is here first placed in the synonymy of this species. trophon multicostatus has been considered to be a more northern species, but there are specimens in the

LACM collection from throughout the distribution. Northern specimens tend to reach larger sizes and may

I have more numerous axial lamellae than those occurring in southern California. The prof'de is broader and more cylindrical than that ofBoreotrophon triangulatus and the shoulder

spines are more upturned.

I

Boreotrophon bentleyi Dall, 1908

I Figure 1.16C

Boreotrophon bentleyi Dall, 1908:249.--Abbott, 1974:190 [listed only]. 1

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I Trophon (Neptunea) bentleyi: Oldroyd, 1927:333.

Material Examined. California: Santa Mafia Basin, Phase I, sta. 17, 654 m (1 immature); Phase II,

I sta. R-2, 161 m (1). Other material: 40 lots in the LACM collection.

Description. Shell white, relatively small, slender; anterior canal long; axial lamellae 9-12 per whorl;

lameltae spinose, upturned and strongly projecting at shoulder, fading across base; spiral sculpture lacking or

I faintly indicated. Length 20-27 ram.
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Type Locality and Type Specimens. San Diego, California, 20 fathoms. Holotype: USNM 110648. I

Distribution. OffGaviota, SantaBarbara County (34°N), to La Jolla, San Diego County, California

(33°N).

till

Habitat. Rocky bottoms, 100-350 m. 1m

Remarks. This species is characterized by its slender profile andprominent upturned spines; the

spines are parallel to the body whorl and not laterally projecting, l

m

Boreotrophon avalonensis Dall, 1902

m

Figure 1.16D,E I

Boreotrophon avalonensis Dall, 1902:546.--Dall, 1921:110, pl. 8, fig. 8.--Abbott, 1974:190, fig. 1980 •

[copy Dall figure]. II

Neptunea callicerata Dall, 1919:338 [holotype unfigured].

Neptunea staphylina Dall, 1919:338 [holotype unfandigured]. !

Material Examined. No material is present in the voucher collection. Other material: 29 lots in the

LACM collection. I Description. Shell white, relatively small; profile moderate, anterior canalrelatively long; axiallamellae

about 7-9, usually spinose and laterally projecting at shoulder, spiral sculpture lacking. Length 15-21 mm. m

Type Locality and Type Specimens. Off Avalon, Santa Catalina Island, California, 80 fathoms. I

Holotype: USNM 109109. Neptunea callicerata: Off Point Loma, San Diego County, California; 120-131

fathoms; holotype: USNM 209914. Neptunea staphylina: Off Santa Barbara Island, California, 302-638 8

fathoms; holotype: USNM 209947. mE

Distribution. Santa Rosa Island, California (34°N), to Islas San Benito, Baja California (28 °N).

Habitat. Rocky bottoms, 80-270 m. II

Remarks. This resembles Boreotrophon triangulatus in its laterally projecting spines, but has a

longer canal, more lamellae, and is smaller. It is broader than B. bentleyi, with the spines lower on the

shoulder, and more laterally projecting. I

II

The synonymy above is introduced here. The holotype of the junior synonymNeptunea callicerata is

similar to that of Boreotrophon avalonensis. The holotype of the junior synonym Neptunea staphylina has I

weak spines. Figure 16E shows aspecimen lacking development of the spines; this specimen is referred toB. I!

avalonensis because the angulate periphery is low on the whorl. u

Boreotrophon eucymatus Dall, 1902 I

Figure 1.16F

I

Boreotrophon (avalonensis var.?) eucymancs Dall, 1902:547.--Da11, 1921:110, pl. 15, fig. 7.

Trophon (Neptunea) eucymatus: Oldroyd, 1927:333, pl. 30, fig. 9. •

Boreotrophon eucymatus: Abbott, 1976, fig. 1984 [not fig. 1989]. II

Material Examined. No material is present in the voucher collection, although the species occurs in •

southern California. Other material: 19 lots in the LACM collection. |

Description. Shell white, relatively small, profile of moderate breadth, anterior canal long; axial

lamellae about 13-19; lamellae aspinose, backfilled; shoulder angulate; spiral sculpture lacking. Length 15- •

26ram.

86 I
!

I Type Locality and Type Specimens. Off San Diego, California, 124 fathoms. Lectotype (here

designated): USNM 109087.

I Distribution. Off SantaBarbara Island (33.5°N) to N of San Clemente Island, California (33°N). Habitat. Rocky bottoms, 100-500 m.

Remarks. This species differs from other species in southern California in lacking spines at the

I shoulder and in the lamellae low and backfilled. The shoulder is the whorl than that of

having higher on

Boreotrophon avalonensis.

I Boreotrophon apolyonis (I)all, 1919)

I Figures 1.17A,B

Neptunea apolyonis Dall, 1919:337.

I Tropho Trophonnopsis (Neptunea apolyon ) apolyonis: Oldroyd, 1927:338. is: Abbott, I974:191 [listed].

I examined: Materi LACaMl Exam 76-313.1, ined.Tanner Not repr Baesnk, ented Califorinnthe Santa Mafia Basin voucher collection. Other material

ia, 390 m (1); LACM 77-250.16, TannerBank California,

511-530 m (1).

I Description. Shell white, size medium, of moderate width, canal deflected to left, length less than aperture length; axial lamellae retractively flexed at shoulder, 13-15 on final whorl, cresting and faintly

spinose at shoulder; spiral sculpture of fine, regularly spaced, incised striae. Length 14-21 mm.

I Type Locality and Type Specimens. "Vicinity of Santa Barbara Islands" [off San Nicolas Island, California], 216-339 fathoms. Lectotype (here designated): USNM 209303.

Distribution. San Nicolas Island (33°N) to TannerBank, California (33.5°N).

I Habitat. Soft bottoms, 390-620 m.

Remarks. Type material is illustrated here for the first time (Figure 17A) and the first subsequent

I rec greater ords depths of the sthan peciethose s are ofalsthe o reported precedingherespecies (Figu.re 17B). The three known records for this species axefrom

Boreotrophon apolyonis has a shell profile suggestive of the boreal type species, Boreotrophon

I clathratus (Linnaeus, 1767), but differs in its smaller size and presence of spiral sculpture.

i BoreotrophonFigure raymondi (Moody, 1916) 1.17C,D

I Trophon raymondi Moody, 1916:53, pl. 1, figs. la, lb.

Trophon (Boreotrophon) raymondi: Grant and Gale, 1931:724.

I Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material: 17 LACM lots.

Description. Shell white, relatively small, of moderate width; body whorl constricted at base, canal

I long straight, length greater length aperture; area concave.

and of canal than of shoulder Axial lamellae

retractively flexed at shoulder, poorly developed on shoulder, 1I-14 on final whorl, sharply spinose at shoulder,

i spines laterally directed; spiral sculpture of fine striae. Length 14-20 ram.
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Figure 1.17. Muricidae. A. Boreotrophon apotyonis (Dall, 1919): 395-620 m, SantaBarbara Islands, California

(USNM 209303, lectotype); height 21 mm. B. Boreotrophon apotyonis (Datl, 1919): 390 m, Tanner •

Bank, California (LACM 76-313.1); height 14.0 ram. C. Boreotrophon raymondi (Moody, 1916): |

390 m, Tanner Bank, California (LACM 76-313.3); height 14.9 ram. D. Boreotrophon raymondi

(Moody, 1916): Off China Harbor, Santa Cruz Island, California, 234 m (Thelma Crow collection); 1

height 18.8 mm. E. Boreotrophon keepi (Strong and Hertlein, 1937): 55-91 m, off W end San I

Nicolas Island, California (CAS 065961, holotype); height 26.9 mm. I

!

Type _ and Type Sl_ee_men._. Upper Pliocene, Fourth and Broadway, Los Angeles, California.

Holotype: UCMP 11088. •

Distribution. Queen Charlotte Sound, British Columbia [LACM 69-71.11 ] (52°N), to San Clemente II

Island, California [LACM 76-387.8] (32.5°N).

Habitat. Soft bottoms, 80-440 m. Occurring at minimal depths in the northern part of the distribution; !

at depths greater than 300 m in southern California. It

Remarks. This species, which was originally described from the Upper Pliocene of Los Angeles, is tm

here first reported in the living fauna. There is variation in size in the living material, as also reported by III

Moody for fossil specimens, as well as variation in spire height and the number of axial lamellae. Figure 17D

w

shows a higher-spired specimen in which the canal tip is missing due to breakage. I

The long canal and overall profile ofBoreotrophon raymondi resembles that ofB. coronatus (H. and I

A. Adams, 1864) [= Trophon muriciformis Dall, 1877], a much larger, northern species from the Chukchi and

Bering Seas. Immature shells of that species comparable in size to specimens ofB. raymondi differ in being It

more slender and in having one less teleoconch whorl, and also differ in lacking the fine spiral sculpture of B.

|

raymondi.

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I Boreotrophon keepi (Strong and Hertlein, 1937)

I Figure I. 17E

Trophon keepi Strong and Hertlein, 1937:170, pl. 35, fig. 8.

i Trophonopsis keepi: Abbott, 1974:191 [listed]

Material Examined. Not represented in the Santa Maria Basin collection. Material examined: holotype

and LACM 75-319.2, off South Point, Santa Rosa Island, California, 115 m (5).

I Description. Shell white, size medium, of slender profile; canal length equal to aperture length; axial

lamellae 10-12, of moderate strength, slightly spinous at shoulder, extending from shoulder to canal, but

i lower andTymore pe Loccrowded ality andon canal; Type spiral Specimesculpture ns. OffofWmicroscopic end San Nicstriae. olas IHeight sland, Calif 26.9ormm. nia, 30-50 fathoms.

Holotype: CAS 065961.

I Distribution. Off Santa Rosa Island (34°N) to San Nicolas Island, California (33°N).

Habitat. Soft bottoms, 55-90 m.

i Remarks. This species has a moderate number of axial lamellae that extend across the body whorl and crest at the shoulder. It is more slender, with more numerous, but less raised lamellae than those of

Boreotrophon multicostatus. It is larger thanB, bentleyi, from which it differs in having less projecting spines

i atthe shoulder and lametlae that extend across the body whorl. It differs fromB, apolyonis in its more slender profile and lack of broadly marked spiralsculpture. It most resembles the more northemB, clathratus (Linnaeus,

1767), from which it differs in its much more slender profile.

I They areTheimmature five specimens (the largest reported measures here fr13.2 om Santa mm inRoheight), sa Islandbutarethetheidentity only addisitiocertain nal specimens because known. of their

extremely slender profile.

I Boreotrophon kabati McLean, new species

i Figure 1.18A

Material Examined. Not represented in the Santa Maria voucher collection. Other material in addition

I to6.2,holoffotype: LongLPoint, ACM 4Santa 1-204.6, Catalina off Long Point, Santa Catalina Island, California, 488-635 m (1); LACM 53- Island, California, 419 m (1); LACM 75-320.5, off South Point, Santa

Rosa Island, California, 103 m (1); LACM 40-170.6, off Gull Island, S side Santa Cruz Island, California,

i 201-256 Desc m (1). ription. Shell white, relatively small, slender; shoulder rounded, high on whorl. Canallong, of

length equal to that of aperture length. Apex eroded on all specimens. Axial lamellae numerous,low, partially

I backfiUed, on shoulder28-37 and onbodyfinalwhorl, whorl well (28 onmarked holotype), but retractive less elevated ly flexed on canal. at shoulder, Spiralbutsculpture not spinoulacking. s, well developed Length of

holotype 17.0 mm.

I Type Locality and Type Specimens. Off Long Point, Santa Catalina Island, California (33°24.7'N, 118°14.1'W), Velero IV, sta. 1402-41,440-488 m, I specimen. Holotype: LACM 2782.

Distribution. Santa Rosa Island (34°N), to Santa Catalina Island, California (33.5°N).

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Figure 1.18. A. Boretrophon kabati McLean, new species: 439-488 m, off Long Point, Santa Catalina Island, I

California (LACM 2782, holotype); height 17.0 ram. B. Boreotrophon tolomius (Dal], 1919): off

i

San Miguel Island, California CLTSNM222441, holotype); height 19 ram. C. Boreotrophon hazardi l

McLean, new species: 219 m, off Isla Vista, Santa Barbara County, California (LACM 2783, I_

holotype); height 20.0 ram. D. Boreotrophon pedroanus (Arnold, 1903): 270 m, off Gaviota, Santa

Barbara County, California (LACM 65582); height 14.5 ram. E. Boreotrophon stuarti (E. A. Smith,

1880): 112 m, off Santa Barbara, California CLISNM122582, holotype of Boreotrophon smithi); •

height 41 ram. l

Habitat. Soft bottoms, 100-490 m. i

Remarks. This species differs from other eastern Pacific species in its slender prof'fle and numerous

axial lamellae. In its numerous ribs it most closely resemblesBoreotrophon tolomius, but that species differs •

in its broader profile, shorter canal and faint spiral sculpture. It also resembles the more northern species B. I

tripherus Dall, 1902, which has traces of spiral sculpture, and axial ribs that are more strongly retractive and

fewer (approximately 14) that fade on the body whorl below the shoulder. •

The name honors Dr. Alan R. Kabat, formerly of the U.S. National Museum of Natural History.

Boreotrophon tolomius (I)all, 1919) i

Figure 1.18B

1

Neptunea tolomia Dall, 1919: 337. I

Trophon (Neptunea) tolomia: Oldroyd, 1927:339. I

Trophonopsis tolomius: Abbott, 1974:191 [listed]. i

Material Examined. Not represented in the Santa Maria Basin voucher collection. Material examined:

holotype, i
I

I Description. Shell white, size medium, profile broad, canal relatively short, less than half length of

aperture. Axial lamellae numerous, 35 on final whorl, low, backf'llled, extending across shoulder and body

i wwhorl, horl, but forming diminished faint nodes on canal. whereShcrossing oulder weakl axialyriangula bs. Length te; spiral of holotype cords w19.3 eak, abmm. out 6 on shoulder andbody

Type Locality and Type Specimens. Off San Miguel Island, California, 376 fathoms. Holotype:

I USNM 222441. Distribution. Off San Miguel Island, California (known only from holotype).

Habitat. Soft bottoms, 690 m.

I Remarks. This species is unlike others from the northeastern Pacific in its broad profile and large

number of axial lamellae. In its numerous axial lamellae, it is similar toBoreotrophon kabati, but differs in its

i broader profile, shorter canal, and faint spiral sculpture. Dali's original description made no mention of the spiral sculpture, but it is apparent in the illustration here.

Although the depth reported for this species is greater than that of other species included in this work,

I annewexception species Bis. made kabatiin. order to figure the holotype for the first time and provide a basis for comparison to the

I Boreotrophon hazardi McLean, new species

Figure 1.18C

I Material Examined. Not represented in the Santa Maria Basin voucher collection. Known only from

holotype and paratype.

I on final De whorl, scripweakly tion. Shell angulate white,onsizeearlymedium, whorls;profile canalmoderatel length equal y broad, to aperture suture colength. nstricted. Axial Whribs orls 4few,, rounded 10 on

final whorl, number of axial ribs constant from whorl to whorl, ribs projecting, backfilled, extending across

i shoulder and body whorl but obsolete on canal. Spiral cords faint, 4, interspaces of similar width, detected only on final whorl and penultimate whorl. Columellar wall not projecting to form raised inner lip. Height of

holotype 20.0 mm.

I (approxima TytepelyLocality 34°23'N,and120Type °00'W), Spec220 imens. m. Coll OffeNctedaplesbyPoint, RalphIslHaaVista, zard onSanta drgboBaatrbarKaildee Co,unJanua ty, Calif ry o19rni74a.

Holotype: LACM 2783. There was an additional paratype specimen (length 18.8 mm) from the same sample

I retained the canalbiny ththee late holotype Thelmais broken, Crow, butthethecanal whereabouts apparentlyofretains that specimen its original is nolength. w unknown. Although the tip of

Distribution. Off Isla Vista, Santa Barbara County (34.5°N); known only from type locality.

I Habitat. Soft bottoms, 220 m.

Remarks. This species is unlike other northeastern Pacific species in its relatively few, massive,

backfilled axial ribs, faint spiral sculpture, and deeply impressed suture.

I This species is named after Ralph Hazard, skipper of the dragboat Kildee, who saved numerous

shells for the late Thelma Crow, who in turn donated them to the LACM collection.

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Boreotrophon pedroanus (Arnold, 1903) I

Figure 1.18D

Trophon (Boreotrophon pedroana Arnold, 1903:251, pl. 6, fig. 12. •

Trophon (Boreotrophon) orpheus var. pedroana: Grant and Gale, 1931:723. i

Boreotrophon pedroanus : Woodring, 1946:76. I

Trophon (Boreotrophon) stuarti var. praecursor Arnold, 1903:253, pl. 6, fig. 5.

m

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material 1

examined: 21 LACM lots.

Description. Shell light tan with darker aperture, relatively small; spire high, suture deeply impressed; i

whorls shouldered, canal moderately long. Axial lamellae low, 16-18 on fmal whorl, crossed by 4 spiral cords I

of equal strength, one defining the shoulder, one at periphery and 2 below. Early whorls showing the shoulder

cord and the peripheral cord. Height 14-20 mm. i

m

Type Locality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene. Holotype

not located. Trophon stuarti var. praecursor: San Pedro, California, Pleistocene; holotype: USNM 162546. in

Distribution. Santa Rosa Island, California (34°N), to Isla Todos Santos, Baja California (32°N) I

[LACM 41-25.16].

Habitat. 40-270 m, soft bottoms, i

Remarks. This species is here reported living for the first time, having been first described as a

Pleistocene fossil. Grant and Gale (1931) and Woodring (1946) associated the two names. Although most

specimens in the LACM collection were not live-collected, one lot (LACM 150844) has immature specimens 1

that were live-collected.

Boreotrophon pedroanus resembles the more northern species B. orpheus (Gould, 1849), which i

differs in having more numerous spiral cords of lesser strength than that of the axial varices. I

Boreotrophon stuarti (E. A. Smith, 1880) i

Figure 1.18E

Trophon stuarti E. A. Smith, 1880:481, pl. 48, fig. 6. I

i

Trophon (Neptunea) stuarti: Oldroyd, 1927:333, pl. 33, fig. 13.

Trophon (Boreotrophon) stuarti: Arnold, 1903:252, pl. 6, fig. 4. I

Boreotrophon stuarti: Abbott, 1974:189 [not fig. 1973, smithO.

Boreotrophon (stuarti vat.?) smithi Dall, 1902:542. n

Trophon (Neptunea) smithi: Dall, 1921:111, pl. 13, fig. 8.----Oldroyd, 1927:336, pl. 33, fig. 14 [copy Dall i

fig.].

Boreotrophon smithi: Abbott, 1974:189, fig. 1973 [as synonym ofB. stuarti]. 1

i

Material Examined. Not represented in the Santa Maria Basin voucher collection, but occurring

offshore in California. Material examined: 53 LACM lots.

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I Description. Shell white, relatively large; spire high, whorls shouldered, canal moderately long.

Axial lamellae 7-11 per whorl, thin, strongly projecting, crossed by narrow, raised spiral cords, 2 on early

I whorls, lip thin, araised, bout 5 onouterbodylipwhorl with thin (butproje sometimes cting edge. obsolete Lengthon bod 35-60 y whmm. orl), producing coarse cancellations; inner

Type Locality and Type Specimens. Vancouver Island, British Columbia; holotype: Natural History

I Mus USNMeum,12Lo25ndo 82.n. Boreotrophon stuarti var.smithi: off Santa Barbara Island, California, 61 fathoms; holotype

Distribution. PribilofIslands, Bering Sea (58°N); Kenai Peninsula, Alaska, to Newport Beach, Orange

I County, California (33°N).

Habitat. Rocky bottoms, 0-100 m, occurring in the intertidal and offshore in Alaska, but deeply

submergent in California.

I Remarks. Specimens incollections fromCalifornia aresmaller thanthose from more northernlocalities.

The variety Boreotrophon smithi differs in lacking spiralsculpture in later whorls and in its broader profile

i and wider lamellae. It occurs throughout the range ofB. stuarti

Genus Scabrotrophon McLean, new genus

I Type species (here designated): Trophon maltzani Kobelt and Kiister, 1878. Northeastern Pacific.

Diagnosis. Shell small to medium in size, canal moderately long, open; protoconch paucispiral, of

I 1.5 rounded, well-separated whorls. Sculpture axial and spiral; early sculpture dominated by 2 strong spiral cords; spiral cords scabrous, overriding axial ribs of mature sculpture.

Remarks. Scabrotmphon is characterized by the two dominant spiral cords of the early sculpture

I and strongly scabrous spiral cords of mature whorls. It differs from Boreotrophon, the dominant

in which

early sculpture is axial, in having spiral sculpture that overrides the axial ribs at all growth stages. In

Boreotrophon, spiral sculpture (if present) does not override the axial ribs.

I Eastern Pacific species of this genus have previously been allocated to Trophonopsis Bucquoy,

Dautzenberg, and Dollfus, 1882 [type species (OD): Murex muricatus Montagu, 1803; Europe], and, more

recently The to Nipgenus ponotro Trophonopsis phon Kurodais inapprop and Habe, riate19be71cause [typethespenorth cies Bore European otrophonand echinus Mediterranean Dall, 19type 18; Jsapepan] cies.

(see Egorov, 1993, figs. 34E, F) has dominant axial rather than spiral sculpture in the early whorls. The very

I pr Moreover, ominent axial thereriarebs are prominent overridenlabial by spiral denticles, sculpwhich ture thatarefonot rmsseen beadsin athet intersecti northeastern ons withPacifi thec axial species. sculpture.

Radwin and D' Attilio (1976: 82) used the genus Nipponotrophon, but that genus is characterized by

I aNipponotrophon strong rnid-whorl has angul been afollowed tion at alwith l growth out dissctages. ussionAlthough by subseqthe uentassignment authors, theofnortheastern eastern PaciPacific fic species speciesto

differ in having the two spiral cords in the early teleoconch and more extensive spiral sculpture at maturity. I

I dBoreotrophon o not consider , I cNi onsider pponotrScabrotrophon ophon to be toapbeplicrestricted able to antoythenoNorthern rtheasternHemisphere. Pacific species. Like the genus

Five species are treated here, including 2 new species. Four of these species were considered variants

I of the following species by such authors as Willett (1938) and Myers and D'Attilio (1980).

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Scabrotrophon maltzani (Kobelt and Kiister, 1878) I

Figure 1.19A

"Trophon tenuisculpta Carpenter, 1866."---of authors.

Trophon maltzani Kobelt and K_ster, 1878:301, pl. 75, fig. 17,18. •

Nipponotrophon maltzani: Houart, 1985:61, figs. 2-5. II

Trophon subserratus Sowerby, 1880 [1842-87], vol. 4, Trophon, p. 65, pl. 2, fig. 32-33.

Nipponotrophon subserratus.--Roth, 1981:58, figs. 1, 2. I

U

"Trophon (Trophonopsis) lasius (Dall, 1919)".--in part of Willett, 1938:10, pl. 1, figs. 2-5 [not 1, 6].

"Trophonopsis lasius (Dall, 1919)".--in part of Abbott, 1974:191, fig. 2000 [left only], n

"Nipponotrophon lasius (Dall, 1919)".---ofRadwin and D'Attilio, 1976:84, pl. 13, fig. 6-8.

Material Examined. Not represented in the voucher collection, but occurring offshore in southern •

California. Material examined: 138 LACM lots. II

Description. Shell white, thick and heavy, medium-sized; spire high, whorls weakly shouldered,

canal moderately long. Axial ribs 10-12, rounded to sharp, strongly developed on early whorls, sometimes

faint on final whorl. Spiral cords numerous, even, separated by narrower interspaces; cords finely imbricate.

Length 30-45 ram.

Type Locality and Type Specimens. Kodiak, Alaska; type material in Berlin Museum (Houart,

1985). Trophon subserratus: Vancouver Island, British Columbia; syntype material in NHML (Roth, 1981). HI

Distribution. Aleutian Islands, Alaska, to Cortes Bank, California [LACM 41-121.6] (32.5 °N).

Habitat. Rocky bottoms, 0-300 m. Occurring in the intertidal zone in Alaska, but deeply submergent I

at depths below 80 m in California.

Remarks. Early authors used the name Trophon tenuisculpta Carpenter, for this species, but syntype

material figured by Palmer (1958, pl. 23, figs. 9-12) has a short canal; the syntypes are here identified as an

immature stage of Ocenebrina interfossa (Carpenter, 1864), in which the canal had not achieved its sealed

mature form. I Dall missed two early names for this species. Roth (1981 ) recognized Trophon subserratus Sowerby,

1880, from British Columbia, and Houart (1985) showed that the earliest name for this species is Trophon

maltzani Kobelt and K/Jster, 1878, which has an Alaskan type locality. However, I do not follow Houart R

(1985) in placing Trophon (Boreotrophon) scitulus Dall, 1891, and Neptunea (Trophonopsis) lasia Dall,

1919, in the synonymy of Scabrotrophon maltzani.

The LACM collection has 20 lots of this species from offshore localities in southern California.

These specimens show the same range of variation as those from more northern localities. u

Scabrotrophon eerritensis (Arnold, 1903) U

Figure 1.19B

m

Trophon (Boreotrophon) cerritensis Arnold, 1903:249, pl. 6, fig. 6 [Pleistocene]. I

"Nipponotrophon scituIus (Dall, 1891)."---_'npart ofMyers and D'Atfilio, 1980:84, figs. 6, 7 [not figs. 1-5, 8-14].

II

Material Examined. No specimens are represented in the voucher collection from the Santa Maria I

Basin. Other material: 9 lots in the LACM collection.

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Figure 1.19. Muricidae. A. Scabrotrophon maltzani (Kobelt and Kiister, 1878): 265-274 m, SE of Santa Catalina

Island, California (LACM 40-146.6); height 21.5 ram. B. Scabrotrophon cerritensis (Arnold, 1903):

I 270 off San Clemente Island, California (LACM 150961); height 21.5 mm C. Scabrotrophon

m,

lasius 03all, 1919): 366-375 m, off SE end of San Nicolas Island, California (LACM 77-132.4);

height 22.9 mm. D. Scabrotrophon grovesi McLean, new species: 79-140 m, off NE side of Santa

I Ca newtalina specIislesan: d213, Cmaliforni , off aMon(CteASrey,07Mont 4962);erehei y County ght 28,.1Califo nun.rniE.a Scabrotrophon (LACM 2774, hclarki olotype)McL ; heiean, ght

43.4 ram. F. Ocenotrophonpainei 03all, 1903): 91 m, off Avalon, Santa Catalina Island, California

I (USNM 109306, holotype); height 14.2 ram.

I Description. Shell white, medium-sized; spire high but suture not deeply impressed, whorls shouldered, canal moderately long. Axial ribs 13-15; spiral cords two on early whorls, with relatively few on body whorl,

interspaces of same width; cords finely imbricate, producing cancellations on crossing axial sculpture. Length

20-33 mm.

Type Locality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene.

Holotype: USNM 162545. I

tI Distribution. Santa Rosa Island (34°N) to San Diego, California (33°N).

Habitat. Rocky bottoms, 110-270 m.

I Remarks. Scabrotrophon cerritensis was based on Pleistocene material and the name has been ignored in the literature. Here it is introduced for a species living offshore in southern California. As noted in the

synonymy above, Myers and D'Attilio (1980) figured a specimen of this species identified asNipponotrophon

scitulus [see comparisons under Scabrotrophon grovesi below].

I

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This species resembles Scabrotrophon maltzani in profile, but differs in its coarser and more cancellate !

sculpture and broader interspaces between the cords. The shoulder area has axial sculpture only, rather than

the three spiral cords of S. maltzani, and the body whorl has fewer and larger cords. !

Scabrotrophon lasius (Dall, 1919) m

Figure 1.19C I

Neptunea (Trophonopsis) lasia Dall, 1919:338.--Willett, 1938:11, pl. 1,fig. 6 [holotype]. 1

[] Trophon (Neptunea) lasius: Oldroyd, 1927:337.

Material Examined. Not represented in the Santa Maria Basin voucher collection, but occurring []

offshore in California. Material examined: 20 LACM lots. II

Description. Shell white, small to medium-sized; spire high, whorls rounded, canal moderately long.

Axial ribs irregular, low, sometimes sharply raised, sometimes lacking on f'mal whorl. Spiral cords freely _l

imbricate, numerous, even, separated by equal interspaces. Length 20-30 mm.

Type Locality and Type Specimens. Off Point Pifios, Monterey County, California, 198-495 fathoms. /

Holotype: USNM 210087.

m

Distribution. Off Graham Island, Queen Charlotte Islands, British Columbia (54°N), to N of San

Clemente Island, California (33°N).

Habitat. 90-550 m, bottom conditions unknown. 11

Remarks. Although this has been considered a synonym of Scabrotrophon maltzani, it differs in

having a thinner shell, finer spiral sculpture, and rounded rather than shouldered whorls. It usually occurs in •

deeper water than does S. maltzani, but depth ranges of the two species overlap. Willett (1938:11, pl. 1,fig. []

6) figured the holotype. Other shells illuslxated by Willett are here identified as S. maltzani (pl. 1,figs. 2-5)

and S. grovesi, new species (pl. 1, fig. 1). i

Scabrotrophon grovesi McLean, new species

Figure 1.19D II

"Trophon Iasius (Dall, 1919)."--in part ofWillett, 1938:10, pl. 1, fig. 1 [not figs. 2-5]. I

"Nipponotrophonscitulus (Dall, 189I)."--inpartofMyersandD'Atlflio, 1980:84, figs. 1-5, 8 [not figs. 7, 9-13].

Material Examined. Santa Mafia Basin, Phase I, sta. 16, 591 m (1). Other material examined: I

LACM 39-4.9, off White Cove, Santa Catalina Island, California, 73-146 m (1); LACM 41-9, off San Pedro,

Los Angeles County, Cafifornia, 148-152 m (1); LACM 41-19, off Wilson Cove, San Clemente Island,

California, 95-112 m (1); LACM 41-20, off San Diego, California, 143-148 m (1); LACM 41-56, off Point I

Dume, Ventura County, California, 86-88 m (1); SDNHM 73594, off La Jolla, San Diego County, California,

91-123 m (1); SDNHM 22951, off Santa Catalina Island, California (I).

Description. Shell white, medium-sized; spire high, whorls 5; canal long, open, length of canal equal 1

to length of aperture. Spiral sculpture of 2 strong cords on early whorls, uppermost strongest. Uppermost cord

with open, laterally projecting spines developing by third whorl. Penultimate whorl with shoulder cord and 1-

3 lesser cords (3 in holotype). Body whorl with strong shoulder cord and 5-7 additional primary cords; canal _l

with about 5 weak cords; cords becoming spinose on crossing axial sculpture; spines of much lesser strength []

than those of shoulder cord; interspaces between cords of nearly same breadth as spiral cords, uppermost

I
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I interspaces broader. Axial ribs 11-12 on mature shell, forming square cancellations with spiral cords of early

whorls; axial sculpture strongly developed only where crossing spiral cords. Canal with imbricate fasciole

I Length produced28.1bymmsuccessive (holotype). growth stages corresponding to production of axial sculpture. Inner lip thin, raised.

Type Locality and Type Specimens. Off NE side Santa Catalina Island, California (33°N, 118°W);

I 7LACM 9-140 m2781; on pebbles 1 paratype, and stones. SBMNH7 specimens. 142897; 1Holotype paratype USNM CAS 074962; 887572.3 paratypes CAS 075613; 1 paratype

Distribution. Point Buchon, San Luis Obispo County (35°N), to La Jolla, San Diego County, California

I (33ON)"

Habitat. 70-590 m, hard bottoms.

i Remarks. This species has previously been figured, as noted in the above synonymy. Willett (1938) considered it an "ornately frilled" form of "Trophonopsis" lasius (Dall, 1919) and Myers and D'Attilio

(1980), treated it as "Nipponotrophon" scitulus (Dall, 1891).

I by RadwinScabrotro and D'phon Attilliogro(1974, vesi differs pl. 14,from figs. the 10, Alaskan 11), in itsspec smaller ies Ssize, . scituIus more, whi slender ch has profile, best been shorterilluandstrated less

sloping shoulder, and presence of lamellar spiral cords on the canal, which are not present in S. scitulus.

i Spiral cords are fewer and interspaces broader than those of S. maltzani, in which the interspaces between cords are narrower than the cords.

There are forms of Scabrotrophon maltzani in which there is a prominent shoulder angulation (see

I Radwin are and D'Attilio, 1974, pl. 13, fig. 7), but S. grovesi differs from such specimens in having spines that longer and has fewer spiral cords on the body whorl.

The holotype specimen has relatively short spines compared to others in the same lot. Other specimens

have even shorter spines (LACM 41-9; LACM 41-19; LACM 41-20; LACM 41-56, as cited above). Such specimens may be distinguished from other species of Scabrotrophon in having higher, more elongate spires,

with the suture more tightly constricted.

I This is named after Groves of the Section, Los Museum species Lindsey Malacology Angeles County

of Natural History.

Scabrotrophon clarki McLean, new species

I Figure 1.19E

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material in

addition to holotype: LACM 41-172.13, 6.5 miles ESE of South Point, Santa Rosa Island, California, Velero

I 17I,sta. 1, m ( juvenile, length mm).

1392-4 104 1 9 m4

Description. Shell white, large for genus, spire moderately high; whorls five, tabulate; canal long,

I open, diameter length1.3ofmm. canal Earlyequalsculpture to lengthofof2 a

strong perture.spirProtoconch al cords, crossed paucispiral, by axial whorls ribs1of.2,lesser sharplystrength defined,thatmaximum produce

square cancellations. Second teleoconch whorl with upturned spines, which become long and re,curved toward

i cowithiling axis on third whorl. Body whorl with strong shoulder cord and four additional cords; base andcanal 5 cords of lesser strength. Cords forming open spines on crossing fluted axial ribs; recurved spines of

shoulder about 3 times longer than other spines. Spaces between cords with fine spiral striae, except those of

base and canal. Axial lamellae 14 on final whorl and on penultimate whorl. Canal with imbricate fasciote

produced by growth stages corresponding to production of axial sculpture. Columella white, concave, inner

lip thin, raised, reflected over projecting lamellae of base and canal. Length 43.4 mm (holotype).

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Type Locality and Type Specimens. Off Monterey, California, 213 m (700 feet), in shrimp pot of I

Tom Ghio (WatsonviUe, California), received from Roger Clark, 1994. Exact position unknown. The holotype

was evidently live-collected, butthe operculum was not saved. Holotype LACM 2774. t

Distribution. Monterey (37°N) to Santa Rosa Island, California (34°N).

Habitat. 100-210 m, bottom conditions unknown. The shallow limit is based on the juvenile specimen

cited above and the lower limit is based on the collection depth of the holotype. 1

m

Remarks. This species has not been previously illustrated. It is known from two specimens, the

holotype from Monterey and ajuvenile shell of 2.4 whorls from S of Santa Rosa Island. I1

Although Scabrotrophon clarki resembles S. grovesi inits tabulate shoulder with upturned spines, it I

differs in overall size, size of protoconch, whorl profile, and details of sculpture. The holotype ofS. clarki is

approximately twice as long as the known specimens ofS. grovesi, but it is not a larger than usual specimen •

of that species because it has the same number of whorls (5) and a larger protoconch. Compared to that I

species, the early whorls ofS. clarki are more vertically compressed and spines develop on the second rather

than third whorl. The recurved spines, which are bent toward the coiling axis, are characteristic and do not i

occur in S. grovesi. The usual number of axial lamellae orS. clarki is 14, compared to 12 in S. grovesi.

|

Scabrotrophon clarki also resembles the Alaskan species S. scitulus (Dall, 1891), from which it

differs in its larger size, having a more slender profile, having recurved spines, and having strong cords on the •

canal, which are lacking in S. scituIus. !1

The species is named after Roger Clark, who provided the holotype.

Genus Ocenotrophon McLean, 1995

Ocenotrophon McLean, 1995. Type species (OD): Murex (Ocinebra?)painei Dall, 1903. California. I

I

Diagnosis. Shell small, sculpture of numerous lamellar cords of unequal prominence, crossed by

sharply raised ribs, forming nodes at intersections; early teleoconch sculpture of two spiral cords, canal short, I

sealed. i

Remarks. This genus differs from bothBoreotrophon and Scabrotrophon in having the canal closed

at maturity. D'Attilio (1980:6) discussed the anomalous shell characters of the type species, placing it in •

Trophon Mont-fort, 1810, on the basis of radular characters, although the closed canal is more like that of U

Ocinebrina. The genus is monotypic.

In

Ocenotrophonpainei (Dall, 1903) I

Figure 1.19F I

Murex (Ocinebra?)painei Dall, 1903:174.--Da11, 1921:107, pl. 6, fig. 1.

Tritonaliapainei: Dall, 1921:107, pl. 6, fig. 1.--Oldroyd, 1927:318, pl. 32, fig. 7. I

? Ocenebra painei: Radwin and D'Attilio, 1976:123, pl. 20. fig. 8.

Trophon painei: D'Attilio, 1980:6, figs. 1-4. I

Oc enotrophon painei: McLean, 1995:40. !1

Material Examined. Not represented in the Santa Mafia Basin voucher collection, but occurring it

offshore in southern California. Material examined: 31 LACM lots. |

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I Description. Shell white, small; spire of moderate height, whorls strongly shouldered, suture deep,

canal short, sealed, fasciole broad. Axial ribs about 15, narrow and projecting, crossed by spiral cords of

slightly lesser strength, forming weak nodes at intersections. Inner lip sharply raised, outer lip smooth within.

I Length 12-15 mm.

Type Locality and Type Specimens. Off Avalon, Santa Catalina Island, California (from specimen

I label). Lectotype (here designated): USNM 109306. Distribution. Queen Charlotte Sound, British Columbia [LACM 70-87] (51°N) to Punta Rompiente,

Baja California [LACM 71-168] (28°N).

I Habitat. Rocky bottoms, 110-270 m.

Remarks. This species can not be confused with any other. It recalls species of Ocenebrina, but

i differs in lacking color and denticles within the outer lip.

i Family Turbinellidae Swainson, 1835

Diagnosis. Shell fusiform, usually with long anterior canal; with or without up to 4 columellar

plications; protoconch paucispiral, bulbous. Radular ribbon small, rachidian tricuspid, laterals unicuspid or

bicuspid.

Biology. The animals feed on polychaete or sipunculid worms. The paucispiral protoconchs indicate

I direct development. Remarks. Harasewych (1987) reviewed anatomical characters and outlined a classification for the

family. Recognized subfamilies are Vasinae, Columbariinae, Ptychatractinae and Turbinellinae. Only the

I subfamily Ptychatractinae is represented in the northeastern Pacific.

I Key to Species of Turbinellidae

IA. Penult_nate whorl bulging, axial ribs strong .................................................. Exilioidea rectirostris

I 1B. Penultimate whorl faintly rounded, axial ribs weak ............................................... Exilioidea kelseyi

I Subfamily Ptychatractinae Stimpson, 1865

i DBioiaglonos gy.is.Genera Shell small, inhabitfucold siform,waters, high-spire at bathyal d; aperture to abyssal narrowdepths, ; columellar exceptteethfor 0-4. the more shallow

occurrence of boreal groups.

I Remarks. Harasewych (1987) provides further notes on the subfamily.

Genus Exilioidea Grant and Gale, 1931

I Exilioidea Grant and Gale, 1931.Type species (OD): Chrysodomus rectirostris Carpenter, 1864. Northeastern

Pacific.

I Diagnosis. Shell fusiform, tall-spired, with long anterior canal. Sculpture of retractively curvedaxial

ribs and f'mespiral cords. Operculum elongate, filling aperture,nucleus anterior. Radular ribbon extremely

minute, total length not exceeding 0.14 mm.
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Figure 1.20. Turbineltidae, Buccinidae. A. Exilioidea rectirostris (Carpenter, 1864): 426 m, Santa Barbara i

Channel, California (USNM 213359); height 33.7 mm. B. Exilioidea ketseyi (Dall, 1908): 91 m,

off San Diego, California (USNM 110631, holotype); height 33.5 mm. C. Neptunea amianta (Dall, I

1890): 757 m, off Santa Barbara Islands, California (USNM 96556, holotype); height 72 mm. D.

Neptunea tabulata (Baird, 1863): 137 m, off Anacapa Island, California (LACM 150502); height

91.2 mm. I

Remarks. Bouchet and War6n (1988: I73) first aUocated Exilioidea to the Turbinellidae, having i

found the extremely minute radula that had eluded Bartsch ( 1945:61), who had tentatively placed the genus in

the Fusinidae (now Fasciolariidae). Bouchet and War6n concluded that there is a single highly variable Im

northeastern Pacific species, E. rectirostris, but I do not agree and here reinstate E. kelseyi as a separate i

species, as discussed below.

gxilioidea rectirostris (Carpenter, 1864) I

Figure 1.20A I

Chrysodomus rectirostris Carpenter, 1864: 664.--Arnold, 1903:228, pl. 7, fig. 7.

Tritonofusus (Plicifusus) rectirostris: Dall, 1902: 525, pl. 34, fig. 2. i

il Exilia rectirostris: Oldroyd, 1927:206, pl. 6, fig. 7 [copy Dall fig.].

Exilioidea rectirostris: Grant and Gale, 1931:665, pl. 28, fig. 5.--Bentson, 1940:224, pl. 2, figs. 23 [holotype],

24.--Bartsch, 1945:61, pl. 7, figs. 2, 3, 6.--Palmer, 1958:215.--Abbott, 1974:216.--Bouchet and i

War6n, 1988:174, figs. 16, 18, 22 [not figs. 17, 19-21].

i

Plicifusus (Microfusus) obsoletus Talmadge, 1971: 42, figs. 3,4. I

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I Material Examined. Not represented in the voucher collection from the Santa Mafia Basin. Other

material examined: 27 lots in the LACM collection.

I less thanDehalfscripshell tion. length, Fusiform, axial sculpture whorls 6, ofapinumerous cal area eroretractively ded in maturecurved specimens; fibs, obsolete siphonaloncanal lowerlong, half aofperture shell;

spiral sculpture of fine raised striae. Shell white with mid-whorl band of light brown. Periostracum dark

I yellow-brown, lighter in color across base, columella white. Length 25-30 mm. Type Loeatity and Type Specimens. Puget Sound, Washington. Holotype: USNM 4515. Plicifusus

obsoletus: OffEureka, Humboldt County, California, 450 fathoms; holotype: CAS 063623 (ex CAS 13319).

1

:_ Distribution. Behm Canal, Alaska (DaU, 1921:92) (55°N), to San Diego, California [USNM 122621]

I

(33°N).

i Ha Rebmarks. itat. Soft Specimens bottoms,from60-800 depthsm. below Below200220mmarein relatively southern California. broad with inflated whorls and strongly

projecting axial fibs, as typified by the population described by Talmadge (1971 ) under the name Plicifusus

I obsoletus.

Exilioidea kelseyi (Dall, 1908)

I Figure 1.20B

Tritonofusus (Plicifusus) kelseyi Dall, 1908:249. Exilia kelseyi: Dall, 1921:92.--DaU, 1925:17, pl. 1, fig. 6.

Exilioidea kelseyi: Bartsch, 1945:62, pl. 7, figs. 1, 4, 5.--Abbott, 1974:216, fig. 2388.

I Material Examined. Not represented in the voucher collection from the Santa Mafia Basin. Other

material examined: 15 lots in the LACM collection.

I Description. Slender, 7, apical area mature specimens; siphonal long, aperture

whorls eroded in canal

less thanhalf shell length; axial sculpture of numerous slightly curved ribs, becoming obsolete onlater whorls

and dorsal side of mature shell, not extending to lower half of shell, spiral sculpture of faint striae. Shell

I cream-colored with mid-whorl band of light brown, columeUa white. Periostracum dark brown. Length

grayish

25-30 mn_

Type Locality and Type Specimens. Off San Diego, California, 50 fathoms. Holotype: USNM

I 110631.

Distribution. Strait of Georgia, British Columbia ILACM 63-17] (50°N), to Redondo Beach, Los

!1 Angeles County, California [LACM 10935] (34°N).

Habitat. Soft bottoms, 90-200 m.

Remarks. This is more slender and has less convex whorls than Exilioidea rectirostris. The

I periostracum is darker than that of E. rectirostris. The species is here reinstated and removed from the

synonymy ofE. rectirostris, under which it was placed by Bouchet and Wartn (1988:173), Specimens figured

by Bouchet and Wartn that I identify as E. kelseyi are shown in their figures 17, 19-20. Although slender

i specimens with strong axial fibs are known, the ribs become nearly obsolete on the dorsal side of the final whorl. Both species occur together at some stations and are morphologically very close, but separation to two

species is possible.

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I 101
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Family Buccinidae Rafmesque, 1815 I

terminal.DRadula iagnosis.rachiglossate; Shell small torachidi large,anwithwithhigh1-5spire; cusps; canal laterals long towithshort. 2-6Operculum cusps. ovate, nucleus usually I

Biology. Buccinids are carnivores that either feed on living prey or are scavengers.

Remarks. Ponder and Wartn (1988) placed three additional groups usually treated as full families I

(Nassariidae, Melongenidae, Fasciolariidae) as subfamilies of Buccinidae, but I retain the traditional separation

of the families of the recommendation of M. G. Harasewych. Subfamily classification of Buccinidae proper is

unsettled and no division into subfamilies is utilized here. I

Key to Species of Buccinidae I

1A. Whorls rounded ..................................................................................................... Neptunea amianta II

lB. Whorls shouldered ................................................................................................ Neptunea tabulata I!

Genus Neptunea Rtcling, 1798 I

Neptunea R&ting, 1798. Type species (SD, Sandberger, 1861); Fusus antiquus Linnaeus, 1758. Northeastern

Atlantic. I Golikovia Habe and Sato, 1973. Type species (OD): Neptuneafukuae Kira, 1959. Japan.

Diagnosis. Shell large, sturdy, profile broad, with twisted siphonal canal; sculpture chiefly spiral, II

protoconch paucispiral, smooth; operculum large, f'filing aperture, nucleus terminal. 1

Biology. Neptunes in Alaska are chiefly predators on living bivalves and large tubiculous polychaetes

(Shimek, 1984). In feeding they make use of a highly extendible proboscis. Upright egg capsules are laid in l

clusters. II

Remarks. The neptunes are large and important gastropods of the Arctic and northern oceans. They

have been monographed inan unpublished thesis by C. Nelson (1974) and by Golikov (1963). Nelson (1976:139) 1

corrected earlier errors in the type designation forNeptunea. The origin of the genus was discussed by Nelson I

(1978). Northern species are highly variable and the taxonomy remains unsettled.

!

Neptunea a_nta (D_I, 1890)

Figure 1.20C I

Chrysodomus amiantus Dall, 1890:321, pl. 5, f. 10.---Oldroyd, 1927:96. I

Neptunea amianta: Abbott, 1974:213, fig. 2358. I

Material Examined. Not represented in the voucher collection from the Santa Maria Basin. Other 1

material examined: 102 lots in the LACM collection. II

III

Description. Shell of moderate size for genus; whorls 4, thin, chalky white under thin brown

periostracum. Early whorls with strong peripheral carination, mature whorls rounded. Spiral sculpture of l1

numerous narrow, low cords of unequal strength and spacing; axial sculpture of growth increments only. |

Canal short, lip usually thin and easily broken, seldom flaring when mature. Length 60-90 mm.

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102 I
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I Type Locality and Type Specimens. "SantaBarbara Islands," California, 414 fathoms. Lectotype

(here selected): USNM 96556.

I CaliforniaDistr [LACM ibution. 93-76.3] Pribilof(31Islands, °N). Alaskan BeringspeSeacimens , Alaskaare[UfewSNMin c,olLAC lectioM] ns.(57The°N)species , toPunbecta oSan mesJocsr, ommBaojna

south of the Queen Charlotte Islands, British Columbia.

I Habitat. Soft bottoms, 300-1500 m.

Remarks. Neptunea amianta resembles N. pribiloffensis (Dall, 1919), a much larger species of

i broader profile that ranges from the Bering Sea to British Columbia.

Neptunea tabulata (Baird, 1863)

i Figure 1.20D

Chrysodomus tabulatus Baird, 1863:66, pl. 1, f. 1.--Dall, 1902:524, pl. 36, fig. 5.--Arnold, 1903:228, pl.

I 7, fig. 6.--Oldroyd, 1927:227, pl. 4, fig. 5, pl. 18, fig. 4.

Neptunea (Sulcosipho) tabulata: Grant and Gale, 1931:658.--Abbott, 1974:215, fig. 2374.

I Material Examined. Not represented in the voucher collection from the Santa Maria Basin. Other

material examined: 53 lots in the LACM collection.

I periostracum. Description. Suture Shell deeplymodimpressed, erately large shoulder for gestrongly nus; whotabulate, rls 7, sturdy, smooth, cream-c marked oloredbyunde sharpr thin, projecting brown

carination; sculpture below shoulder of numerous, narrow, closely spaced, imbricate spiral cords; canal long,

i twisted; TyfascpeioleLocality producing and Tyumbpeilical Specchink; lip thick and slightly flaring at maturity. Length 80-110 mm. imens. Vancouver Island, British Columbia. Holotype: Natural History

Museum, London.

I Distribution. Petersburg, Alaska [LACM 147302] (57°N), to Newport Bay, Orange County, California

[LACM 70-69.13] (33.5°N).

Habitat. Soft bottoms, 50-200 m.

I Remarks. This is the type and only living species of the subgenus Sulcosipho, Dall, 1918, which

denotes the deep suture and tabulate shoulder. There is a Miocene predecessor, Chrysodomus bairdii Dall,

i 1909, which Grant and Gale (1931:658) placed in synonymy.

I Family Nassariidae Iredale, 1916

Diagnosis. Shell small to medium in size, with high spire, ovate aperture, and short anterior canal;

i posterior anal canallacking, columella short andtwisted, usually with well-formed columellar callus. Operculum ovate, edge often serrate.Radula stenoglossate, with multicuspid central and two-pronged lateral teeth.

Biology. The animals are carnivorous and are chiefly scavengers on sand and mud bottoms. Eggs are

I laid in caRpsules emarks.andThe in mmost ost species comprehensive there are free swimming veliger larvae. source of information is the major monograph of Cemohorsky

(1984) in which three subfamilies were defined: Nassariinae, Dorsaniinae, and Cylleninae. Cemohorsky treated

I inform. full the Indo-Pacific species of Nassariinae and Cylleninae and all others were treated in at least checklist

I

i 103
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Key to Species of Nassariidae I

1A. Columellar wall with prominent pustules .......................................................... Nassarius insculptus I

1

lB. Columellar wall smooth ................................................................................................................... 2

2A. Shell length 18-22 mm ..................................................................................... Nassarius perpinguis I

2B. Shell length 22-35 mm ......................................................................................... Nassarius rhinetes

Genus Nassarius Dtimeril, 1806 I_

Nassarius Dfmeril, 1806. Type species (SM, Froriep, 1806): Buccinum arcularia Linnaeus, 1758. Indo- I_

Pacific. !1

Diagnosis. Base of shell with deep groove or fossa; columella concave; siphonal notch deep, siphonal 1

canal short; sculpture spiral and axial, often cancellate. Operculum with serrate edge on side adjacent to outer 1

lip.

Remarks. Demond (1952) and Cernohorsky (1975) reviewed the northeastern Pacific species of

Nassarius. Addicott (1965) treated fossil and living species of the northeastern Pacific. A large number of

taxa were recognized as subgenera of Nassarius by Cemohorsky; those in use for the northeastern Pacific

species are mentioned under the remarks section of each species. I

An additional species that may be present at depths to 200 meters is the well-known shallow-water

species Nassarius mendicus (Gould, 1850) and its forms N. cooped (Forbes, 1852), and N. indisputabiIis

(Oldroyd, 1927). This species has been treated in detail by Demond (1952:308), Addicott (1965:3), Abbott 1

(1974:225) and Cemohorsky (1975:124).

Nassarius perpinguis (Hinds, 1844) I

Figure 1.21A

Nassa perpinguis Hinds, 1844 [1844-45]:36, pl. 9, figs. 12, 13. I

Nassarius (Schizopyga)perpinguis: Grant and Gale, 1931:673, pl. 26, figs. 51, 52.

Nassarius perpinguis: Demond, 1952:305, pl. 2, figs. 4, 5.---Cernohorsky, 1975: 123, figs. 1, 2 [holotype].-- I[

McLean, 1978:48, fig. 26.3.

u

Nassarius (Hinea)perpinguis: Abbott, 1974:224, fig. 2460.---Cemohorsky, 1984:35.

Nassa perpinguis var. bifasciata Berry, 1908:39. I

Material Examined, California: Santa Mafia Basin, Phase I, sta. 21, 49 m (1 immature). Other Ill material: 231 lots in the LACM collection.

Description. Shell of moderate size, white to light brown under closely adherent periostracum, with

darker bands of brown; whorls evenly rounded, fossa deep; axial and spiral sculpture of nearly equal strength, [i

producing fine, even cancellations; mature lip only slightly thickened, deeply lirate within; columella with fine I!

raised lines on lower portion; parietal shield with single raised fold. Length 18-22 mm.

Type Locality and Type Specimens. Bahia Magdalena, Baja California Sur (probable error), l!

Holotype, NHML 44.9.23.5. Nassa perpinguis var. bifasciata: San Pedro, Los Angeles County, California;

lectotype (here designated): CAS 064634 (Figure 1.18A).

Distribution. Point Reyes, Marin County, California (38°N) to Isla Cedros, Baja California (28°N) I

104 |
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I Figure 1.21. Nas California sariidae,(CAS Fascio064364, lariidae. leA.ctotype Nassarius of perpinguis Nassa perpinguis (Hinds, vat. 1844): bifasciatus San Pedro, ); Lheight os Angele 21.2s nuConun. ty, B.

Nassarius rhinetes Berry, 1953:68 m, Monterey Bay, California (USNM 212133); height 33 ram.

C. Nassarius insculptus (Carpenter, 1864): 174-247 m, off Point Loma, San Diego County, California

I (USNM 209046,Alectrion insulptus vat. eupleura, holotype); height 15 nun. D. Fusinus barbarensis (Trask, 1855): 91 m, S of Santa Catalina Island, California (SBMNH 118915); height 98 ram.

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Habitat. Soft bottoms, intertidal to 60 m.

I Remarks. This is the type species of the subgenus Caesia H. and A. Adams, 1853, which has been

used for most of the northeastern Pacific species of Nassarius (see Cernohorsky, 1984:32). Type material of

two additional synonyms from erroneous localities was illustrated by Cemohorsky (1975).

I

Nassadus rhinetes Berry, 1953

I Figure 1.21B

"Nassa californiana (Conrad, 1856)."----ofDall, 1891: 177.--Dall, 1921:102, pl. 11, fig. 4.

I "'Nassarius (Schizopyga) californianus."--ofGrant and Gate, 1931:672, pl. 26, fig. 49.--Oldroyd, 1927:264,

pl. 26, fig. 13.

i Nassarius "'Nassarius californianus."--ofDemond, 1952:206, pl. 2, fig. 6. (Schizopyga) rhinetes Berry, 1953: 415, pl. 28, f. 7.

Nassarius (Caesia) rhinetes: Addicott, 1965:10, pl. 2, fig. 28.

I Cemohorsky, 1984:35.

Nassarius rhinetes:

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material:

I 13 lots in the LACM collection.

I 105
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Description. Shell large for family, light brown with lighter bands under closely adherent periostracum; I

whorls evenly rounded, fossa deep; axial ribs numerous, slanted; spiral cords 6 on penultimate whorl; axial

and spiral sculpture of nearly equal strength, producing beaded intersections; columella concave, projecting I

and twisted at base; mature lip only slightly thickened, lirate within. Length 22-33 mm. |

Type Locality and Type Specimens. Off Moss Landing, Monterey County, California, 40 fathoms.

Holotype: SBMNH 34536. I

U Distribution. Off Punta Delgada, Humboldt County, California [LACM 71-127.2] (40°N), to Bahfa

Blanca, Baja California [LACM 51-13.9] (29°N). m

Habitat. Soft bottoms, 10-91 m. I

immff

Remarks. Early authors misidentifed this species as Nassa californiana Conrad, 1856, a species

shown by Berry (1953) and Addicott (1965:3) to be extinct. It is closest to Nassarius perpinguis, but is larger •

and has coarser clathrate sculpture. Interspersed spiral striae as inN. perpinguis are not present. This species |

is much less common than N. perpinguis. It is a more characteristic species of northern and central California

than of southern California. I

II

Nassarius inscu_tus (Carpenter, 1864)

Figure 1.21C 1

Nassa insculpta Carpenter, 1864:662 [unfigured]. •

Nassarius insculptus: Demond, 1952:317, pl. 2, figs. 1, 3.--Palmer, 1958:214.---Cemohorsky 1984:34.

Nassarius (Uzita) insculptus: Addicott, 1965:11. ==

Nassarius (Nassarius) insculptus: Abbott, 1974: 224, fig. 2455. I

Alectrion insculptus var. eupleura Dall, 1917: 576.

Nassarius insculptus gordanus Hertlein and Strong, 1951: 81, pl. 8, fig. 6. I

Material Examined. Not represented in the voucher collection, but occurring abundantly offshore in

southern California. Material examined 376 LACM lots. I

Description. Shell of moderate size, white under closely adherent greenish-brown periostracum; !1

whorls 7, whorls slightly inflated; axial sculpture of strong ridges on early whorls, usually fading on later

whorls; spiral sculpture of shallow grooves, about 10 per whorl; fossa shallow; outer lip inflated and thickened,

inner lip with broad shield of callus, one strong basal fold and smaller raised folds becoming pustular near

termination of inner lip callus. Length 18-25 mm.

Type Locality and Type Specimens. Santa Catalina Island, California. Type material not found by

Palmer (1958:214). Alectrion inscuIptus var. eupleura: Off Point Loma, San Diego County, California, 95-

135 fathoms (type locality from holotype label); holotype: USNM 209046. Nassarius insculptus gordanus:

Gorda Bank, off S tip Baja California Sur, 60 fathoms; holotype: CAS 065081. I

m

Distribution. Point Arena, Mendocino County, California (39°N), to Gorda Bank, Baja California

(23°N). Also in the Gulf of California and at Isla Clarion, Islas Revillagigedos, Mexico.

Habitat. Soft bottoms, 50-530 m. This is perhaps the most abundant gastropod of its size occurring 1

offshore in southern California.

Remarks. This species is the onlyNassarius in the northeastern Pacific with the shell nearly smooth I

except for the incised sculpture.It was placed by Addicott (1965:11 ) in the subgenus Uzita H. and A. Adams, |

1853, characterized by the fine plaits on the inner lip. Related species occur off west Africa.

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i FamilyFasciolariidae Gray, 1853

I Diagnosis. Shell of medium to large size, anterior canal usually long. Animal red colored. Biology. Carnivores, mostly preying on mollusks.

Remarks. Subfamilies are: Fusininae, Fasciolafiinae, and Pefistemiinae. The genus Fusinus is assigned

I to the Subfamily Fusininae Wrigley, 1927.

I Genus Fusinus Rafinesque, 1815

Fusinus Rafmesque, 1815. Type species (SM, Lamarck, 1799; ICZN Opinion 1765): Murex colus Linnaeus,

1758. Indo-Pacific.

I Gracilipurpura Jousseaume, Type species (OD): strigosus Lamarck, 822; = rostratus

1880. Fusus 1 Murex

Olivi, 1792, not Solander in Brander, 1766.

i Bar Harfordia barofususDall,Grabau 1921.andTypeShimer, spcies1909. (M):TypFusus e speharfordi cies (OD):Steams, Fusus 1871. barbarensis EasternTrPacific. ask, 1855. Eastern Pacific.

i and Diagnosis. Shell elongate, fusiform, with tall spire and long siphonal canal; sculpture of axial ribs spiral cords; columella without folds. Protoconch paucispiral, early teleoconch whorl with strong axial

fibs. Rachidian with three short cusps; laterals multicuspid.

I and War6nRem(1985:160), arks. The gewho nus did has not not been distinguish reviewedsubgenera worldwide. except The geforneric the shallow-water synonymy aboveAptyxis is fromTroschel, Bouchet

1868.

I Fusinus barbarensis (Trask, 1855)

i Figure 1.21D

Fusus barbarensis Trask, 1855: 55.--Arnold, 1903:224, pl. 4, fig. 10.

I Fusinus Woodring, barbarensis: Dall, 1915:55.--Oldroyd, 1927:177.----Grant and Gale, 1931:639, pl. 27, fig. ll.-- 1946:73, pl. 34, fig. 9.--Abbott, 1974:2519, fig. 230.

I materialMat examin erialed:Examined. 129 lots inNthoet LACM represented collecintion. the voucher collection from the SantaMaria Basin. Other

Description. Shell large; whorls 7, rounded; canal long, of same length as aperture; lip lirate within;

I axial fibs 8-12, strong on early whorls, usually lacking on later whorls; spiral cords 4 on early whorls,

secondary cords arising in interspaces so that final whorl is covered by numerous cords of irregular strength

and spacing; color white to tan under brown periostracum, cords often darker brown. Length 50-110 mm.

I and Type Specimens. Santa Barbara, Pleistocene; material lost (Woodring,

TypeLocality type

1946:73). The type specimen was not figured and was lost in the fire at the California Academy that followed

the 1906 earthquake.

I Distribution. Monterey Bay, California [LACM 60-21] (36.5°N), to Gorda Bank, off Cabo San

Lucas, Baja California Sur [LACM 40-7] (23°N).

I Habitat. Remarks. Soft Fusinus bottoms, 50-350 m. barbarensis is highly variable in number of axial ribs and strength and spacing of

the spiral cords. Large specimens are not common in collections although there are many immature specimens.

I the Specime vicinity ns frofomIslaBaCedros. jaCalifornia are somewhat smaller, although there are specimens that reach 90 mm from

i 107
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Family ColumbeUidae Swainson, 1840 i

Diagnosis. Shell minute to small; sturdy, outer lip thickened andusually denticulate within; columella II

smooth or plaited, siphonal canal short. Radula rachiglossate, rachidian rectangular, lacking cusps, lateral I

teeth elongate, bearing 2-3 cusps near tip.

Biology. Little is known of the biology of columbellids. There are both carnivorous and herbivorous

groups. Egg capsules are attached to differing substrata, including shells of the same species. Veliger larvae

are produced; the veliger stage occurs before hatching in most species. Boss (1982:1015) summarized the

recent literature on the anatomy and biology of the family. I

Remarks. Two subfamilies were recognized by Radwin (1977, 1978) in his review of western Atlantic

genera and species: Columbellinae, with the lateral teeth of the radula broad and flat and with strongly flexed

basal flanges, and Pyreninae, with the lateral teeth sigmoid and unflanged. I

I!

Species formerly assigned to Mitrella Risso, 1826, are now distributed among three genera in two

subfaroalies:Alia H. and A. Adams, 1853, which occurs in shallow water (Columbellinae); and,in the Pyreninae, lira

Astyris H. and A. Adams, 1853, and MitrelIa. MitreUa occurs in tropical waters and is characterized by its

columellar denticles (Radwin, 1978:337). l

Key to Species of Columbellidae I

l

1A. Sculpture lacking except for spiral grooves on canal ....................................................................... 2 I

lB. Sculpture of axial ribs and spiral cords ........................................................................................... 3

I

2A. Outer lip lirate ....................................................................................................... Astyris gausapata

2B. Outer lip not lirate ................................................................................................ Astyris permodesta !

3A. Whorl profile slightly concave below suture ........................................................... Amphissa undata •

3B. Whorls rounded ................................................................................................................................ 4 II

I

4A. Axial ribs > 15 per whorl ................................................................................... Amphissa reticulata

4B. Axial ribs < 14 per whorl ....................................................................................... Amphissa bicolor

I

Subfamily Pyreninae Surer, 1919

Diagnosis. Shells usually narrow, fusiform. Lateral teeth of radula sigmoid. .I

Remarks. Western Atlantic genera and species of Pyreninae were reviewed by Radwin (1978).

I

Genus Astyris H. and A. Adams, 1853 '_

Astyris H. and A. Adams, 1853. Type species (OD): Buccinum rosaceum Gould, 1840. Arctic Ocean. 1

Diagn_is. Shell small, high-spired, whorls smooth, lacking axial and spiral sculpture, except for

spiral incisions on base; aperture narrow, canal short but deeply notched; anal notch lacking; protoconch •

paucispiral. |

108 I
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I Biology. As for family. Species occur at shallow to offshore depths.

Remarks. The diagnosis given here is tentative, as Radwin (1978:331) neglected to diagnose this

I genus. He provided only a list of western Atlantic species that he allocated to it. Species that I group in Astyris lack or have weak labial denticles and have a paucispiral protoconch

indicative of direct development. Astyris aurantiaca (Dall, 1871) is a shallow-water member of the genus

I having a paucispiral protoconch. In contrast, shallow-water species of the columbelline genusAlia H. and A. Adams, 1853, including Alia carinata (Hinds, 1844) and A. tuberosa (Carpenter, 1864), have strong labial

denticles and a multispiral protoconch indicative of planktotrophic development.

I

AsCyris ga_apa¢_ (Gould, 1850)

I Figure 1.22A

Columbella gausapata Gould, 1850:170.--Gould, 1852:267, pl. 19, fig. 337.--Johnson, 1964:82 [citation

I of type material].

Nitidella gouldii Carpenter in Gould and Carpenter, 1857:208.---Oldroyd, 1927:277.--Abbott, 1974:198,

fig. 2103.

I Mitrella gouldii: Grant and Gale, 1931:695.

Columbella (Nitidella ?) dalli E. A. Smith, 1880:287 [unfigured].

I Nitidella ? lutulenta Dall, 1919:331 [unfigured].

Alia casciana Dall, 1919:330 [unfigured].

I Material Examined. California: Santa Maria Basin, Phase I, sta. 85, 113 m (1); sta. 102, 99 m (1); Phase II, sta. R-2, 161 m (1 immature); sta. PJ-7, 123 m (2 immature); sta. PJ-14 (1). Other material: 271 lots

in the LACM collection.

I Description. Shell small, thick, 5, except microscopic striations; spiral

whorls smooth for base with

sculpture; protoconch paucispiral, usually deeply eroded in mature specimens; lip with weak lirae away from

edge; periostracum adherent, brown; shell color light yellowish brown, with darker chevron or mottled markings.

I Length 8-13

mm.

Type Locality and Type Specimens. Puget Sound, Washington. Lectotype selected by Johnson

i (1964:82): USNM 5732. Nitidella gouldii: Santa Barbara, California; type material not located by Palmer (1958:210). CoIumbella dalli: Vancouver Island, British Columbia; type material in NHML. Nitidella lutulenta:

Gulf of the Farallones, off San Francisco, California, 24 fathoms; lectotype (here selected): USNM 211068.

Alia casciana: Off La Jolla, San Diego County, California, 110-199 fathoms; lectotype (here selected): USNM

I 209456.

Distribution. Bechevin Bay, NW side Alaska Peninsula, Alaska [LACM 90-197.1] (55°N, 163°W),

i to Punta Habitat. San PabloSo, fBaja t bottCoamslif,ornia 30-20Sur 0 rn[LACM . 71-176] (27°N).

Remarks. Gould's name Columbella gausapata is the older name for this species, based on a

I redetermination of the holotype, although authors including Grant and Gale (1931:695) had incorrectly considered it a synonym of the shallow-water species Alia carinata (Hinds, 1844). This species is better

known under the name Mitrella gouldii (Carpenter), as used by Grant and Gale (1931:695) and Abbott

I (1974:198). It is placed in Astyris because the paucispiral protoconch is indicative of direct development. Dali's taxaNitidella lutulenta andAlia casciana have been ignored in the literature but are here placed in the

synonymy of this species.

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Figure 1.22. Columbellidae. A.Astyris gausapata (Gould, 1850):82-91 m, offLong Point, Santa CatalinaIsland, 1

California (LACM 41-185.4); height 10.5 mm. B. Astyris permodesta (Dall, 1890): 505 m, off II

SantaBarbara Islands, California(USNM 96526,lectotype);height t2.5 mm. C.Amphissa reticulata

Dall, 1916:113-335 m, off Point Loma, San Diego County,California (USNM 210004, lectotype);

height 16 mm. D.Amphissa bicolorDall, 1892:563 m, off PointSur, Monterey County, California 1

(USNM 106877, syntype); height 13.8 mm. E. Amphissa undata (Carpenter, 1864): 68-73 m, S U

side Santa Cruz Island, California (LACM 40-165.31); height 12.0 mm.

!

AsCyris pe_odesta (Dall, 1890)

Figure 1.22B I

Columbellapermodesta Dall, 1890: 327, pl. 5, fig. 4. I

ColumbelIa (Astyris) permodesta: Oldroyd, 1927:275. I

MitreIla permodesta: Abbott, 1974:200, fig. 2141.

Material Examined. Califomia: Santa Maria Basin, Phase I, sta. 89, 471 m (18); sta. 99, 540 m (5); 'l'-

Phase II, sta. R-7,565 m (6). Other material: 35 lots in the LACM collection.

Description. Shell of moderate size for genus, broadly inflated, thin, white under greenish brown •

periostracum, but often coated with dark brown mineral deposits; apex usually eroded; sculpture lacking

except for weak spiral cords on base of shell. Lip thin, lacking lime; columella simple, translucent. Length 10-

14mm. I Type Locality and Type Specimens. Santa Barbara Islands, California, 276 fathoms. Lectotype

(here designated): USNM 96526 (Figure 1.19B).

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I Distribution. Monterey Bay, California, to Bahia Magdalena, Baja California Sur [USNM 265899]

(24.5 °N).

I Habitat. Soft bottoms, 300-1000 m. The species is characteristic of anoxic bottoms of offshore basins. It has also been recorded along with species of Calyptogena Dall, 1891, large vesicomyid bivalves

that are characteristic of reducing environments.

I Remarks. The genetic assignment is based on the paucispiral protoconch and the thin lip without lime. It differs from Astyris gausapata in lacking the lirae and in having the shell thinner and broader.

I Genus Amphissa H. and A. Adams, 1853

Amphissa H. and A. Adams, 1853. Type species (SD, Dall, 1913): Buccinum corrugatum Reeve, 1846, not

1 Brocchi, 1814 [=Amphissa columbiana Dall, 1916]. Northeastern Pacific.

Diagnosis. Shell large for family, high-spired with inflated whorls and deep suture; columella smooth,

I siphonalBcanal short, sculpture of axial ribs and incised spiral lines; lip lirate within; protoconch multispiral.

iology. The multispiral protoconch is indicative of a planktonic veliger stage. Eggs are laid in

capsules that release veliger larvae.

I Remarks. Amphissa is a characteristic genus of cool waters of the northeastern Pacific. Two species

occur in shallow water: A. versicolor Dall, 1871, and A. columbiana (Dall, 1916). There is also a single

species that occurs in the North Atlantic (Radwin, 1978:329).

I

Amphissa reticulata Dall, 1916

I Figure 1.22C

Amphissa versicolor vat. reticuIata Dall, 19t6: 27.

I Amphissa reticulata: Oldroyd, 1927:283.--Abbott, 1974:203.

Material Examined. California: Santa Maria Basin, Phase I, sta. 2, 200 m (1). Other material: 133

I lots in the LACM collection.

Description. Shell light tan under greenish brown periostracum, some with 2 light brown bands;

i whorls moderately inflated; axial ribs about 16 per whorl, nearly straight, fading across base; spiral cords strong, overriding axial ribs, producing beads at intersections; lip lirae relatively poorly developed. Length

12-16mm.

I LectotypeType (hereLocdalit esigynataednd):TUSNM ype Spe210004 cimens.(Figure Off Po1.22C). int Loma, TheSanoriginal Diegodescription County, California, cited only the 62-183 catalogm.

number; the type locality is taken from the specimen label.

I (28 oN). Distribution. Kenai Peninsula, Alaska (59.5°N, I51.5°W), to Islas San Benito, Baja California

Habitat. Soft bottoms, 30-300 m.

I

I

I

I 111
!

Remarks. Surprisingly, this species has not previously been figured. It differs from the shallow- I

water species Amphissa versicolor in its more slender profile, lack of color markings on the shell, and weak

development of the labial lirae. Unlike A. versicolor, it occurs offshore, frequently with Astyris gausapata. I

Amphissa bicolor Dall, 1892

Figure 1.22D l

Amphissa bicolor Dall in Williamson, 1892:213, pl. 20, fig. 4.----Oldroyd, 1927:282.--Abbott, 1974:203. I

Ii

Material Examined. California: Santa Maria Basin, Phase I, sta. 68, 390 m (11); sta. 72, 401 m

(21); Phase II, sta. R-6, 410 m (5). Other material: 65 lots in the LACM collection.

Description. Shell light tan under tan periostracum, marked with 2 broad brown bands, 1at midwhorl I[

and 1 on base; whorls strongly inflated; axial ribs 11-13 per whorl, fading across base, slanted to left; spiral

sculpture weak, consisting of incised striae overriding axial ribs; lip lirae not developed. Length 12-14 mm.

Type Locality and Type Specimens. Off Point Sur, Monterey County, California, 298 fathoms II

(from label). Lectotype (here designated): USNM 106877, based on the catalog number cited in the original

caption. I Distribution. Farallon Islands (37.5°N) to San Diego, California (33°N). Distribution limits from

Dall (1921:105). The LACM collection has specimens from Monterey to Newport Bay, Orange County,

California. I

Habitat. Soft bottoms, 90-600 m.

Remarks. Amphissa bicolor is easily recognized by its banded color pattern, broadly spaced axial II

ribs, subdued spiral sculpture, and lack of lip tirae. I

This species is more characteristic of greater depths than those known forAmphissa reticulata andA.

undata. Like Astyris permodesta, it has been found in association with Calyptogena in anoxic, reducing •

environments. However, unlikeA, permodesta, there are numerous lots known from normal habitats. II

I

Amphissa undata (Carpenter, 1864) i_

Figure 1.22E

aa

Amycla undata Carpenter, 1864: 224. l

Amphissa undata: Dall in Williarnson, 1892:214, pl. 20, fig. 8.--Oldroyd, 1927:282. Abbott, 1974:203,

fig. 2183.--Palmer, 1958:211, figs. 21, 22 [holotype]. l

w

Amphissa ventricosa Arnold, 1903: 242, pl. 5, fig. 11 [Pleistocene].

Material Examined. California: Santa Mafia Basin, Phase I,sta. 4, 393 m; Sta. R-4, 90 m (1). Other I

material: 272 lots in the LACM collection. g

Description. Shell tan with variegated pattern of lighter and darker brown markings under light tan

periostracum; whorls strongly inflated, suture deep; axial ribs 12-14, projecting strongly at periphery, fading !

across base; spiral cords strong but narrow, overriding axial fibs, interspaces wider than cords; lip lirae

moderately developed. Length 12-16 mm. I

Type Locality and Type Specimens. Santa Catalina Island, California, 40 fathoms. Holotype: USNM I

23284.Amphissa ventricosa: San Pedro, Los Angeles County, California; Pleistocene; holotype USNM 162550.

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!

I Distribution. Monterey Bay, California (36.5°N), to Isla Asuncitn Island, Baja California (27°N).

There are two lots in the LACM collection from the Gulf of California (off Isla Espiritu Santo and Puerto

I EscondidHo)a.bitat. Soft bottoms, 30-90 m.

Remarks, This species is characterized by its color pattern, deep suture and narrow spiral cords.

I There is a slightly concave shoulder, which imparts a profile that differs from that of the other two species treated here.

i Woodring (1946:77) first placed Arnold's Amphissa ventricosa in the synonymy ofA. undata.

I Family Cystiscidae Stimpson, 1865

Diagnosis. Shell small to minute, glossy, with low spire, thickened outer lip and columellar plications;

internal whorls partially resorbed; protoconch paucispiral. The mantle of living animals envelops the shell,

I its surface The radula is with rachidian tooth; lateral teeth keeping glossy. rachiglossate multicuspid a

lacking

in most genera.

i CoovertBiology. Cystiscids are carnivorous. The paucispiral protoconch is indicative of direct development.

and Coovert (1995) treatedthe anatomy and biology of the marginelliform families Cystiscidae and

Marginellidae.

I were separated Remarksat. theUntsubfamily il the worklevel of Coovert in the Marginellidae. and Coovert (1995), Thesetheauthors familiesdescribed Cystiscidae significant and Marginellidae anatomical

and radular differences in addition to the shell character distinction that cystiscids partially resorb the internal

I whorls, whereas Coovert margine and Coovert llids do not. (1995) recognized four subfamilies in the Cystiscidae. The genus that follows

was placed in the subfamily Plesiocystiscinae Coovert and Coovert, 1995.

I Genus Plesiocystiscus Coovert and Coovert, 1995

I Plesiocystiscus Coovert andCooverL 1995. Type species (OD): Marginellajewettii Carpenter, 1857. California.

Diagnosis. Shell minute, spire low, not concealed, anterior margin rounded, without a siphonal notch;

lip thickened, Remarks. smootEastern h within;Pacific lateralspecies teeth ofwere raduformerly la presentplaced . in Cystiscus Stimpson, 1865, a genus now

limited to South Africa. Eastern Pacific species of Plesiocystiscus were treated by Coan and Roth (1966) and

I Roth and Coan (1968).

Piesiocystiseus myrmeeoon (Dall, 1919)

I Figure 1.23A

I Hyalina Marg myrmecoon Dall, 1919: 308. inella (Hyalina ) myrmecoon: Oldroyd, 1927:165.

i CystiscusMaterial myrmecoon: Coan and Roth, 1966:291, pl. 51, fig. 65. Examined. California: Santa Maria Basin, Phase I, sta. 16, 591 m (2). Other material: 37

lots in the LACM collection.

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Figure 1.23. Cystiscidae, Cancellariidae. A, Pleisocystiscus myrmecoon (Dall, 1919): 591 m, off Point Buchon, =w

San Luis Obispo County,California (SBMNH 142804,Santa MariaBasin, sta. 16);height 3.9mm.

B. Cancellaria cooperi Gabb, 1865:124 m, off Santa Barbara, California (USNM 106860); height 1

56mn'LC. CancellariacrawfordianaDall, 1891:OffRedondoBeach,Los AngelesCounty,California II

(LACM 72505); height 49.8 mm. D. Admete californica (-Dall,1908): 1103 m, off San Diego,

California (USNM 110626,lectotype); height 15.8 ram. E. Admete graciIior (Carpenter, 1869): 69 •

m, off Point Fermin, Los Angeles County, California (LACM 57-90.1); height 11.4mm. I

1

Description. Shell small, white, smooth; whorls 3, columella with 3 prominent plaits and 3 smaller, m

more posterior plaits. Profile of lip straight to slightly concave. Length 3.3-4.8 mm. I

Type Locality and Type Specimens. San Diego, California. Holotype USNM 9440. 1

Distribution. Pacific Grove, Monterey County, California (36° N),to Ptmta Rompiente, Baja California

Sur (27.5 ° N).

Habitat. Intertidal to 40 m. The voucher specimens from 591 m were live-collected, which suggests

that there was a station error, as there are no specimens in the LACM collection from comparable depths.

Remarks. This species is here reinstated and removed from the synonymy ofPlesiocystiscus politulus 1

(Dall, 1919), in which it had been placed by Roth and Coan (1968:64). Earlier, Coan and Roth (1966:291) J

had illustrated the holotype, but had been uncertain as to its validity.

Plesiocystiscus myrmecoon is characterized by its relatively slender profile and by its size, which !

ranges from 3.3-4.8 ram, intermediate between that of the larger P.jewetti (Carpenter, 1857) and the smaller

gum

P. politulus (Dall, 1919). Specimens ofP. myrmecoon in collections had previously been thought to be smaller- I

sized specimens of P.jewetti. Distributions of the three species overlap in southern California. •

J

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I Superfamily Cancellarioidea

I Family Cancellariidae Forbes and Hanley, 1851

Diagnosis. Shell small to large, spire elevated; sculpture both axial and spiral; columella with 2-4

I weak to slxong plaits; canal short, operculum lacking; protoconch paucispiral. Radula nematoglossan, of long central teeth only.

Biology. The biology of cancellariids is poorly known. Recent papers are those of Hamsewych and

I Petit (1982, 1986) and Petit and Harasewych (1986). The elongate teeth of the nematoglossan radula are unlike that of other neogastropods, accounting for the placement of this family in its own superfamily. All

cancellariids are believed to feed suctorially on internal fluids of prey animals, using the tong teeth to penetrate

i the bodies of the prey.

Cancellariids live on soft substrates from shallow to deep water. Egg capsules are attached by long

stalks. Planktotrophic veligers may be released or development may be direct (Pawlik et al., 1988).

I Remarks: Petit and Harasewych (1990) published a catalog of all generic and specific level taxa that

have been proposed in the family. Itis still premature to offer a complete classification because so few genera

i are known anatomically. Two subfamilies are known, Cancellariinae and Admetinae.

i Key to Species of Cancellariidae

1A. Columellar plaits strong ................................................................................................................... 2

I lB. Columellar plaits weak .................................................................................................................... 3

I 2A. Color pattern of narrow dark bands on lighter ground ...................................... Cancellaria cooperii

2B. Color pattern lacking ................................................................................ Cancellaria crawfordiana

I 3A. Whorl profile angulate ......................................................................................... Admete californica

3B Whorl profile rounded ............................................................................................. Admete gracilior

!

Subfamily Cancellariinae Forbes and Hanley, 1851

I Diagnosis. Shell of medium to large size, columellar plicae well developed.

Remarks. Cancellariinae are typically a more tropical group than the Admetinae.

!

Genus Cancellaria Lamarck, 1799

I Cancellaria Lamarck, 1799. Type species (M): Voluta reticulata Linnaeus, 1767. Caribbean.

Diagnosis. Shell of moderate to large size, spire of moderate height, sculpture both axial and spiral;

I col operucmella ulum with lacking. well-de Radula veloped nematoglos calussan, and ofthreelongplcentral aits, theteethuppermost only. the strongest; canal usually short;

Biology. As described for family.

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Remarks. Shell characters provide useful distinctions atthe subgeneric level. Most authors have not •

elevated them to full genera, however. The applicable subgenera for the two species treated here are mentioned

in the remarks section for each. I

u

Caneellaria eooperii Gabb, 1865 m

Figure 1.23B I

Cancellaria (Narona) cooperii Gabb, 1865:186 [unfigured]. I_

Cancellaria (Progabbia) cooped: Dall, 1921:83, pl. 5, figs. 3, 4.--Oldroyd, 1927:152, pl. 11, fig. 14.-

Abbott, 1974:247, fig. 2694.

Cancellaria cooperi: Grant and Gale, 1931:619.--McLean, 1978:51, fig. 28.1. i

n

Material Examined. Santa Maria Basin, Phase I, sta. 31,100 m (1). Other material: 51 lots in the

LACM collection. I

Description. Shell large, high-spired; shoulder slightly concave; columella with 2 strongplicae and a

basal fold; lip strongly lirate within. Ground color yellow brown, with low thin spiral cords of darker brown.

Axial sculpture of broad ribs, forming projecting nodes but not extending across the shoulder. Length 40-90 I[

m II I/L

Type Locality and Type Specimens. San Diego and Monterey, California. Type material not located I1

(Coan and Bogan, 1988:278). l

Distribution. Monterey Bay, California (36.5°N), to Punta San Pablo, Baja California Sur (27°N).

Habitat. Soft bottoms, 30-280 m. 1

Ill Remarks. The color markings of this species are characteristic andit canbe confused with no other.

O'Sullivan et al. (1987) reported that Cancellaria cooperi parasitizes the California electric ray, as

it rests on the sea bottom, but not other bottom fishes. In the absence of electric rays the snails remain buried •

in the sand. This is the only member of the Cancellariidae for which the feeding relationship is known.

Reproduction in this species was described in detail by Pawlik et al. (1988). Im

Gabb's original spelling of the name cooperii is used, because the current rules of zoological 1

nomenclature do not allow for emendation to the preferable spelling cooped.

This species is the type species of Progabbia Dall, 1918 (Petit and Harasewych, 1990:7). I

Cancellaria crawfordiana Dall, 1891

Figure 1.23C I

Cancellaria crawfordiana Dall, 1891:182, pl. 6, fig. 1. •

CancelIaria (Progabbia) crawfordiana: Grant and Gale, 1931:614.---Oldroyd, 1927:153. U

Cancellaria (Crawfordina) crawfordiana: Abbott, 1974:247, fig. 2692.

l

Cancellaria ghiorum Costa, 1993:9, figs. 1-4. I

Material Examined. California: Santa Maria Basin, Phase II, sta. R-6, 410 m (1); sta. PJ-16, 130 m 1

(1immature). Other material: 28 lots in the LACM collection. •

U

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I Description. Shell moderately large, high-spired; whorls rounded, suture impressed, upper part of

whorl slightly tabulate; columella with 3 plications. Axial sculpture of strong ribs, extending to canal; spiral

I sculpture Shell creaofm narrow colored cords, under overriding a thick, darkaxibrown al ribs periostracum. and forming nodes Lengthatintersections. 30-48 mm. Lip strongly lirate within.

Type Locality and Type Specimens. Drake's Bay, Matin California, 24 fathoms. Holotype: USNM

I 122400. Cancellaria ghiorum: Carmel, Monterey County, California, 300 m; holotype: CAS 086776. Distribution. Coos Bay, Oregon (43.5°N), to Punta Rompiente, Baja California Sur (27.5°N)

Habitat. Soft bottoms, 40-290 rn_Feeding habits are unknown. The bathymetric range exceeds that

I of Cancellaria cooperi.

Remarks. The illustrated specimens of the recently proposed Cancellaria ghiorum are well within

I the variation This of is the C. crawfordiana type species,ofandthethesubgenus name isCrawfordina here added toDall, the synonymy. 1891 (Petit and Harasewych, 1990:5).

I Subfamily Admetinae Troschel, 1866

I Diagnosis. Shell small to medium in size; columellar plaits reduced; radula lacking. Remarks. Members of this group occur in cooler or deeper water than the more tro _icalCancellariinae.

I Genus Admete Mtller, 1842

Admete Mtller, 1842. Type species (M): Admete crispa Mtller, 1842; = Tritonium viridulum Fabricius,

I 1780. North Atlantic.

Diagnosis. Shell relatively small, thin, spire high; sculpture axial and spiral; columella concave, with

I weak development Bio of plaits; operculum lacking; radula lacking. logy. Anatomy of the type species was studied by Haresewych and Petit (1986). Despite the loss

of the radula, other anatomical characters of the digestive system were comparable to those of Cancellaria

I anuknown. d Haresewych and Petit considered the admetines to feed suctorially, although the prey organisms are

Remarks. The type species Admete viridula is broadly distributed throughout the Arctic Ocean and

I the North Atlantic, as welI as the northeastern Pacific south to Oregon (in the LACM collection). It is characterized by broad spiral cords with narrower interspaces. It is highly variable in profile and numerous

synonyms have been proposed.

I There has been debate in the literature about the of the viridula for the To

use name type species.

settle the matter, Bouchet and War_n (1985:258) designated a neotype, figured specimens representing a

range of variation, and listed a number of additional synonyms. Further arguments in support of the type

I species asAdmete viridula were provided by Harasewych and Petit (1987:48). The species is often cited

type

as A. couthouyi Jay, 1839, which is another synonym.

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Admete californica (Daft, 1908) •

Figure 1.23D

CancelIaria (Admete ?) caIifornica Dall, 1908:296, pl. 4, fig. 4. II

Admete califomica: Oldroyd, 1927:159, pl. 11, fig. 6.--Abbott, 1974:248. i

i

Material Examined. California: Santa Mafia Basin, Phase I, sta. 46, 597 m (1). Other material: 42

tots in the LACM collection.

Description. Shell of medium size, whorls 4, spire high, suture deeply impressed, whorls with tabulate 1

profile defined by peripheral angulation; shoulder area slightly convex; columella with concave profile and

pronounced umbilical chink; columella plaits weak; axial and spiral sculpture producing square cancellations 1

with beads at intersections; 4 spiral cords on early whorls and 10 more across base. Shell white under olive |

brown periostracum. Length 12-16 mm.

Type Locality and Type Specimens. Off San Diego, California, 603 fathoms. Lectotype (here •

designated): USNM 110626. 1

Distribution. Off Send Kruzof Island, Alaska (57.5°N), to Islas Coronados, Baja California (32.5°N).

Dall (1921:84) reported the southern distribution to extend to the Gulf of California, but the locality information I

for the single lot [USNM 211379] is suspect and the record is discounted.

Habitat. Soft bottoms, 170-2,000 m. i

Remarks. Admete califomica differs from the type speciesA, viridula in having a pronounced umbilical i

chink, a strongly tabulate shoulder at least in early whorls, and in having evenly cancellate sculpture rather

than sculpture dominated by axial ribs. The umbilical chink is apparent on large specimens, but is not evident i

specimens under 10 mm in length. The species is variable in the strength of the axial and spiral elements. •

on

II

Admete gracilior (Carpenter, 1869) i

m

Figure 1.23E

Cancellaria gracilior Carpenter in Gabb, 1869 [1866-69]:50---Palmer, 1958:223, pl. 26, fig. 6, 7 [holotype]. i

Admete couthouyi graciIior: Oldroyd, 1927:158.

Admete woodworthi Dall, 1905:123.---Oldroyd, 1927:160 [unfigured]. i

Admete rhyssa Dall, 1919:306.--Oldroyd, 1927:156 [unfigured].

Admete seftoni Berry, 1956:155, figs. 2, 9. i

Material Examined. California: Santa Maria Basin, Phase II, sta. R-9, 410 m (1 juvenile); sta. PJ8, 150 m (1 immature). Other material: 74 lots in the LACM collection. HI

Description. Shell small, high-spired; whorls 4, suture deeply impressed, whorls evenly rounded; I

columella with concave profile, plaits weak; umbilical chink usually closed by columellar lip; axial sculpture

of 9-14 strongly projecting ribs, crossed by narrow, raised spiral cords. Shell white under olive brown m

periostracum. Height 10-12 mm. 1

Type Locality and Type Specimens. Santa Barbara, middle Pleistocene. Holotype: UCMBP 15560.

Admete woodworthi: Monterey Bay, 10-45 fathoms; holotype: USNM 181822. Admete rhyssa: off South •

Coronado Island, Baja California, 155 fathoms; holotype: USNM 211241. Admete seftoni: N of Anacapa i

Island, California, 46-58 fathoms; holotype: SBMNH 34555.

Distribution. Monterey Bay (36.5°N), to Punta San Pablo, Baja California Sur (27°N). i

118 i
I

I HabitaL Soft bottoms, 60-250 m.

Remarks. Four taxa are here grouped as a species variable in breadth and in number of axial ribs.

I The species is characterized by rounded, projecting axial ribs, narrow spiral cords anda deep suture. Carpenter's taxon, based on a specimen from the middle Pleistocene, was thought to be extinct, but is represented by Living

specimens of slender profile. Admete woodworthi was based on a slender form with a minimal number of

I axial ribs, whereas A. rhyssa and A. seftoni were based on type material of broader profile.

i Superfamily Conoidea

Until recently, the superfamily Conoidea included three main families: Turridae, Terebridae, and

Conidae. The family Turridae (which hadbeen defined on the shell character ofa subsutural sinus) as used by

I authors, has recently been subdivided result of phylogenetic analysis by Taylor, et aL (1993).

previous as a

Based on a comparative studyof foregut anatomy andcorresponding radular characters, the old family Turridae

has been subdivided into five families (here called turriform families) that were previously regarded as

I subfamilies: Drilliidae, Pseudomelotomidae, Strictispiridae, Turridae, and Conidae (Conidae now includes

some turriform subfamilies). Sysoev (in Taylor et al., 1993, Appendix 2) assigned all turriform genera to the

revised classification of Taylor et al.

I Two turriform families, the Turridae, and the redefined Conidae, are represented in the benthic fauna

of southern California. There is a major difference in radular structure and function; but, on shell characters,

they can not readily be distinguished. Genera of both families are therefore placed in the same key based on

I shell characters.

I Key to Species of Turridae and Conidae

I 1A. Anal sinus not bordered by thick callus ........................................................................................... 2

lB. Anal sinus bordered by thick callus on both sides ......................................................................... 21

I 2A. Shell nearly smooth, lacking axial and spiral sculpture ................................................................... 3

213. Shell with spiral or axial sculpture or both ...................................................................................... 4

I 3A. Shell sinistral .................................................................................................... Antiplanes cataIinae

i 3B. Shell dextral ........................................................................................................... An@lanes thalea

4A. Sculpture spiral only ........................................................................................................................ 5

I 4B. Sculpture both axial and spiral 8

I 5A. Canal nearly as long as aperture ...................................................................................................... 6 5B. Canal short, less than length of aperture .......................................................................................... 7

I 6A. Three spiral cords at periphery ............................................................................. Antiplanes briseis

6B. Single spiral cord at periphery ............................................................................. Carinoturris fortis

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I 119
I

7A. Whorls strongly inflated .................................................................................. Pseudotaranis strongi I

7B. Whorls only slightly inflated .......................................................................... Pseudotaranis hyperia

I

8A. Early whorls with axial nodes on peripheral carination ................................................................... 9

8B. Early whorls not having axial nodes on peripheral carination ....................................................... 16 I

9A. Columella lacking single carination ............................................................................................... I0

9B. Columella with single carination ..................................................................................................... 11 I

10A. Shell length over 17 mm ................................................................................. Rhodopetoma diauIax I

10B. Shell length under 16 mm ............................................................................... Rhodopetoma renaudi

11A. Axial nodes persisting on final whorl ............................................................................................. 12 I

12B. Axial nodes not persisting on final whorl ....................................................................................... 13

I

12A. Shell relatively slender, profile from whorl to whorl convex .............................. Borsonella bartschi

12B. Shell relatively broad, profile from whorl to whorl straight .............................. Borsonella merriami I

13A. Periphery angulate ......................................................................................................................... 14 HI

13B. Periphery not angulate ................................................................................................................... 15 I

i

14A. Shoulder concave ............................................................................................ Borsonella coronadoi i

I 14B. Shoulder fiat-sided ............................................................................................... Borsonella hooved

15A. Whorl profile rounded ........................................................................................ Borsonella omphale I

15B. Whorl profile constricted below suture ........................................................... Borsonella pinosensis

I

16A. Whorl profile rounded .................................................................................................................... 17

16B. Whorl profile angulate ................................................................................................................... 20 I

If

17A. Axial sculpture on early whorls, lacking on final whorl .................................... Daphnella clathrata

17B. Axial and spiral sculpture retained at all growth stages ................................................................ 18 •

18A. 18B. Mature Mature shell > 20 mm ................................................................................... Ophiodermella inermis I shell < 15 mm .................................................................................................................... 19

I

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I 19A. Profile broad, aperture length nearly half the length of shell ................... Ophiodermella cancellata

19B. Profile slender, aperture length about one-third the length of shell ........... Ophiodermellafancherae

I

20A. Outer lip thin at edge ................................................................................................... Kurtzina beta

I 20B. Outer lip thickened at edge ........................ ................................................................ Kurtzia arteaga

21A. Final whorl lacking axial fibs ........................................................................................................ 22

I 2lB. Final whorl with axial ribs ............................................................................................................. 23

I 22A. With strong peripheral carination ........................................................................ Crockerella evadne 22B. Lacking strong peripheral carination .............................................................. Crockerella philodoce

I 23A. Spiral cords of penultimate whorl broader than interspaces .......................................................... 24

23B. Spiral cords of penultimate whorl narrower than interspaces ........................................................ 25

!

24A. Interspaces deeply incised ............................................................................... Crockerella tridesmia

I 24A. Interspaces not deeply incised ........................................................................ Crockerella castianira

25A. Axial and spiral sculpture equal in strength .................................................. Crockerella crystallina

I 25B. Axial ribs stronger than spiral cords ..............................................................................................

26

I 26A. Shoulder tabulate ........................................................................................................................... 27 26B. not tabulate ..................................................................................................................... 28

I 27A. Profile slender, axial ribs about 10 per whorl .............................................. Crockerella conradiana

27B. Profile broad, axial ribs about 12 per whorl .......................................................... Crockerella scotti

I 28A. Spiral cords alternating in size ........................................................................ Crockerella cymodoce

tl 28B. Spiral cords not alternating in size ................................................................................................. 29

29A. Whorls rounded, not shouldered ........................................................................ Crockerella eriphyle

I 29B. Whorls with shoulder CrockerelIa Iowei

concave

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Family Turridae Swainson, 1840 i

Diagnosis. Shell tall-spired, aperture elongate, with anterior canal or notch; and sinuous posterior •

notch; sculpture varied. Odontophore present, radular teeth attached to membrane, usually of wishbone type

of marginals only; venom gland present.

Biology. This group of turriform gastropods uses the venom gland to paralyze prey organisms. Although i

the rachididan and lateral teeth (if present) are vestigial, and the functional teeth consist of the marginal teeth; II

the odontophore and radular ribbon are still functional. Reproduction can be either direct or with a planktotrophic

veliger stage, i Remarks. In the classification of Taylor et al. (1993), the newly redefined Turridae retains five

subfamilies (Ctavatulinae, Crassispirinae, Zonulispirinae, Cochespirinae, Turrinae) and a large number of

genera, i

Subfamily Cochlespirinae Powell, 1942 !

Diagnosis. Shell medium to large, fusiform, anterior canal moderately long, anal sinus on shoulder. i

Remarks. This group was previously known as Turriculinae, but the type genus Turricula Schumacher, 1

1817, was transferred to the Clavatulinae by Sysoev in Taylor et al. (1993:160).

I

Genus Antiplanes Dall, 1902 i

I

Antiplanes Dall, 1902. Type species (OD): Pleurotoma (Surcula)perversa Gabb, 1865, not Philippi, 1847 i

[= Pleurotoma (Antiplanes) catalinae Raymond, 1904]. Northeast Pacific. HIlll$

Rectiplanes Bartsch, 1944. Type species (OD): Pleurotoma (Antiplanes) santarosana Dall, 1902. Northeast

Pacific. I

Diagnosis. Shell large, fusiform, dextral or sinistral; axial sculpture absent; spiral sculpture of cords

or lacking; anal sinus deep, U-shaped, on shoulder; siphonat canal long. Protoconch paucispirai. Operculum

filling aperture, nucleus terminal. Radula teeth of bifurcated marginals and vestigial rachidian, i

!

Biology, The paucispiral protoeonch is indicative of direct development. The circular egg capsule

was figured by Kantor and Sysoev (1991, fig. 101).

Remarks. Kantor and Sysoev (1991) reviewed the northwestern Pacific members of this genus, i

Dextral and sinistral species are known and subgenera have been based on this distinction. However, Kantor

and Sysoev did not consider the distinction worthy of recognition. The significance of this difference is not j

understood. Antiplanes has been used as a subgenus for the sinistral species and Rectiplanes Bartsch, 1944, |

for the dextral species.

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122 i
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I Figure 1.24. Turddae. A. Antiptanes catalinae (Raymond, 1904): 229 m, off Santa Catalina Island, California

(USNM 192087, lectotype); height 25.1 ram. B. Antiplanes thalea (DaU, 1902): 461 m, off San

I Luis 1902):Obispo, 97 m, offCalifSanta orniaRosa(USNM 122568, holotype); height 40 mm. C. Antiplanes thalea (Dall, Island, California (USNM 109198, holotype ofA. santarosana); height

36 mm. D. Antiplanes briseis Dall, 1919:150-161 m, Avalon Bay, Santa Catalina Island, California

I (LACM 40-123.7); height 25.0 mm. E. Carinoturrisfortis Bartsch, 1944:227-229 m, N of Anacapa Island, California (LACM 41-52.3); height 14.3 mm.

I Antiplanes catalinae (Raymond, 1904)

i Figure 1.24A

Pleurotoma (Surcula) perversa Gabb, 1865:183 [not Philippi, 1847].

I Anu'planesperversa: Oldroyd, 1927:82, pl. 11, fig. 9, pl. 28, fig. 8. Pleurotoma (Antiplanes) catalinae Raymond, 1904:2 [unfigured].

Antiplanes major Bartsch, 1944:29 [unfigured].

I "Antiplanes voyi (Gabb, 1866)."-- of Abbott, 1974:267.

Antiplanes gabbi Kantor and Sysoev, 1991:122 [unnecessary new name for Pleurotoma perversa Gabb]

I Material Examined. Not represented in the voucher collection, but occurring offshore in southern

California. Other material examined: 232 lots in the LACM collection.

I moderatelyDescimripptires on.sed; Shespiral ll moscdulpture erately oflarge, fine sini straestral; , axialspiresculpture high, whoforlsf'me8-growth 10, evenly lines; rosinus uded,deesuture p, Ushaped, lip greatly protracted below sinus; protoconch paucispiral, of 1.5 rounded whorls. Periostracum tan;

I shell light brown, with lighter band at periphery; interior glossy brown, showing light peripheral band. Length 40-55 mm.

I 123
!

Type Locality and Type Specimens. Pleurotoma perversa: Santa Catalina Island, California, 60 •

fathoms. Syntypes: UCMP 15929. Pleurotoma catalinae: Santa Catalina Island, California, 125 fathoms;

lectotype (here designated) USNM 192087 (Figure 1.22A). Antiplanes major: off Bodega Head, Sonoma •

County, California, 62 fathoms; holotype: USNM 224347. l

Distribution. Queen Charlotte Sound, British Columbia (51.5°N), to San Diego, California (33°N).

Habitat. Soft bottoms, 90-270 m. l

m

Remarks. The earliest name for this species is preoccupied. The species therefore takes the name of

the next available synonym. Although Raymond claimed that his species Antiplanes catalinae was more m

slender thanAntiplanes perversa, separation into two species of the large number of specimens in collections •

im is not possible.

Grant and Gale (1931:553) placed all sinistral and dextral taxa named from the eastern Pacific under •

the name Antiplanes perversa (Gabb), believing that dextral and sinistral forms could occur in the same l

species, but that view is ruled out by the size discrepancies between the dextral and sinistral forms.

Abbott (t974:267) incorrectly used the taxonAntiplanes voyi (Gabb, 1866), for this species, but that []

name is not applicable, as it was based on a dextral species of stout profile that is now considered to be I

extinct.

Antiplanes vinosa (Dall, 1902) is a similar sinistral species described from the Aleutian Islands and •

occurring in the Northwest Pacific to the Okhotsk Sea, Russia (Kantor and Sysoev, 1991:137). As there is no []

sinistral species known from the Gulf of Alaska or southeast Alaska, the two species A. catalinae and A.

vinosa are allopatric and can be considered distinct, although they are certainly closely related. I

Antiplanes thalea (Dall, 1902)

Figures 1.24B,C B

Pleurotoma (Antiplanes) thalea Dall, 1902:514.--Dall, 1919:37, pl. 11, fig. 6.--Dall, 1921:72, pl. 1, fig. 8

[not fig. 9].--Dall, 1925:4, pl. 22, fig. 1.

Antiplanessantarosana Dall, 1902:515.--Dall, 1919: 36, pl. 11,fig. 3.--Dall, 1921:71, pl. 1, fig. 9 [not fig. 8].

Antiplanes (Rectiplanes) santarosana: McLean, 1971:120, fig. 39 [radula]. I

Pleurotoma (Spirotropis) smithi Arnold, 1903:216, pl. 6, fig. 13 [not Pleurotoma smithii Forbes, 1840].

Antiplanes rotula Dall, 1921:71 [new name for Pleurotoma smithi Arnold, not Forbes; incorrectly cited as

"Dall, 1919"].

Antiplanes (Rectiplanes) willetti Berry, 1953: 419, pl. 29, fig. 2.

Material Examined. California: Santa Maria Basin, Phase II, sta. R-2, 161 m (1 immature). Other

material: 94 lots in the LACM collection

Description. Shell of moderate size, dextral, spire high, whorls 8-11, evenly rounded, suture moderately I

impressed; spiral sculpture of fine striae, axial sculpture of fme growth lines; sinus deep, U-shaped, lip m

greatly protracted below sinus; protoconch paucispiral, of about 1.5 rounded whorls. Periostracum light

olive-green, frequently with a thin darker line at midwhorl; shell white to light tan under periostracum. Length l

30-40 mm.

Type Locality and Type Specimens. Off San Luis Obispo, California, 252 fathoms. Lectotype

(here designated): USNM 122568. Pleurotoma santarosana: Off Santa Rosa Island, California, 53 fathoms; 1

lectotype (here designated): USNM 109198. Pleurotoma smithi: San Pedro, Los Angeles, California,

u

Pleistocene; holotype: USNM 162566. Antiplanes willetti; Forrester Island, Alaska, 50 fathoms; holotype:

CAS 064665. I

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I Distribution. Petrel Bank, N of Semisopochnoi Island, Aleutian Islands, Alaska [LACM 86-326.3]

(52°N, 180°W), to Islas San Benito, Baja, California (28°N).

I Habitat, Soft bottoms, 130-400 m.

Remarks. One species of smooth-shelled, dextral Antiplanes seems to occur in the northeastern

Pacific. The lectotypes of Antiplanes thalea and A. santorosana represent extremes of variation: that of A.

I thalea (Figure 1.24B) is relatively broad, with fewer whorls than that of santarosana (Figure 1.24C), which

is slender with more whorls. Attempts todivide the LACM materialbetween two species have not succeeded.

This species is the type species of the subgenus Rectiplanes, based on its dextral shell, although the

I of such distinction is not clear.

significance a

I An@lanes briseis Dall, 1919

Figure 1.24D

I Anu'planes briseis Dall, 1919:35, pl. 22, fig. 1.--Oldroyd, 1927:85.--Abbott, 1974:267 [listed].

Material Examined, Not represented in the Santa Maria Basin voucher collection. Other material

I examined: 16 LACM lots.

Description. Shell tall-spired, slender, aperture length less than one-third shell length; shoulder slightly

i concave; periphery marked by single prominent spiral cord; base with numerous spiral cords; axial sculpture

lacking except for growth increments. Sinus deep, filling area above peripheral cord. Length 18-25 mm.

Type Locality and TypeSpecimens. Off Drakes Bay,Matin County, California, 30 fathoms. Holotype:

i USNM 212329. Distribution. Drakes Bay, Matin County (38°N), to San Diego, California [LACM 41-20] (33°b0.

Habitat. 100-230 m, soft bottoms.

I Rhodopetoma diaulax lacks the axial nodes of that species and has

Remarks. This resembles but

more prominent spiral cords on the base.

The type locality is in northern California, but all specimensin the LACM collection are from southern

I California, San Miguel Island to San Diego.

I Genus Carinoturris Bartsch, 1944

Carinoturris Bartsch, 1944. Type species (OD): Cryptogemma adrastia Dall, 1919. California.

I Diagnosis. Shell small (length 14-20 mm); canal long, sinus on shoulder just above periphery; spiral sculpture of single peripheral carinatiort; axial sculpture lacking. Radula of wishbone type.

Remarks, The shell and radula of the type species were figured by Powell (1966:49, pl. 6, fig. 10,

I text fig. B36). The type species occurs in much deeper water than does the following species and is therefore

not treated here.

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Carinoturrisfortis Bartsch, 1944 I

Figure 1.24E

Carinoturrisfortis Bartsch, 1944:60.--Abbott, 1974:266 [listed]. •

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material 1

examined: 4 LACM lots.

Description. Shell white under light brown periostracum; shoulder convex to nearly fiat; canal of

moderate length; spiral sculpture of single peripheral carination; axial sculpture lacking. Protoconch eroded R

on available specimens. Length 14-16 mm.

n

Type Locality and Type Specimens. Off Point Sur, Monterey County, California, 298 fathoms. •

Holotype: USNM 212323. I

Distribution. Off Eel River, Humboldt County [LACM 147411 [ (41 °N),to Anacapa Island,California

[LACM 41-52] (34°N). •

Habitat. 180-550 m, soft bottoms.

Remarks. This species is illustrated here for the first time. It differs from the type species of

Carinoturris in having a longer aperture, less concave shoulder, less vertically compressed whorls, a weaker I

projecting carination and a less deeply impressed suture. It occurs in shallower water than the type species. I

Genus Rhodopetoma Bartsch, 1944 I

Rhodopetoma Bartsch, 1944. Type species (OD): Borsonella rhodope Dall, 1919 [= Turris (AntipIanes)

diaulax Dall, 1908]. Northeastern Pacific. I

Diagnosis. Shell tall-spired, canal twisted to left; sinus on excavated shoulder; spiral sculpture of

peripheral cord low on whorl; axial sculpture of weak swellings on peripheral cord; protoconch paucispiral. •

Radula of wishbone type marginals.

Remarks. Rhodopetoma differs from AntipIanes chiefly in having axial swellings; the radula, as

illustrated by Powell (1966) and McLean (1971) is like that of Antiplanes. The tall profile and axial swellings I

resemble those of Borsonella, but there is no plication on the columella, the periostracum is thin and the

microscopic vermiculate sculpture of Borsonella is lacking.

One additional species, Rhodopetoma renaudi (Arnold, 1903) is here transferred to Rhodopetoma. I

1

Rhodopetoma diaulax (DAB, 1908) I

Figure 1.25A,B

(Antiplanes) diaulax Dall, 1908:247 [unfigured]. I

Turris

Antiplanes diaulax: Oldroyd, 1927:84.--Abbott, 1974:267 [listed].

Borsonella rhodope Dall, 1919: 39, pl. 12, f. 3. 1

Rhodopetoma rhodope: Bartsch, 1944:59.---Powell, 1966:32, pl. 2, fig. 16; text fig. B13 [radula].--McLean,

1971:119, fig. 42 [radula].--Abbott, 1974:265 [listed].

1

Material Examined, Not represented in the Santa Maria Basin voucher collection. Other material I

examined: 7 LACM lots.

I

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I Description. Shell with thick, dull-surfaced, olive-colored periostracum; canal short, twisted to left;

sinus on excavated shoulder; axial sculpture of weak peripheral swellings, becoming obsolete in final whorl;

i spiral scuTylptpuereLoca of lbitroy adandperipheral Type Speci cordmeand ns. nume San roDusiego,spiral Califorcordnia,s below 34 fathpeoms. riphery. HoloLength type: USNM 18-23 mm. 110609.

Borsonella rhodope: off Santa Rosa Island, 82 fathoms; holotype: USNM 212361.

I Distribution. Santa Rosa Island (34°b0, to Cortes, Bank, California (32.5°N).

Habitat. 200-570 m, soft bottoms.

i Remarks. The oldest name for this species (for which the holotype is here first figured) is used. Dall did not mention the axial nodes of the holotype of Rhodopetoma diaulax, but these show clearly in Figure

25A; the dimensions and other details of this specimen match the description, so it is clear that the specimen,

i which isThis labeledis the"type," first report is correctly of newidenti records fied. for the species subsequent to the original material described by

Dall, under either the names Rhodopetoma diaulax or R. rhodope.

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Figure 1.25. Turfidae. A. Rhodopetoma d/au/ax (DaU, 1908): 62 m, off Islas Los Coronodos, Baja California

I (holotype, 110609); height ram. Rhodopetoma (Dall, 1908): m,

USNM 19.0 B. diaulax 150 off

Santa Rosa Island, California (holotype of B. rhodope, USNM 212361); height 19 mm. C.

Rhodopetoma renaudi (Arnold, 1903): 159-174 m, SE of Santa Catalina Island, California (LACM

I 41-209. California 12); (LACM height 12.3 77-250.22); ram. O.height Pseudota13.1ranram. is strongi E. Pseudotaranis (Arnold, 190hyperia 3): 511-(D53a0U,m,1919): Tanner177-190 Bank,

m, off San Nicolas Island, California (LACM 40-99.6); height 15.0 mm.

I

i 127
!

Rhodopetoma renaudi (Arnold, 1903) I

ill

Figure 1.25C

DriIlia renaudi Arnold, 1903: 208, pl. 8, f. 5. !

Clathrodrillia renaudi: Oldroyd, 1927:70.

Spirotropis (Typhlomangelia) renaudi: Grant and Gale, 1931:549, pl. 26, fig. 29. I

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

examined: 23 LACM lots. [I

Description. Shell slender, light brown, small for genus; shoulder flat to slightly concave; spiral

sculpture of peripheral carination on lower third of early whorls, higher on Final whorl; axial sculpture of

oblique nodes cresting on carination, usually obsolete on final whorl. Length 11-16 ram. 1

II

Type Locality and Type Spechnens. San Pedro, Los Angeles County, California, Pleistocene.

Holotype: USNM 562537. I1

Distribution, San Miguel Island (34°N) to Cortes Bank, California (32.5°N). i

Habitat. 130-390 m, soft bottoms.

Remarks. This species, which was described originally from Pleistocene material, was reported

living at San Pedro by Dall (1921). It is here assigned to Rhodopetoma.

Rhodopetoma renaudi is characterized by its relatively small size, rrfid-whod carination and noded

sculpture on the carination, particularly on the early whorls. It differs from R. diaulax in its smaller size, •

longer canal, and finer periostracum.

II

Subfamily Crassispirinae Morrison, 1966 I

Diagnosis. Shell small to medium-sized; canal short, anal sinus on shoulder, parietal callus usually

well developed; protoconch paucispiral. Radula with marginal teeth of wishbone type or long flat teeth with II

accessory limb.

Remarks. Numerous genera occur in tropical, shallow-water habitats. Despite the lack of the parietal

callus characteristic of most genera, the following genus is placed in Crassispirinae on characters of the short 1

canal, sinus on shoulder, paucispiral protoconch and long, flat marginal teeth. See definition of subfamily I1

given by Taylor et al. (1993:159). 1

Genus Pseudotaranis McLean, 1995

Pseudotaranis McLean, 1995. Type species (OD): Mangelia (Taranis) strongi Arnold, 1903. Northeastern !

Pacific.

Diagnosis. Shell small, spire high, anterior canal short, whorls 6. Axial sculpture lacking, spiral •

sculpture of 2 cords emerging onteteoconch and3 cords onbase; anal sinusshallow, atperiphery andcoinciding

with uppermost cord; lip not projecting. Protoconch paucispiral, of 1.2 low, rounded whorls. Radula of

marginal teeth attached to membrane; teeth of long, flat type. •

Remarks. Authors prior to 1971 used Taranis Jeffreys, 1870, for the type species, but the presence of

a radula precludes placement in that genus (Powell, 1966:55). Earlier (McLean, 1971) I illustrated the radula

and assigned it toAntiplanes (Rectisulcus) [now in Cochlespirinae]. Pseudotaranis differs in having the sinus II

more shallow, the lip less protracted, the canal shorter, and the protoconch lower and more compressed. The

radula lacks the vestigial rachidian tooth and the marginals are not of the bifurcated type indicated forAntiplanes

by Kantor and Sysoev (1991:122). I
!

I In addition to the type species, there is one other species in this genus: Pseudotaranis hyperia (Datl,

1919), as treated here.

I sharp contrast Some specimens to the moreofdarkly both species pigmentedmayperiostracum have the periostracum of the spireofand the base upperwith portion a lightof pigmentation, the body whorl.in

I Pseudotaranis strongi (Arnold, 1903)

Figure 1.25D

Mangelia (Taranis)strongi Arnold, 1903:215, pl. 9, fig. 7 [Pleistocene].

Taranis strongi: Dall, 1921:83.--Oldroyd, 1927:151.---Grant and Gale, 1931:572, pl. 26, fig. 37.

I Antiplanes (Rectisulcus) strongi: McLean, 1971:120, fig. 40 [radula].

Pseudotaranis strongi: McLean, 1995:81.

I Borsonia incuIta Moody, 1916:54, pl. 1, figs. 2a, 2b [Upper Pliocene]. Taranis incultus: Grant and Gale, 1931:572.

I 45 lots inMaterial the LACMExamined collection. . Not represented in the Santa Mafia voucher collection. Other material examined:

Description. Shell small, spire elevated; anterior canal short; whorls 6, shoulder concave; spiral

I sculpture of 2 projecting cords whorl, 2 defining base and 3 fine cords on canal; axial sculpture of fine

per

growth lines; sinus shallow, centered on uppermost cord; lip not greatly produced. Protoconch paucispiral of

1.2 low, rounded whorls. Periostracum dark brown, often light colored on base; shell cream colored. Radula

I as described for genus. Length 12-15 mm.

Type Locality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene.

Holotype: USNM 162549. Borsonia inculta: Upper Pliocene, Fourth and Broadway, Los Angeles; holotype:

I UCMP: 11091.

Distribution. Biorka Island, Alaska [LACM 87-361.2] (56.0°N), to Islas San Benito, Baja California

i [LACM H41-5 abi1tat. ] (28°N). Soft bottoms, 100-400 m.

Remarks. This species is characterized by its strongly projecting spiral cords. Most specimens have

I the apex deeply eroded. The shell varies in breadth and strength of the spiral cords.

i Pseudotaranis hyperia (Dall, 1919) Figure 1.25E

Antiplanes hyperia Dall, 1919:35, pl. 9, fig. 6.---Oldroyd, 1927:87, pl. 3, fig. 2.

Pseudotaranis hyperia: McLean, 1995:81.

I lots in theMat LACM erial collection. Examined. California: Santa Maria Basin, Phase I, sta. 53, 196 m (1). Other material: 14

Description. Shell small, spire elevated; anterior canal short, whorls 5; spiral sculpture of 2 low

I cords whorl, 2 base and about 5 low cords canal; axial sculpture of f'me growth lines; sinus

per defining on

shallow, centered on uppermost cord; lip not greatly produced. Protoconch eroded in material on hand.

Periostracum olive green, often light colored on base; shell cream colored. Length 12-17 mm.

I

i 129
I

Type Locality and Type Specimens. Drakes Bay, Marin County, California, 30 fathoms. Holotype: I

USNM 212330.

Distribution. Fort Bragg, Mendocino County [LACM 71-138] (39.5°N), to Cortes Bank, California _l

[LACM 41-123] (32.5°N).

Habitat. Soft bottoms, 130-270 m.

Him

Remarks. This species has not previously been discussed; references to the name have been checklist 1

entries that repeated Dall (1921).

I

Pseudotaranis hyperia is more slender and with spiral cords more subdued than P. strongi. The fact 1

that it has a more restricted distribution than that ofP. strongi provides an argument in support of separation 1

at the species level. The two species are also separated bathymetrically; P. incultus occurs in lesser depths

where distributions of the two species overlap. I

inn

Family Conidae Fleming, 1822 II

I

Diagnosis. Shell coniform or turriform. Odontophore lacking, radular membrane lacking; radula of

hollow marginal teeth only; venom gland present. •

Biology. The coniform and turriform snails grouped here are capable of injecting venom into their 1

prey through hollow teeth that are used one at a time like hypodermic needles. There are exceptions, as some

have lost the radula [for example the subfamily Taraninae]. 1

Remarks. Taylor eta/. (1993:160) placed all turriform groups having the hollow marginal teeth ("the

higher turrids" of previous authors) into a family that also includes the Conidae of previous classifications.

Subfamilies recognized in the newly def'med Conidae are Clathurellinae, Conorbinae, Coninae, Oenopotinae, I

Mangeliinae, Daphnellinae and Taraniinae. []

Subfamily Clathurellinae H. and A. Adams, 1858 I

Diagnosis. Shell turriform, sculpture primarily spiral. Radula with awl or harpoon-shaped marginal _l

teeth, the tooth cavity opening terminally at the proximal end. II

Remarks. Taylor et aL (1993:160) included in this loosely defined subfamily three turriform groups

treated as subfamilies by McLean (1971): Borsoniinae, Mitrolumninae, and Clathurellinae, instead recognizing I

less formal groups: the bathytomid, borsoniid, clathurellid, mitromorphid, and tomopteurid groups. A more

[]

meaningful classification awaits knowledge of anatomy of more genera.

Genus Borsonella Dall, 1908 I

Borsonella Dall, 1908. Type species (OD): Pleurotoma (Borsonia ) dalli Arnold, 1903; not Pleurotoma dalli []

Verrill and Smith, 1882; = Borsonella omphale Dall, 1919. []

Diagnosis. Shell size small to medium (length 12-30 mm), sinus on shoulder, columella with broad •

shield and strong to weak plication; sculpture spiral, with axial ribs or nodes; shell surface with adherent 1

glossy brown periostracum; minute sculpture of vermicular markings in oblique rows. Protoconch paucispiral.

Operculum lacking. Radular teeth of slender, simple marginals with expanded bases. •

1

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130 I
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I Remarks. Borsonella species may be distinguished from other turriform genera by the minute, oblique,

vermicular pattern of sculpture, which has been mentioned in most species-level descriptions, but until now

i ofhasthenotcolumellar been emphasized plait mayas depend a genericonchar theaviewing cter. Theangle; broad ifcoltheumellar shell isshield rotatedis atolsolook characteristic. deeper within, The the strengplait th

will appear stronger. Radular teeth of BorsonelIa species were figured by Powell (1966:60, text figs. E114-

117).

I The type species of Borsonella was misidentified by Powell (1966:60, pl. 8, fig. 14), an error that

traces back to Dall (1908:25 8, pl. 13, fig. 9), who misidentified Arnold's (1903) species B. dalli. Moreover,

I match as indicated for theabove, specimen Arnold's ofB. name omphaleis preoccupied, Dall, as illustrated but thehere. actual Dali's typeillustration ofB. dalli of(USNM B. dalli162534) is here identi is a good fied

as B. coronadoi (Dall, 1908).

I and neglected The species in recent of Borsonella literature, with axe wonly ell represented one figured inbysouthern Abbott California (1974). ThebutLACM have been collection poorlycontains known

numerous lots, which has enabled the following review of the six species occurring in depths between 100 and

I 400 following m in southern were firstCalifor namednia.fromTherePleistocene are othermaterial species from by Arnold greater(1903) depthsandthatagain axenot named treatedfromhere. livingFourmaterial of the

by Dall, who did not realize the extent to which the Pleistocene species described by Arnold would likely be

I exhibits still living. a range Grantofandvariation Gale (1931) that hasplaced made many speciesnames resolutions in synonymy difficult.but Species did notareillustrate arrangedthem. here Each in decreasing species

order of abundance in the LACM collection.

i Borsonella bartsehi (Arnold, 1903)

I Figure 1.26A

Pleurotoma (Borsonia) bartschi Arnold, 1903: 200, pl. 9, f. 1

l Borsonia bartschi: Grant and Gale, 1931:545, pl. 26, figs. 27, 28.

BorsoneUa bartschi: Oldroyd, 1927:93.--Abbott, 1974:268 [listed].

Borso Borsonella neiIa civ nicoli itellaDall, Dall,1919:39, 1919:30pl. 2 [un 12,fifig. gured].---Oldroyd, 1.---Oldroyd, 1927:95.--Abbott, 1927:94.--Abbott, 1974:268 1974:268[listed]. [listed].

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other rnaterial

i examined: 45 LACM lots.

Description. Shell slender, shoulder concave, axial sculpture of about 10 elongate, oblique nodes,

I pfromrojecting whorlatoperiphery; whorl; columellar oblique vermif plait strong orm stoculmoderately pture apparedeveloped. nt under magnifi Lengthcati18-23 on; mm. profile of spire convex

Type l_aw.ality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene.

i Holotype: USNM 162535. Borsonella civitella: off Point Loma, San Diego County, California, 71-75 fathoms; lectotype (here designated): USNM 209034. Borsonella nicoli: SW of San Nicolas Island, California, 158

fathoms; holotype: USNM 198925.

Distribution. San Miguel Island (34°N) to San Diego, California (32.5°b0.

Habitat. 140-260 m, soft bottoms.

I profile with Remarks. Borsonella bartschi is recognized by its persistent axial nodes and moderately slender the spire profile convex from whorl-to-whorl. Nojustification for the proposal of the synonyms B.

civitella and B. nicoli was given by Dall; holotypes of both are well within the variation exhibited by this

i species.

i 131
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Figure 1.26. Conidae. A. Borsonellabartschi(Amold, 1903): 214-234 m, SEofSanta Catalina Island, California I

(LACM 40-125.11); height 21.0 mm. B. Borsonella coronadoi (Dall, 1908): 229-240 m, Cortes

Bank, California (LACM 41-114.3); height 22.0 mm. C. Borsonella omphale Dall, 1919:274-311 !

m, off Pyramid Cove, San Clemente Island, California (LACM 39-126.3); height 19.8 mm. D. I

Borsonella merriami (Arnold, 1903): 104 m, S of Santa Rosa Island, California (LACM 41-172,12);

height 18.1 mm. E. Borsonella hooveri (Arnold, 1903): 511-530 m, NW of Tanner Bank, California I/

(LACM 77-250.24); height 21.1 ram. F. BorsoneUa pinosensis Bartsch, 1944:119-165 m, offEmerald !

Bay, Santa Catalina Island, California (LACM 39-5.7); height 24.8 ram.

i

Borsonella coronadoi (Dall, 1908) I

Figure 1.26B I

Borsonia (Borsonella) coronadoi Dall, 1908: 277, pl. 14, f. 2.

/

Borsonella coronadoi: Oldroyd, 1927:91.--Abbott, 1974:268 [listed]. 1

Borsonella nychia Dall, 1919: 38, pl. 12, f. 2.----Oldroyd, 1927:94.--Abbott, 1974:268 [listed].

Borsonia nychia: Grant and Gale, 1931:546. j

I

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material

examined: 42 LACM lots II

Description. Shell slender to moderately broad, shoulder concave; axial sculpture of faint nodes on I

peripheral cord of early whorls, obsolete on final whorl; peripheral cord on spire strongly projecting on lower

third of whorl, but higher on penultimate and final whorl. Oblique vermiform striae well marked. Collumellar •

plait well developed. Length 17-22 mm. il

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I Type Locality and Type Specimens. Islas Coronados, Baja California, 34 fathoms; holotype: USNM

110608. Borsonella nychia: Off Point Loma, San Diego County, California, 101 fathoms; lectotype (here

I designated) D : USNM 208891. istribution. San Miguel Island (34°N), California, to Islas Coronados, Baja California (32.5°N).

Habitat. 90-300 m, soft bottoms.

i Remarks. This is another abundant member of the genus in southern California. It is characterized

by a concave shoulder, weak nodes on early whorls, a strong peripheral carination and a strong columellar

plication. Broad and slender forms are known; these forms would be considered separate species were it not

i the factthat most specimens are of moderate The was (Borsonella)

for width. slender form described asBorsonia

coronadoi. Dall (1908, pl. 16, fig. 9) illustrated the relatively broad typical form of this species, incorrectly

identified as Borsonia (Borsonella) dalli Arnold, 1903, a preoccupied name that is here synonymized with

I Borsonella omphale.

I BorsoneUa ompha/e DaIl, 1919 Figure 1.26C

I Pleurotoma (Borsonia) dalli Arnold, 1903: 201, pl. 6, f. 2 [not Pleurotoma dalli Verrill and Smith, 1882].

Borsonia (Borsonella) daUi: Dall, 1908:275 [not pl. 13, fig. 9].

Borsonella dalli: Oldroyd, 1927:91 [not. pl. 6, fig. 8, copy DaU, 1908]. Borsonella omphaIe Dall, 1919: 38, pl. 12, f. 4.--Oldroyd, 1927:90, pl. 19, fig. 6.--Abbott, 1974:268.

Spirotropis (Borsonella) omphale: Grant and Gale, 1931:552.

I Borsonella angelena Hanna, 1924:158 [new name for Pleurotoma dalIi Arnold, 1903].--Abbott, 1974:268.

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material

i examined: 10 LACM lots.

Description. Shell of medium profile, whorls rounded, shoulder area flat or only weakly concave;

I ofspiral lowscul projections pture of weak on peripheral peripheralangulation, angulation, bestlowdeveloped on whorl onandearly bestwhorls. developedOblique in earlyvermiform whorls; axial striaesculpture present.

Columellar plait weakly developed, visible if shell is rotated. Length 16-21 mm.

i USNM 162534. Type LocBorsonella ality and TyomphaIe pe Sp:eciOffPoint mens. Borsonella Loma, Sandalli Diego : SanCounty, Pedro,California, California, 78Pleistocene; fathoms. Holotype: holotype:

USNM 209174.

I (33°N). Distribution. DrakesBay, Matin County [USNM 212324] (38°N) to San Clemente Island, California

Habitat. 100-200 m, soft bottoms.

Remarks. This species is characterized by its lack of a strongly concave shoulder, its tall spire,

slender profile and minimal spiral and axial sculpture, compared to other species of Borsonella.

I

replacement As noname, ted above, but thatArnold's was notori needed, ginal nbecause ame for this Dall'sspecies (1919)isname preocBorsonella cupied. Hannomphale a (1924)is aprovi synonym ded a

that takes precedence.

I species, This the concept species ofis the the type genusspedoes ciesnot of Bochange. rsonella. Although previous authors have misidentified the type

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I 133
!

Borsonella merriami (Arnold, 1903) I

Figure 1.26D

Ii

DrilIia merriami Arnold, 1903: 207, pl. 8, f. 7. _l

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material /

examined: 8 LACM lots. I!

Description. Shell relatively broad; shoulder markedly concave; axial sculpture of about 10 vertically illl

oriented nodes projecting at periphery andpersisting at all growth stages. Profile from whorl to whorl straight. 1

1

Columellar plait weakly developed, visible if shell is rotated. Length 16-27 ram.

Type Locality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene. /

Holotype: USNM 162536 [not USNM 561571, which also is labeled "type"]. il

Distribution. Santa Rosa Island (34°N) to San Clemente Island, California (32.5°N).

Habitat. 70-290 m, soft bottoms, i

Ill

Remarks. This is another species described by Arnold (1903) from the Pleistocene of San Pedro that

is here reported to occur in the living fauna. .Ib

This species is characterized by its broad profile that is straight from whorl to whorl, prominent axial I

nodes that may be elongate without being oblique, and weak development of the columellar plait in mature

IIIII

shells. I

BorsoneIla merriami differs from B. bartschi in its broader profile. The spire prof'fle from whorl to I

whorl is straight rather than convex. Although mature specimens have a poorly developed plait, the plait is

evident on immature specimens. 1

Borsonella hooveri (Arnold, 1903)

Figure 1.26E I

Pleurotoma (Borsonia) hooveri Arnold, 1903: 201, pl. 10, f. 1

II Spirotropis (Borsonella) hooveri: Grant and Gale, 1931:550.

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

examined: 5 LACM lots. |

Description. Shell profile broad, aperture large, nearlyhalf length of shell; shoulder areastraight to

convex; spiral sculpture of peripheral angulation on lower third of whorl, becoming faint in mature specimens;

axial sculpture of weak nodes on peripheral angulation of early whorls. Oblique vermiform sculpturepresent. I1

Columellar plait of moderate strength, visible if shell is rotated. Length 15-21 mm.

Type Locality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene. I

Holotype: USNM 162533. II

Distribution. Santa Catalina Island (33.5°N) to Cortes Bank, California (32.5°N). I

Habitat. 150-500 m, soft bottoms. I

Remarks. This species was described from Pleistocene material but is here recognized in the living

fauna. It is a rare species characteristic of depths greater than those of the other species treated here. I

II

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I Borsonella hooveri is characterized by its relatively long aperture (nearly half the length of the shell),

inflated whorls and lack of a concave shoulder area; mature specimens may have a peripheral carination, but

I the shouThe lder Pleistocene area is flat and the carination is higher on the whorl than that ofB. coronadoi. type specimen of BorsonelIa hooved is not fully grown; it measures 12.8 mm in

height.

I Borsonella pinosensis Bartsch, 1944

I Figure 1.26F

Borsonellapinosensis Bartsch, 1944:63 [unfigured].--Abbott, 1974:268 [listed].

I Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

examined: 5 LACM lots.

centeredDesc onupper ription. thirdShe of wllhorl; slendperiphery er, tall-spired rounded , whor ; axial ls wesculpture akly roof unded, vrticalshonodes ulder onwithearlynarro whorwls, const becoming riction

obsolete on final whorl; sinus on shoulder, moderately deep. Oblique vermiform striae present. Columellar

i plait weakly developed, visible if shell is rotated. Length 21-27 ram. Type Locality and Type Specimens. Off Point Pifios, Monterey County, California, 49-50 fathoms.

Holotype: USNM 210634.

I Distribution. Point Pifios, Monterey County (36.5°N), to Santa Catalina Island, California (33.5 °N).

Habitat. 90-300 m, soft bottoms.

locality. ReIt mmost arksclosely . This sresembles pecies is fiBorsonella gured for theomphale first time , fromandwhich the distributi it differson inis itsextegreater nded frolength, m themore type

slender proffde and having the constriction on the uppermost third of the whorl. This species as well as B.

omphal Californeia.are the only members of Borsonella known to occur off central California as well as southern

i Genus Ophiodermella Bartsch, 1944

Ophiodermella Bartsch, 1944. Type species (OD): Pleurotoma inermis Reeve, 1843. Northeastern Pacific.

I Diagnosis. Shell fusiform, whorls rounded, suture moderately impressed; axial and spiral sculpture

equally developed, producing square cancellations with beads at intersections; sinus on shoulder, moderately

deep, U-shaped, its position marked by sinuous axial fibs; lip protracted below sinus; canal long; protoconch

I paucispiral, of 1.5 separated and rounded whorls.

Biology. Shimek (1983) studied the biology of Ophiodermella inermis and O. cancellata. Both

i species feed exclusively on different species of owenid polychaetes. The paucispiral protoconch is indicative of direct development, although Shimek (1983:306) reported that the late veliger stage is mobile and feeds

actively for up to five weeks.

I Remarks. Three species are known in the northeastern Pacific and each is treated here.

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i

Figure 1.27. Conidae. A. Ophiodermelta inerrnis (Reeve, 1843): 18 m, off San Diego, California (USNM 110644,

holotype of halcyonis); height 23 ram. B. Ophiodermella cancetlata (Carpenter, 1864): 198-199 m, iBm

off Long Point, Santa Catalina Island, California (LACM 41-164.21); height 8.5 mm. C. il

Ophiodermellafancherae (Dall, 1903): 86-95 m, NW of Anacapa Island, California (LACM 41-

47.17); height 10.0 mm. D, Kurtzina beta (Dall, 1919): 102 m, off Point Afio Nuevo, San Mateo 'll

County, California (USNM 206554, holotype); height 5.0 mm. E. Kurtzia arteaga (DaU and Bartsch, i

1910): 22-27 m, off Point Loma, San Diego County, California (USNM 150993, holotype of var.

roperi); height 6.5 ram. ,,.

I

11

Ophiodermella inermis (Reeve, 1843) I

Figure 1.27A

I

Pleurotoma inermis Reeve, 1843 [1843-44], pl. 8, fig. 64.--Keen, 1966:271, pl. 47, fig. 9 [holotype].

Ophiodermella inermis: Shimek, 1983:281, fig. 1A. i

Drillia incisa Carpenter, 1864:657.--Palmer, 1958:226. I

Moniliopsis incisa: Dall, 1919:28, pl. 12, fig. 7.

Surcula ophioderma Dall, 1908:247 [unnecessary new name for Pleurotoma inermis Reeve]. B

Ill

Moniliopsis incisa var. ophioderma: Grant and Gale, 1931:566.

Ophiodermella ophioderma: McLean, 1978:54, fig. 30.2 i

Turris (Surcula) halcyonis Dall, 1908: 248.--Da11, 1919:28, pl. 8, fig. 1.

Ophiodermella montereyensis Bartsch, 1944:62 [unfigured]. l

Material Examined. Not represented in the voucher collection. Other material: 197 lots in the LACM R

collection.

i

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I Description. Shell large,whorls 8, nearly flat-sided on first 4 whorls, weakly rounded on later whorls;

axial and spiral sculpture strong on immature specimens, becoming faint on mature specimens; protoconch of

Length 1.5 separated and rounded whorls; shell light brown with axial flamules of brown under brown periostracum. 25-40 mm.

Type Locality and Type Specimens, Bahfa Magdalena, Baja California Sur, Mexico. Holotype:

I N12630. HML 79Turris .2.26.halcyonis 98 DriIIia: Coronado incisa: NeahBeach, Bay, Clallam San DiegoCounty, County, Washington; California,lectotype 10 fathoms; (here selected): holotype USNM USNM

110644. Ophiodermella montereyensis: Monterey Bay, California, 13 fathoms; holotype: USNM 214251.

I Distribution, Skidegate, Queen Charlotte Islands, British Columbia ILACM 73-78] (53°N) to Rancho Inocentes, Baja California Sur [LACM 71-19] (24.5°N).

Habitat. Soft bottoms, intertidal to 70 rm

I Remarks. Although usually credited to Hinds, Keen (1966:271) showed that the name of the type

species inermis was first published by Reeve. Dall's replacement nameSurcula ophioderma was unnecessary

because Some "Drilliaauthors inermhave is Partsch, 1843" was not a published name, as shown by Cemohorsky (1975:318). treated the northern form named incisa as a species distinct from the more southern

Ophiodermella inermis, but there is no justification for doing so, even at the subspecifie level.

I Ophiodermella The name inermis Turris are halcy brownonis , thereby Dall,differing is basedfromon more O. cancellata slender andoffshore O.fancherae forms.. Small Specimens specimens occurringof

in the intertidal zone tend to be large and may have worn sculpture, whereas those from deeper water are more

i slender and more crisply sculptured.

Ophiodermella cancellata (Carpenter, 1864)

I Figure 1.27B

Drillia canceUata Carpenter, 1864:658 [unfigured]. Ophiodermella cancelIata. Shimek, 1983:281, fig. 2C.

i PIeurot Surculaomarhinesvancouvere Dall, 1908:248 nsis E. A. [unnecessary Smith, 1880:286 new name [unfigure for Drillia d]. cancellata].

Moniliopsis rhines: Dall, 1919:28, pl. 8, fig. 5

i Moniliopsis chacei Berry, 1941:6, pl. 1, f. 2.

Material Examined. Not represented in the voucher collection from Santa Mafia Basin. Other material:

,_m 30 lots in the LACM collection. Description. Shell small, whorls 6, rounded, axial and spiral sculpture producing coarse clathrate

pattern; protoconch of 1.5 separated and rounded whorls; shell white under thin, nearly transparent periostracum.

i Length 11-12 ram.

Type Locality and Type Specimens. Puget Sound, Washington. Type material not located by Palmer

(1958:226). Pleurotoma vancouverensis: Vancouver Island, British Columbia; syntypes: NHML: 1880.6.23.9-

I Moniliopsis Pleistocene, Marl, Hilltop Quarry, Pedro, California; holotype:

10. chacei: Middle Lomita San

SBMNH 34515.

Distribution. Kachemak Bay, Alaska [LACM 57-11.44] (59.5°N), to Tanner Bank, California [LACM

I 76-313] (32.5°N).

Habitat. Sandy bottoms, 50-500 m. Specimens from southern California are recorded from depths

I below 200 m.

i 137
!

Remarks. This species is characterize by its small size,inflated whorls and coarse cancellate sculpture. I

Palmer (1958:225) showed that Dali's replacement name Surcula rhines was unnecessary, although his

illustration (Dill, 1919) was a good representation of a typical specimen, i

OphiodermeUafancherae (Dill, 1903)

Figure 1.27C I

Mangilia fancherae Dill, 1903:172. I

Moniliopsisfancherae I : Dill, 1919:28, pl. 8, fig. 3

Moniliopsis incisa var.fancherae: Grant and Gale, 1931:567. in

Ophiodermella fancherae: Skoglund, 1994:127, fig. 1. I

Material Examined. Santa Maria Basin, Phase II, sta. R-4, 92 m (2). Other material: 68 lots in the

LACM collection. !

Description. Shell small, whorls 6, nearly flat-sided; early whorls with peripheral carination; spiral

cord of shoulder weak; mature sculpture finely clathrate, protoconch of 1.5 separated and rounded whorls;

shell white

to tan under light brown periostracum. Length 8-17 mm. l[

Type Locality and Type Specimens. Off Avalon, Santa Catalina Island, 50 fathoms. Lectotype

(here selected): USNM 109303. [l_

Distribution. Off Point Pifios, Monterey California [LACM 59-98] (36.5°N), to Bahia San Bartolom6,

Baja California Sur ULACM40-2] (27°N); also known in the Gulf of California (Skoglund, 1994:127).

Habitat. Soft bottoms, 50-200 m. i

Remarks. This species is also small but much more slender than Ophiodermella cancellata. Confusion

in the literature is due to Dali's later figure said to be Moniliopsisfancherae (1921, pl. 6, fig. 3), which does Ill

not agree with the holotype and is evidently based on a specimen ofO. canceIlata (Carpenter, 1864). Specimens I

ofO. fancherae can be separated from immature examples of the more slender offshore form ofO. inermis in

being more slender and lighter in color than O. inermis. The specimen from the Gulf of California figured by

Skoglund (1994, fig. 1) is the largest known specimen at 17.7 mm in length. I

Subfamily Mangeliinae Fischer, 1884

III

Diagnosis. Shell small, fusiform, anterior canal short; axial and spiral sculpture well developed; anal

sinus on shoulder; outer lip usually swollen, sometimes denticulate, operculum lacking. Marginal teeth of m

radula hollow, with solid base, some with semi-enrolled shaft; tooth canal open laterally. I

Remarks. Most have elaborate multispiral protoconchs, indicative of planktotmphic veligers. Generic

descriptions emphasize differing kinds of protoconch sculpture. I

Genus Kurtzina Bartsch, 1944 Ill

Kurtzina Bartsch, 1944. Type species (OD): Mangelia (Kurtziella) beta Dall, 1919. California. I

Diagnosis. Shell small, with spiral and axial sculpture and midwhorl carination; sinus shallow, not II

bordered with callus; lip not thickened. Protoconch relatively large, whorls 3, tip immersed, second protoconch I

whorl smooth, third with cancellate sculpture with beaded intersections.

Remarks. There are two species, the type species and Kurtzina cyrene (Dill, 1919) in the Panamic Ill

province (see McLean in Keen, 1971:749). !1

138 I
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I Kurtzina beta (Dall, 1919)

Figure 1.27D

i Mangilia (Kurtziella) beta Dall, 1919: 64, pl. 22, fig. 4

i Mangilia KurtzieIla beta (Kurtzina : Oldroyd, ) beta1927:135. : McLean in Keen, 1971, fig. 1792.

I Ill Material Examined. California: Santa Maria Basin, Phase I, sta. 22, 99 m (1); sta. 52, 98 m (1);

! [] Phase II, sta. R-8, 90 m (12). Other material: 127 lots in the LACM collection.

Description. Whorls 4, spiral cords even, with nearly equal interspaces; spiral cords numerous on

I shoulder, microscopically beaded, producing an overall frosted effect Length: 5 mm. Type Locality and Type Specimens. Off Point #,rio Nuevo, San Mateo County, California, 56

fathoms. Lectotype (here selected): USNM 206554.

i Distribution. Point/kilo Nuevo, San Mateo California to Isla La Plata, Ecuador

County, (37°N),

[LACM 34-317] (l°S).

i Habita Remarks. t. SoTfthisbospttecies oms, 2is0common -150 m. throughout its range.

i Genus Kurtzia Bartsch, 1944

] Kurtzia Bartsch, 1944. Type species (OD): Mangelia arteaga Dall and Bartsch, 1910. California.

i

I Diagnosis, Shell small, with axial and spiral sculpture and midwhorl carination; sinus well marked,

lip thickened. Protoconch of three whorls, tip immersed, second whorl smooth, third with cancellate sculpture;

axial fibs of protoconch more numerous than those of teleoconch.

I Remarks. This genus resembles Kurtzina, except that the lip is thickened and the anal sinus is also

well defined. The multiwhorled protoconch is indicative of development with a planktotrophic veliger stage.

I Kurtzia arteaga (Dall and Bartseh, 1910)

t Figure 1.27E

"'Mangelia sculpturata (Dall, 1887)."---of Arnold, 1903: 214, pl. 6, fig. 17.

I Mangelia arteaga Dall and Bartsch, 1910: 11, pl. 2, fig. 4.--Oldroyd, 1927:140. Kurtzia arteaga: McLean in Keen, 1971:754, fig. 1810.

Mangelia (Kurtziella) arteaga roped Dall, 1919:64, pl. 22, fig. 5.----Oldroyd, 1927:140. I

I Kurtzia gordoni Bartsch, 1944:65 [unfigured].

Material Examined. California: Santa Mafia Basin, Phase I, sta. 22, 99 m (1); sta. 73, 98 m (1);

I Phase U, sta. R-4, 92 m (3); sta. PJ-13, 144 m (1). Other material: 174 lots in the LACM collection.

Description. Shell small, elongate with long anterior canal. Aperture length less than half length of

I swhorls, hell, ouprotoconch ter lip thicken1Ied,withposterior 2 spiralnotthreads ch broad, andshallow. curvedSaxial uture dfibs. eeplyMature impres

ssed. culpture Protocoofnstrong ch of 1.5peripheral smooth

carination and less prominent spiral cord; base with numerous spiral cords; strong axial ribs extend from

i suture to suture and across base; fine axial and spiral striae produce a frosted surface throughout teleoconch. Length 7-10 mm.

I 139
Type Locality and Type Specimens. Barkley Sound, Vancouver Island, British Columbia, 8-34 i

fathoms. Lectotype (here sdected): USNM 211605. Mangelia arteaga roperi: Off Pt. Loma, San Diego

County, California; 12-15 fathoms; holotype: USNM 150993. Kurtzia gordoni: Off Santa Cruz, California, it

43-46 fathoms; holotype: USNM 331115. |

Distribution. Vancouver Island, British Columbia (49°N), throughout the Gulf of California and

south to the Gulf of Tehuantepec, Mexico (15°N). I

!1 Habitat. Common on soft bottoms, 20-90 m. Nothing is known about the feeding biology of this

species.

Remarks. This is the type species of the genus. None of the synonyms are worthy of recognition as i

geographic subspecies and none have come into general usage.

lira

Genus Crockerella Hertlein and Strong, 1951 I

CrockereUa Hertlein and Strong, 1951. Type species (OD): ClathureUa crystallina Gabb, 1865. California. il

Diagnosis. Shell small (7-13 mm), white under thin periostracum; microscopic sculpture throughout i

of extremely minute pustules; anterior and posterior canals deeply notched and bordered with callus; outer lip

thickened but smooth within; sculpture both axial and spiral; protoconch paucispiml, projecting and elongate, j

with deep suture, of 1.5-2 whorls, smooth except last half whorl with a faint carination. Radula of tong, II

slender marginals without barbs, tooth bases expanded.

Biology. The paucispiral protoconch is indicative of direct development or at least a very short !

planktonic stage, unlike such related genera as CIathureUa and Glyphostoma, which have multispiral !

protoconchs indicative of planktotrophic development. The radula was illustrated by Powell (1966:109, fig.

E143). i

Remarks. This genus may be distinguished from all others treated here by its deep anal sinus with

raised borders. The genus is speciose in the offshore waters of southern California, but like Borsonella has i

been neglected andpoorly understood. Most of the species described by Dall were figured but the illustrations B

were not copied by Oldroyd (1927) and Abbott (1974). To my knowledge, the extremely microscopic pustular

II

sculpture is noticed here for the first time. m

As is the case with Borsonella, the type species (ClathureIIa crystallina Gabb) of Crockerella has H

been misidentified, owing to the fact that Gabb's holotype has never been figured. The specimen illustrated by

Dall (1919, 1921), and by Abbott (1974), is here identified as Crockerella lowei (Dall), a name that DaU had IBm

dismissed as a synonym of ClathureUa crystallina. The genus Crockerella has previously been used in the 11

literature only for the species crystalIina. Despite this misuse of the name, the concept of the genus does not

change. Nine species are treated here; all but one of the names are allocated to Crockerella for the first time. in

Species are arranged here in decreasing order of abundance in the LACM collection. The diagnoses J

do not repeat the generic characters of the large siphonal notch and anal sinus, and the microscopic pustular

sculpture, i

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I

I

I

I

i

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i

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I Figure 1.2.8, Conidae. A. Crockerella lowei (Dall, 1903): 95 m, S of East Point, Santa Rosa Island, California

(LACM 41-326.5); height 9.9 mm. B. Crockerella lowei (DaI1, 1903): 96 m, off San Miguel Island,

l California (LACM 75-412.2); height 9.2 mm. C. Crockerella crystallina (Gabb, 1865): 101 m, E

of Santa Rosa Island, California (LACM 41-327.6); height 8.6 mm. D. Crockerella eriphyle (Dall,

1919): 247-256 m, SW of Balboa, Orange County, California (LACM 4t-6.3); height 11.8 mm. E.

Crockerella tridesmia (Berry, 1941): 196 m, E of Long Point, Santa Catalina Island, California

I (LACM 41-146.6); height 10.2 ram. F. CrockerelIa evadne (Dall, I919): 91 offRedondo Beach,

m,

Los Angeles County, California (LACM 17351); height 9.0 ram.

I

i CrockereFigur Ua elowei s 1.28A0D,Ball, 1903)

I Clathurella lowei Dall, 1903:172 [unfigured].

"'Philbertia crystallina (Gabb, 1865). "---of Dall, 1919:62, pl. 13, fig. 1.---Dall, 1921:79, pl. 6, fig. 4.--

Oldroyd, 1927, pl. 3, fig. 5.

Im "Clathurella crystallina Gabb, 1865."---of Abbott, 1974:283, fig. 3294.

Philbertia hesione Dall, 1919:54 [unfigured].---Oldroyd, 1927:130.

I Glyphostoma hesione: Abbott, 1974:283 [listed].

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material

l examined:Desc27 LACM lots. ription. Shell profile broad, whorls 4; shoulder broad, smooth, flat to somewhat concave; axial

ribs projecting at periphery and usually extending toward base; spiral cords narrow, overriding axial ribs;

l base with numerous spiral cords. Length 9-12 mrn.

I 141
!

Type Locality and Type Specimens. Off Avalon, Santa Catalina Island, California [from label]. I

Holotype: USNM 109302. Philbertia hesione: off Santa Rosa Island, California, 53 fathoms; holotype:

USNM 211333.

!1

Distribution. San Miguel Island (34°N) to San Clemente Island, California (33°N). I

Habitat. 80-250 m, soft bottoms.

Remarks. This species is characterized by its relatively broad profile, broad shoulder, prominent l[

axial ribs and angulate periphery. Shortly after the description of Clathurella lowei, Dall dismissed it as a

w

synonym of C. crystallina Gabb, the type of which had not been figured. Dall (1919, 1921) figured this 1ira

species under the name C. crystallina. Now there is need for a name for a species that is larger andof broader _l

profile than C. crystallina, and the name Crockerella lowei is here resurrected from synonymy, taking precedence

over the unfigured Philbertia hesione Dall, a later synonym. i

There is extensive variation in the strength and spacing of the axial ribs and spiral cords, as shown by I

the two extremes of sculpture illustrated here. The shell, however, retains a relatively broad profde and a

peripheral carination and does not exhibit the fine clathrate sculpture of Crockerella crystallina. I

I

Crockerella crystallina (Gabb, 1865)

Figures 1.28C I

Clathurella crystallina Gabb, 1865:184.--Grant and Gale, 1931:607.--Abbott, 1974:283 [not fig. 3294].-- al

Coan and Bogan, 1988:278. II

Philbertia crystallina: Dall, 1919:62 [not pl. 13, fig. 1].--Dall, 1921:79 [not pl. 6, fig. 4].---Oldroyd,

1927:129[not pl. 3, fig. 5, copy Dalt, 1921]. I

g

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material

examined: 25 LACM lots. Ill

Description. Shell profile slender, whorls four; shoulder narrow, smooth, slightly concave; axial ribs n

numerous, oblique, low, narrow, not extending across base; spiral cords three at periphery and visible on early

whorls, the middle the most projecting, producing sharp, oblique cancellations with the axial sculpture; base _l

with numerous spiral cords. Length 8-9 mm. Ii

Type Locality and Type Specimens. Santa Catalina Island, California, 40 fathoms. Holotype: UCMP

15959. I Distribution. Santa Rosa Island, California (34°N), to Isla Todos Santos, Baja California (32°N).

Habitat. 50-210 m, soft bottoms. Ill

Remarks. The hallmark of this species is the relatively small size, narrow profile, and clathrate I

sculpture. Axial ribs are numerous but not thick or strongly projecting.

This is the type species of CrockerelIa, figured here for the first time. Dall's illustrations of 1919 and 1

1921 and that of Abbott (1974) that purport to be this species represent C. lowei. The holotype of ClathureUa J

crystalIina has not been figured. The specimen illustrated here is very similar to the holotype.

I

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142 I
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i Crockerella eriphyle (DalI, 1919)

I Figure 1.28D

Mangilia eriphyle Dall, 1919:69 [unfigured].--Oldroyd, I927:136.

i Mangelia eriphyle: Abbott, 1974:279 [listed].

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

examined: 14 LACM lots.

i Description. Shell prof'de narrow, whorls 5; shoulder narrow, smooth except for growth increments;

whorls rounded at periphery; axial ribs 10-12 per whorl, oblique, extending below level of anal sinus; spiral

i cords narrow, Type overridi Localityng anaxial d Type ribs;Sbapecim se withens.nuOff merous Santaspiral RosacoIsland, rds. LenCalifornia, gth 10-12 mm. 53 fathoms. Lectotype

(here selected): USNM 211326.

I Distribution. Santa Cruz Island (34°N) to Cortes Bank, California (32.5°N).

Ill Habitat. 80-250 m, soft bottoms.

i Remarks. This species is characterized by its slender profile, relatively large size, rounded periphery, and broadly spaced axials. It is more slender than Crockerella lowei and does not have the angulate periphery

of that species.

I lot) is notCrocke illustrated reUa ehere riphyle because is figured the shell hereisfbroken or the first in front. time. The tectotype shell (the largest in the original

i Crockerella tridesmia (Berry, 1941)

Figure 1.28E

I Clathurella (Glyphostoma) tridesmia Berry, 1941:8, pl. 1, f. 3.

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

I examined: 13 LACM lots.

Description. Shell profile moderately broad, whorls 4; periphery angulate; shoulder broad, appearing

I below smoth,shoulder bt bearing ad onnubase, merousseparated sharply byincised deeplyspiralncisedstriae; grooves axial ribs of lesser broad; width. spiralLength sculpture 9-11ofmm. broad cords

Type Locality and Type Specimens. San Pedro, Los Angeles County, California, Pleistocene.

I Holotype: SBMNH 34518. Distribution. Santa Rosa Island (34°N) to Cortes Bank, California (32.5°N).

i Habi Remarks. tat. 100Th-is50sp0 emci,essoisftcbotto haract mse.rizedby itsbroad shoulder, smooth except for well-marked, incised

striae, and the more prominent spiral sculpture of the body whorl andbase, which is separatedby deep spiral

grooves. Described No other species has the spiral cords separated by such deep grooves. from Pleistocene deposits, but here recognized in the living fauna. This is a species

characteristic of greater depths (300-500 m) than those recorded for other species of Crockerella.

I

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I 143
!

Crockerella evadne (Dall, 1919) I

Figure 1.28F

m

Mangilia evadne Dall, 1919: 69, pl. 23, f. 2.--Oldroyd, 1927:137. 1

Spirotropis ? evadne: Grant and Gale, 1931:548. Ill

Mangelia evadne: Abbott, 1974:279 [listed]. II

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material Im

examined: 7 LACM lots. I

Description. Shell profile narrow, whorls 4.5; shoulder broad, smooth, flat; sculpture dominated by

sharply projecting peripheral angulation with raised cord at crest; base with relatively few, low spiral cords, ill

Length 8-12 mm. |

Type Locality and Type Specimens. Off Santa Rosa Island, California, 53 fathoms. Holotype:

USNM 211334. I Distribution. Santa Cruz Island (34°N) to Redondo Beach, Los Angeles County, California (34°N).

Habitat. 90-230 m, soft bottoms. m

Remarks. This species is characterized by a single projecting cafination. Some specimens have oblique, I

narrow axials below the carination on the early whorls, but these become obsolete in the final whorl.

There is anerror in the original measurements: the holotype is 7 mm in length, not 12 ram. I

Crockerella conradiana (Gabb, 1869) Ill

Figure 1.29A ]l

Clathurella conradiana Gabb, 1869 (1866-69):7, pl. 1, fig. 12.--Grant and Gale, 1931:606, pl. 26, fig. 11. l

n Man gilia (Clathurella ) conradiana: Arnold, 1903:210.

Glyphostoma conradiana: Oldroyd, 1927:126. I

Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material

examined: 4 LACM lots. mt

Description. Shell profile slender, whorls 5; shoulder narrow, concave, sculpture of about 10 thick g

and strongly projecting axial fibs, overridden by relatively few spiral cords; base with relatively few spiral

cords. Length 10-13 mm. ill

Type Locality and Type Specimens. Santa Barbara, California, Pleistocene. Holotype not located g

at UCMP.

Distribution. Santa Rosa Island, California (34°N), to Isla Todos Santos, Baja California (32°N). l_

Ill Habitat. 25-240 m, soft bottoms.

Remarks. The shell is characterized by its slender profile and relatively few but very strong axial al

fibs. The species was first described from Pleistocene deposits, but was reported living by Dall (1921:179). It l_

remains very scarce in collections. I have not seen live-collected specimens, although the specimens are

sufficiently fresh appearing to indicate that the species is a member of the living fauna. I

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144 N
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If

tl

I Figure 1.29. Conidae. A. Crockerella conradiana (Gabb, 1869): 55 m, off Santa CataSna Island, California (LACM 151660); height 10.4 ram. B. Crockerella scotti McLean, new species: 100 m, Santa

Barbara Channel, California (LACM 2776, holotype); height 11.5 ram. C, CrockerelIa philodoce

i (height Dall, 19.5919n):un439-488 m, off Long Point, Santa Catalina Island, California (LACM 41-182.3); . D. Crockerella cymodoce (Dall, 1919): 59 m, off Goleta, Santa Barbara County,

California (LACM 75-463.2); height 12.6 mm. E. Crockerella castianira (Dall, 1919): 104-113 m,

SE of Bennett Point, San Miguel Island, California (LACM 41-176.7); height 11.6 ram. F. Daphnella

clathrata Gabb, 1865:219 off Isla Vista, Santa Barbara County, California (LACM 74-70.1);

m,

height 19.9 mm.

I

O'ockerel_ sco_ McLean, new species

i Figure 1.29B

i Material Examined. Not represented in the Santa Mafia Basin voucher collection. Other material in addition to type lot (none live-collected): LACM 41-48.15, NW Anacapa Island, California, 88-93 m (1);

LACM 40-111.20, off Redondo Beach, Los Angeles County, California, 176-219 m (1).

i Description. Shell profile broad, whorls 5. Protoconch paucispiral, tip small, elevated, whoris 1.5, last half whorl with strong carination on lower third of whorl. First and second teleoconch whorls with

carinate profile; whorl profile of later whorls rounded below concave shoulder. Shoulder broad, concave,

i bearing faint spiral striae but no axial sculpture. Axial ribs 11-12, of similar strength on final whorl, lip with stronger axial rib behind edge. Spiral cords 3 on early whorls; body whorl with 9 spiral cords including those

on base and canal; cords broad, straplike, overriding axial ribs and with narrower, well-defined interspaces.

i Fine pustular sculpture of genus apparent only under high magnification. Anterior and posterior canals deeply

notched and bordered with callus. Height 11.5 mm (holotype).

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i 145
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Type Locality and Type Specimens. Santa Barbara Channel (34°N, 119°W), California, 100 m, I

Velero IV, sta. 23247, 10 November 1975.9 mature and 1immature specimens, none live-collected. Holotype

LACM 2775, 6 paratypes LACM 2776, 1 paratype CAS 102984, 1 paratype SBMNH 142896, 1 paratype 1

USNM 887575. 1

Distribution. SanMiguel Island (34.0°N) to Redondo Beach, Los Angeles County, California (33.8°N).

Habitat. 85-220 m, soft bottoms. I

gl

Remarks. This species is characterized by its strong axialribs andrelatively broad profile. Inbreadth

it is comparable to Crockerella lowei, but it has much more pronounced fibs than known for that species. In Jib

strength of axial ribs it resembles C. conradiana, from which it differs in having a broader profile and in l

having more numerous axial ribs.

u

The name honors Paul Scott of the Santa Barbara Museum of Natural History. I

u

Crockerella philodoce O)all, 1919)

Figure 1.29C 1

Mangilia philodoce Dall, 1919:66, pl. 22, fig. 7.---Oldroyd, 1927:139. Ill

Mangelia philodoce: Abbott, 1974:279 [listed]. |

Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material I

examined: 3 LACM lots. II

Deseription. Shell profile of medium breadth, whorls 4, rounded at periphery; suture not deeply

impressed; shoulder scarcely defined; axial ribs weak, crossed by narrow spiral cords. Anal sinus relatively _1

broad. Length 7-10 mm. !1

Type Locality and Type Specimens. Off Point Pifios, Monterey County, California, 65-203 fathoms.

Holotype: USNM 208916. I

mu

Distribution. Off Point Pifios, Monterey County (36.5°N), to Santa Catalina Island, California

(33.5°N).

Habitat. 120-490 m, soft bottoms. I

Remarks. This species is characterized by its rounded whorls, subdued axial andspiralsculpture and

relatively broad anal sinus. _1_

This is the second report of this species, which is now known from the holotype and three LACM lots U

of single specimens reported here. The holotype and the specimen illustrated here have eroded apices, but

there is one immature specimen that has the characteristic protoconch that confirms the assignment to i

Crockerella. The original illustration of the holotype was enhanced, adding missing portions of the shell, as II

was commonly done for illustrations made at the USNM for publications by DaU and by Bartsch.

i

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I Crockerella eymodoce (Dall, 1919)

Figure 1.29D

I Glyphostoma cymodoce Dall, 1919:54, pl. 17, fig. 6.--Oldroyd, 1927:127.--Abbott, 1974:283 [listed].

I examined:Mate1LACM rial Exlot. amined. Not represented in the Santa Maria Basin voucher collection. Other material

Description. Shell profile slender, whorls 5; shoulder broad, marked with incised striae; sculpture of

I about 10 strongly projecting axial ribs, overridden by spiral cords of alternating strength. Anal sinus deep, bordered by callus; lip thickened, crenate at edge to match spiralsculpture. Length 11-13 mm.

Type Locality and Type Specimens. Santa Barbara,California [depth not cited]. Le_totype (here

i selected): USNM 150569.

Distribution. Monterey (36.5°N) pall, 1921] to Santa Barbara, California (34.5°N).

I Habitat. 60 m, soft bottoms. Remarks, This species is characterized by its relatively strong axial ribs and spiral sculpture of

alternating strength, andthe pronounced tubular aspect of the anal sinus. Inits strength of sculpture it recalls

l some tropical species of Glyphostoma, but it has the protoconch of Crockerella, not Glyphostoma.

The specimen figured here (from 59 m off Goleta, Santa Barbara County, California) is the only

specimen other than the holotype that I have seen. It comes from a locality very close to the original type

I locality.

I CrockereUa castianira (Dall, 1919)

Figure 1.29E

t Clathrodrillia castianira Dall, 1919:15, pl. 2, fig. 1.

CIathurella castianira: Abbott, 1974:283 [listed].

I Material Examined. Not represented in the Santa Maria Basin voucher collection. Other material

examined: 1 LACM lot.

I whorl, above Descripeptriion. phery;Shell axial profile ribs strong, broad, whnumerous; orls 4; shoulder spiral cords concavbroad, e, narroverriding ow; shouldaxial er angribs; ulatiseparated on high obyn

narrower interspaces, 3 cords exposed on penultimate whorl; base with numerous spiral cords. Length 10-

I 11.5 mm.Type Locality and Type Specimens. OffCapeSanMartin, Monterey County, California, 218 fathoms.

Holotype: USNM 214246.

t Distribution. Cape San Martin, Monterey County (36°N), to San Miguel Island, California (34°N). Habitat. 100-400 m, soft bottoms.

i Remarks. This species is characterized by a broad profile, a narrow, concave shoulder and moderately

strong axial and spiral sculpture. It differs from all forms of C. lowei in having broad rather than narrow

spiral cords and in having a high shoulder. All forms of C. lowei have a midwhorl angulation.

I

!

i 147
!

Two specimens are now known, the holotype and one LACM specimen from 104-113 m, SE of I

Bennett Point, San Miguel Island, California, which is a good match for this species (Figure 1.27E). The

holotype is badly eroded, missing the protoconch and the fin'sttwo teleoconch whorls. Dali's illustration of the i

holotype was reconstructed to show sculpture on the second teleoconch whorl that is not there. In doing so, an |

error was made in showing too many axial ribs on the second teleoconch whorl, judging from the lesser

number on the LACM specimen, i

Subfamily Daphnellinae Deshayes, 1863 i

i

Diagnosis. Small to large; anal sinus at suture, reversed L-shaped, sculpture usually cancellate, often

with smooth shoulder. Pmtoconch usually multispiml, usually diagonally cancellate. Operculum absent. Radutar i

teeth with large solid base, barbed or unbarbed tips, opening laterally at the base. B

Remarks. This is another large subfamily, in which there are genera with exceptions to allthe diagnostic

characters (Taylor et al., 1993:162) i

Genus DaphnelIa Deshayes, 1863 I

Daphnella Deshayes, 1863.Type species (SD Herrmannsen, 1847): Pleurotoma lymnaeformis Kiener, 1839- H

40. Caribbean.

Diagnosis. Shell with rounded whorls and fine reticulate sculpture; axial ribs becoming obsolete on l

final whorl; anterior end truncate, lip flaring, thickened in some species; protoconch diagonally reticulate.

I

Biology. The multispiral protoconch is indicative of an extended planktotrophic veliger stage.

Remarks. This is a widespread tropical genus occurring in shallow water.

Daphnella clathrata Gabb, 1865 I

Figure 1.29F

I

Daphnella clathrata Gabb, 1865: 185.---Oldroyd, 1927:150.-----Grant and Gale, 1931:542, pl. 25, figs. 23, i

24.--Hertz, 1980:97, figs. 1-5.----Coan and Bogan, 1988:278.

l

Material Examined. California: Santa Maria Basin, Phase I, sta. 2, 200 m (1); sta. 4, 393 m (1). B

Other material: 18 lots in the LACM collection.

Description. Shell of moderate size, slender; whorls 5, rounded; aperture nearly half length of shell; I

sinus deep, at suture; axial and spiral sculpture coarsely clathrate on early whorls, changing to numerous II

spiral cords on final whorl; protoconch of 3.5 diagonally reticulate, dark brown whorls. Length 15-20 mm.

Type Locality and Type Specimens. Santa Catalina Island, 60 fathoms. Holotype: UCBP 15960. I

Distribution. Santa Mafia Basin (35.5°N), to Isla Cedros, Baja California [LACM 71-158] (28°N).

Habitat. Soft bottoms, 40-150 m. i

Remarks. Daphnella clathrata is the only species of the genus yet known from California. It is more II

characteristic of deeper water than are the tropical species. Hertz (1980) discussed the species in detail.

I

!

148 I
!

I Acknowledgments

I I am grateful to editorPaul Scott for guiding this work to completion. His manipulation of

computer

contrast for the figures served to improve the plates. I appreciate the careful reading of earlier drafts and

helpful suggestions made by a number of reviewers. Sections treating Eulimidae and Muricidae were read by

I Anders Wartn and Emily Vokes, respectively. Text of the complete manuscript was read by Gene Coan, Jerry Harasewych, and Lindsey Groves. Alan Kabat checked the status of many type lots at the USNM, in addition

to reading the manuscript. Bob Van Syoc and Terry Gosliner assisted with specimens at the CAS. Lindsey

I Groves at the LACM assisted with cataloging chores.

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i!

I 2. THEOPISTHOBRANCHIA

I by

i Terrence M. Gosliner 1

I Introduction

I mollusks.TheHowever, Opisthobrecent ranchiaphylogenetic traditionally investigations have been con(Haszprunar, sidered as one1988; of three Gosliner, subclasse 1991, s of1994) gastropod have

demonstrated that the subclass Prosobranchia is paraphyletic. Not all of the descendants of the common

I ancest withinor

thearePulmonata. included inForthatthistaxreason, on, sincgastropod e some areclassi cons fication idered has opisthundergone obranchsfundamental and others are restructuring. contained

There is general consensus that opisthobranchs and pulmonates are more closely related to each other than

i tEuthyneura o other gastropod are parttaxofa. aTogether, larger gastropod the opisthotaxon, branchs the Heterobranchia, and pulmonates are whicuni h t

includes ed into thetheEPyramidellidae, uthyneura. The

Architectonicidae, Mathildidae, Rissoellidae, and Valvatidae. Details of phylogeny of the Heterobranchia

have not been precisely determined at present. It appears that the Opisthobranchia, as presently constituted,

I is monophyletic (Gosliner, 1981).

The Opisthobranchia consists of approximately 5,000 described species. This almost exclusively

I ma completely rine taxonshell-less representsslugs. a morpholog Shell reduction ically diverse and loss group, hascontaining occurred she repeatedly lled snails, andinterme independently diate formswithin and

different lineages. This fact accounts for much of the morphological diversity of this taxon.

I Traditionally, the Opisthobranchia has been divided into five major orders of benthic snails and two additional orders of holoplanktonic organisms. The ranks for all of these orders are likely not equivalent. At

least one order, the Cephalaspidea, probably is paraphyletic and is not diagnosed by any shared derived

I features level. . For this reason, taxa formerly included in the Cephalaspidea are here considered at the family

In general, opisthobranch mollusks can be readily identified to species by their external morphology.

I The shape of the body and color pattern are essential to identification, especially in the brightly colored nudibranchs. For this reason, it is far easier to identify living or freshly fixed material, in which many of

these features are preserved. Preserved material can also be identified, but dissections of the mouth parts

I and details of reproductive anatomy are often required. Since most of the present material has been preserved for long periods of time, every attempt has been made to differentiate species on the basis of external

features that are retained in preservation. In some cases, internal features of the shell (when presen0 and

i radular teeth must also be examined to insure correct identification.

The opisthobranchs of the shallow temperate Pacific coast of North America have been reasonably

well studied. Behrens (199i) provides color photographs of living animals of some 217 species of

I opisthobranchs from Alaska to the outer coast of Baja California. This review permits the identification of

living material, including 26 undescribed species.

I

_Departmenotf Inve_eN'ateZoologyandGeology,CaliforniaAcademyofSciences,GoldenGatePark,SanFrancisco,

I California94118-4599

i 161
!

The opisthobranch species inhabiting North American waters below 100 m depth remain poorly •

studied. Chamberlain and Behrens (1980) recorded five species of opisthobranchs from deep water off San

Luis Obispo County, California. Other reports of deeper water opisthobranchs are scattered in the literature II

(eg. Willett, 1944, MacFarland, 1966). Most of the taxa recorded are shelled species, known only from dead II

shells. In several instances the familial and generic status of these species remains in doubt, because

morphological features of preserved animals are required for more refined systematic placement. In these RI

cases, shell morphology alone does not permit proper systematic placement. II

The present material from the Santa Mafia Basin includes well preserved specimens of 28 species

of opisthobranchs. Of these, half have never been described morphologically, including seven likely •

undescribed species not previously recorded from the Pacific coast of North America. Two genera,

Parvamplustrum and HolopIocamus, were previously known only from the Atlantic. Thus, description of

this material adds significantly to our primary knowledge of the opisthobranch fauna of the Pacific coast of

North America. II

Morphology I

i

The most obvious morphological difference between different opisthobranchs is the presence, •

absence, and form of the shell. The ancestral shell of opisthobranchs is similar to that found in caenogastropods. m

In this instance, the shell is external and thickly calcified, with a well elevated spire that extends beyond the

body whorl. This form of shell is generally found in species of Acteonidae, Ringiculidae, some species of I

Acteocina and isolated representatives of several other opisthobranch taxa. In more derived opisthobranchs

that still retain an external shell, the spire is sunken completely within the body whorl. This bulloid shell is

present in a wide array of opisthobranchs and has been derived independently at least four different times I

within the Opisthobranchia. Many other opisthobranchs retain an internal shell that is situated on the dorsal

surface, directly underneath the dorsal epithelium. Sometimes the shell is visible through the translucent

epithelium, but in other instances dissection may be required to observe the shell. In most cases, an internal i

shell is a relatively flattened, thinly calcified plate. As in the case of the bulloid shell, internal shells have II

been independently derived in several distinct groups of opisthobranchs.

Other external features can be used to differentiate taxa. Many less derived opisthobranchs have a !

body divided into a cephalic head shield and a posterior shield. These two portions of the body are usually g

flanked by a pair of lateral parapodia. Other more modified taxa have a pair of chemosensory rhinophores

situated on the dorsal surface of the head. In the case of the Notaspidea, Anaspidea, and many Sacoglossa, i

the rhinophores are roiled rather than solid. In other sacoglossarts and all nudibranchs the rhinophores are

J

solid. The rhinophores may be smooth or ornamented with lamellae, annulae, or papillae. In notaspideans,

a single pinnate gill is situated on the right side of the body. In nudibranchs, gills are arranged in a wide i

variety of fashions, including a simple circle in most dorids and linear gills along the edge of the notum is RI

most Dendronotacea.

In some instances, internal features provide essential information for identifying and differentiating !

species. The most diagnostic features include the morphology of the jaws and whether chitinous rodlets are

m

present or absent. If rodlets are present their form may be important in differentiating closely related species.

Probably the most important feature used in differentiating species of opisthobranchs is the structure of the R

rasping teeth situated within the buccal mass. These teeth are arranged in distinct patterns forming the

complex structure of the radula. In many instances, a central or rachidian row of teeth is present, while in

others these teeth are entirely absent. The central region is generally flanked on either side by lateral teeth el

that are variable in form and arrangement. Often individual species will have a specific number of rows of 1R

lateral teeth within the radula.

!

162 |
!

I In some less derived opisthobranchs, the region at the posterior end of the esophagus is enlarged to

form a muscular gizzard. Within the gizzard are a series of distinct chitinous or calcareous plates. Their

I relative size Opisthobranchs and shape areareoften characteristic and species specific. largely hermaphroditic and have complex arrangements of reproductive organs.

The structure of these organs is important in determining the systematic placement of many taxa. Owing to

I the de-emphasized difficulty infordetermining the purposesreproductive of this study.morphology, its utility in species identifications has been

I Collection and Fixation

I Opisthobranchs can be collected by most standard collection methods though soft-bodied species

can be damaged during collection and sorting. The preferred method of collection of epifaunal species is

directly by hand or trawling. Living soft-bodied opisthobranchs should be retained in salt water, as this

I causes less damage to specimens. Infaunal material can be collected by a variety of dredging and coting

methods. Careful and prompt sorting will minimize damage to specimens. It is important to make color

notes or photographs of living or fresh material, as color is important in distinguishing closely related

I species.

Once material has been sorted and documented, specimens should be relaxed by gradually adding

an 8% solution of magnesium chloride to a shallow dish containing just enough sea water to cover the

I specimens, magnesium place over a period of 8-12 hours to ensure reduced

Addition of chloride should take

contraction of specimens upon fixation. Probing of specimens is necessary to ascertain whether they are

properly anesthetized. If specimens contract or move when probed, more time must be allowed before

specimens are fixed. Specimens should be fixed in buffered 10% formalin Bouin's fixative. The latter

or

will dissolve shells, but provides better fLxationof soft tissues. Ideally, if shelled specimens are sufficiently

abundant, lots should be split and some representatives should be placed in 75% ethyl alcohol, while the

I remainder should be fixed in Bouins. Unbuffered formalin will also dissolve shells, especially if they are left in formalin for extended periods of time. After 24-48 hours, specimens can be transferred from fixative

to 75% ethyl alcohol for long-term storage.

I

Laboratory Methods

!

Most opisthobranchs are relatively small, less than 30 mm in length. Detailed morphological

i examination details of theofmorphology living and preserved of the jaws,specimens radula andreqgiz uireszard,usespecimens of a dissecting mustmicroscope. be dissected.InToorder examine to examine these

structures, the muscular buccal mass and gizzard are carefully removed from the animal and placed in a

i 10% solution Dissecoftionsodiuofmspecimens or potassiisumgenerally hydroxideconducted for approximately using a dissecting 24 hours.microscope, together with ultra

fine forceps or minute insect needles. Tissue can be teased away from the remaining hard parts using minute

I insect water. needles Permanent . The slides remaining then labial can becutiprepared cle, radulbya, anlight d gizzard stainingplates withshoudilute ld beacidthorofuchsin ughly rinsed and mounted in distilledin

aqueous mounting medium or by gradual dehydration in alcohol, emersion in xylene and mounting in

I balsam. Gizzard plates should not be mounted in acidic aqueous mounting media as this rapidly dissolves the calcium carbonates present in some plates. Specimens for scanning electron microscopy should be airdried on a round microscope coverslip after removal from distilled water.

!

I 163
!

Glossary I

I

Aperture. Opening of the shell into which the Punctate sculpture. Shell texture consisting of a I

animal may retract, series of irregular pits within the surface of

w

the shell. A unique derived feature of

Body oWh°rl f 'the shell.The largest and terminal spiral whorl opisthobranchs retained in many taxa. I

Radula. Ribbon of chitinous teeth used for rasping

Bulloid spire. A shell in which the earlier whorls food. m

axe entirely surrounded by the body whorl. •

Radular formula. A numerical depiction of the 1

Cephalic lobes. Paired sensory projections on the

arrangement of radular teeth. Expressed in

dorsal surface of the cephalic shield, the following fashion: a-b x c.d.e.d.c. The I

Cephalic shield. The anterior portion of the body first numbers (a-b) indicate the number of g

of certain bubble snails used to burrow radular rows times the number of teeth per

through soft substrate, row. In the above example, c represents the 1

number of outer lateral teeth, d the number II

Elevated spire. A shell in which the earlier whorls of inner lateral teeth, and e the rachidian

project posteriorly from the level of the body teeth. A zero indicates that none of that •

whorl, particular type of teeth is present in that |

Half row. The radular teeth forming a row on radula, For example, the number of rachidian

either side of the central portion of the teeth is expressed as either 1 or 0, depending 1

radula, on whether a rachidian row is present or I

absent.

Jaws. Chitinous structures used for grasping and •

cutting food, which are situated at anterior Rachidian teeth. The single line of teeth I

end of buccal mass. occupying the central region or rachis of the

radula.

Lateral teeth. The radular teeth situated on either •

side of the central portion of the radula. Inner Velar tentacles. Appendages that may be present U

lateral teeth are often of different form than along the anterior and lateralmargins of the

outer ones. This may be a gradual or abrupt head. I

transition of tooth form. II

Notum. The dorsal surface of the animal, ll

Oper II culum. The trap door that may cover the

aperture of the shell when the animal is

completely retracted. In most opisthobranchs II

the operculum is absent. It is retained in

many members of the Acteonidae and a few

members of some other taxa. When present I

in opisthobranchs it is a thin, oblong, II

chitinous plate.

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I List of Opisthobranch Species

I Family Acteonidae Family Aldisidae

Rictaxis punctocaelatus (Carpenter, 1864) Aldisa sanguinea (Cooper, 1863)

i Family Acteon Hydatinidae sp. Family Cadlinaflavomaculata Chromodorididae MacFarland, 1905

Parvamplustrum sp. Family Polyceratidae

I Family Cylichnidae Holoplocamus sp.

Cylichna diegensis (Dall, 1919)

Acteocina harpa (Dall, 1871) Family Goniodorididae

I Acteocina eximia Okenia sp.

(Bakd, 1863)

Family Philinidae Family Dendronotidae

I Philine polystrigma (Dall, 1908) Dendronotusfrondosus (Ascanius, 1774) Philine sp. 1 Family Tritoniidae

Philine sp. 2 Tritoniafestiva (Stearns, 1873)

I Tritonia diomedea Bergh, 1894 Family Aglajidae

Aglaja ocelligera (Bergh, 1893) Family Arminidae

i Aglaja sp. Armina californica (Cooper, 1863)

Family Gastropteridae Family Tergipedidae

Gastropteron pacificum Bergh, 1893 Cuthona rolleri Behrens and Gosliner,

I Family Retusidae

1988

Retusa xystrum Dall, 1919 Family Aeolidiidae

i Volvulella Volvulella panamica caIifornica Dall, Dall, 1919 1919 AeolidEliot, iella i905) chromosoma (Cockerell and

Family Diaphanidae Cerberilla mosslandica McDonald and

I Diaphana californica Dall, 1919 1975 Nybakken,

Family Pleurobranchidae

I BPleurobranchaea erthella californicacalifornica (Dall, 1900 MacFarland, )

1966

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Key to Opisthobranch Species I

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1A. Shell external ................................................................................................................................... 2 i

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lB. Shell internal or absent .................................................................................................................. 12

2A. Shell spire elevated above body whorl ............................................................................................ 3 I

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2B. Shell spire bulloid ............................................................................................................................ 8

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3A. Shell with punctate sculpture .......................................................................................................... 4

3B. Shell without punctate sculpture, though spiral lines may be present ............................................ 6 I

4A. Shell with alternating black and white bands .............................................. Rictaxis punctocaelatus II

4B. Shell uniformly white ...................................................................................................................... 5

5A. Radula with numerous minute denticulate teeth; all identical in shape and size .............. Acteon sp. I

5B. Radula with 5 teeth per half-row; outer teeth more elongate juvenile ........ Rictaxis punctocaelatus

6A. Posterior end of foot bifurcate ......................................................................... Diaphana californica I

6B. Posterior end of foot undivided ....................................................................................................... 7

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7A. Edges of whorls straight .......................................................................................... Acteocina harpa

7B. Edges of whorls curved .......................................................................................... Acteocina eximia I

8A. Shell with elongate, pointed posterior end ...................................................................................... 9

8B. Shell flat or rounded posteriorly .................................................................................................... 10 I

9A. Outer lip joining apex of posterior extension ................................................ Volvulella californica I

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9B. Outer lip joining base of posterior extension ................................................... Volvulella panamica

10A. Shell with numerous longitudinal ridges ................................................................... Retusa xystrum I

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10B. Shell smooth or with spiral lines ................................................................................................... 11

l

11A. Cephalic lobes bifid ........................................................................................... Parvamplustrum sp.

1lB. Cephalic lobes rounded ....................................................................................... Cylichna diegensis I

12A. Shell internal ................................................................................................................................. 13

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12B. Shell absent .................................................................................................................................... 19 I

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I 13A. Shell with punctate sculpture ........................................................................................................ 14

13B. Shell smooth or with simple growth lines ..................................................................................... 17

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14A. Gill present on right side of body ..................................................................... Berthella californica

I 14B. Gill hidden within posterior cavity ............................................................. genus Philine ........... 15

15A. Gizzard plates absent ..................................................................................................... Philine sp. 2

I 15B. Gizzard plates present ................................................................................................................... 16

I 16A. Shell with posterior flange ............................................................................................ Philine sp. 1

16B. Shell rounded posteriorly ................................................................................... Philine polystrigma

I 17A. Posterior shield rounded posteriorly .......................................................... Gastropteron pacificum

17B. Posterior shield with rounded posterior lobes .............................................................................. 18

I 18A. Shell occupying most of posterior shield cavity ............................................................... Aglaja sp.

I 18B. Shell occupying only posterior third of cavity ...................................................... Aglaja ocelligera

19A. Circle of gills present on back of animal ...................................................................................... 20

I 19B. Circle of absent 23 gills

I 20A. 20B. Gills Gills retractile not retractileinto.......................................................................................................................... pit ................................................................................................................... 2122

I 21A. Radular teeth short, variable in form ............................................................ Cadlinaflavomaculata

21B. Radular teeth all elongate with pectinate denticles ................................................ Aldisa sanguinea

I 22A. Velar tentacles present ......................................................................................... Holoplocamus sp.

22B. Velar tentacles absent ....................................................................................................... Okenia sp.

I 23A. Gills absent, animal with cerata .................................................................................................... 24

23B. Gills present in some form ............................................................................................................ 26

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24A. Body dorsoventrally flattened .................................................................................. Cuthona rolled

I 24B. Body not dorsoventrally flattened ................................................................................................. 25

25A. Rhinophores smooth, cylindrical .................................................................. Cerberilla mosslandica

I 25B. Rhinophores bulbous, lamellate ................................................................. Aeolidiella chromosoma

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26A. Single pinnate gill present along right side of body ............................ Pleurobranchaea californica i

26B. Gills otherwise ............................................................................................................................... 27

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27A. Gills situated on ventral side between mantle and foot ....................................... Armina californica

27B. Gills branched, situated along lateral margins of notum .............................................................. 28 I

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28A. Velar tentacles branched .............................................................................. Dendronotus frondosus

28B. Velar tentacles unbranched ........................................................................................................... 29 •

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29A. Notum smooth ........................................................................................................... Tritoniafestiva

29B. Notum with low tubercles .................................................................................... Tritonia diomedea

I

Descriptions of Species 1

J

Family Acteonidae

I

Genus Rictaxis Dall, 1871 I

Type Species: Tornatella punctocaelata Carpenter, 1864. i

1

Diagnosis. Shell well calcified with elevated spire and punctate sculpture. Operculum present or

absent. Jaws with well-developed rodlets. Radular formula 5.0.5. Radular teeth elongate with small cusps. •

Reproductive system androdiaulic. Penis armed or unarmed. |

Rictaxispunctocaelams (Carpenter, 1864) im

Figures 2.1A, 2.2A, 2.3A

Tornatella punctocaelata Carpenter, 1864:139. I

Rictaxis punctocaelatus: Dall 1871:136.

Material Examined. California, Santa Maria Basin, off Point San Luis, Sta. 21, rep. 3 (1), dissected; I

off Point Sal, Sta PJ-9, rep. 1 (1).

Description. Living animal translucent white (Figure 2.1A). Shell white with alternating black l

bands in mature individuals. Juveniles with uniformly white shell (Figure 2.2A). Spire moderately elevated. II

Aperture wide, elongate. Body whorl with numerous rows of fme punctations. Thin coriaceous operculum

visible along surface of foot of juvenile specimens. Operculum absent in adults. Jaws with triangular to •

linear rodlets. Radular formula 27-35 x 5.0.5. (Figure 2.3A). Inner 3 teeth per half row short with multifid

cusps. Outer 2 rows of teeth with long denticulate cusps. Outermost tooth denticulate in juveniles, more

elongate, smooth in mature specimens.

Biology. Rictaxis punctocaelams is known from intertidal mud and sand fiats to at least 100 m

depth. This species, like other members of the family, feeds on potychaete annelids.

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Figure 2.2. Parvam Rictaxpluisstrum punctAoca" elprese atUSrved(Carp sp., entx1er8'64)' animal j80uvenil , , e shell, x 40. B. Acteon sp., shell, x 30. C. ii

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Remarks. DalI (1871) described Acteon punctocaelams as the type species of the genus Rictaxis. •

Though the subgeneric distinction was originally based on conchological features, Marcus (1972) determined

that the radula of several species, including R. punctocaelatus, was significantly different from that of other •

acteonids. Rictaxis is characterized by a radula with five rows of elongate teeth with numerous small denticles II

along the main cusp. A rachidian row of teeth is absent. In juvenile specimens, such as the material examined

here, the cusp of the outer lateral tooth is short and bears denticles. In older specimens the cusp is more •

elongate and lacks denticles. II

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I The sympatric "Rictaxis" painei has a rachidian row of teeth and there is not an elongate cusp. In

"R". painei there are eight lateral teeth per half row, as compared to five in species of Rictaxis. It is unlikely

I that acteonid R. painei genus,should based beon considered radular morphology. as a member Theofgeneric this genus. positionThisofspecies this taxondoesremains not fitopen readilyto question. into any

Rictaxis punctocaelatus differs from the four other acteonids recorded from off the California coast

I inandhaving dark spiral prominent bands alternating on the shell,black but theandbands whitearebands not prominent. on the shell. Acteon Acteontraskii traskiialsoalsohashasa radula alternati consisting ng light

of numerous minute radular teeth per half row. Rictaxis painei has a uniformly white or brown shell and a

I different well developed radularsiphonal formula.furrow. Microgyphis estuarinus and M. breviculus have uniformly white shells with a

Type Locality and Type Specimens. San Diego, California; holotype (USNM 14914).

I Distribution. Ketchikan, Alaska to Magdalena Bay, Baja California (Behrens, 1991), intertidal zone to 100 m.

I Genus Acteon Monffort, 1810

Type Species: Bulla tornatilis Linnaeus, 1758.

I Diagnosis. Shell well calcified with elevated spire and punctate sculpture. Operculum present.

Jaws with well-developed rodlets. Radular formula 50-150.0.50-150. Radular teeth small with numerous

I small denticles. Reproductive system androdiaulic. Penis unarmed.

Aeteon sp. Figures 2.2B, 2.3B-C

I Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BSV 42, 1 (1),

rep.

dissected.

i Description. Shell (Figure 2.2B) 3 mm long, uniformly white with 22 finely punctate spiral lines on body whorl. Shell with three whorls. Outer lip of shell broken. Animal with large operculum on foot.

Jaws (Figure 2.3B) with multifid elements. Radula containing numerous (more than 50) teeth per half row.

Teeth minute (Figure 2.3C), with 16-18 denticles along inner edge of tooth. Penis short, digitiform, with

I rounded apex.

Remarks. This species is a member of the genus Acteon, as the radula bears numerous rows of

i

, Acminute, teon traskii finely. deThenticul aperture ate teethofActeon with many sp. isteeth widerperandhalftherowwhole . It apshell pearsistmore o be conc squathothan logicially nA. dis traskii tinct. Afromlso,

A. traskii is colored with rose bands that are absent in the present shell. It is possible that the present

i specimen is a juvenile of A. traskii, but the dissected animal appeared to be sexually mature. The shell is similar in shape toActeonpanamensis Dall, 1908, which is known only from a single shell collected in deep

water from the Gulf of Panama. Further material is required to confirm the identity of this species.

I Distribution. Thus far, this species is known only from the single specimen collected from the Santa Mafia Basin.

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Figure 2.3. A.Rictaxispunctocaelatus (Carpenter, 1864), radular teeth, x 1000.B, C.Acteon sp, B.jaw rodlets, I

x 3000. C. radular teeth, x 4000. D. Parvamplustrum sp., radular teeth, x 2000.

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I Family Hydatinidae

I Genus Parvamplustrum Powell, 1951 Type Species: Parvamplustrum tenerum Powell, 1951.

I Diagnosis. Shell weakly calcified, bulloid. Operculum absent. Jaw rodlets reduced to chitinous patches. Radular formula 1.0.1. Lateral teeth flattened, without denticles. Reproductive system androdiaulic.

Penis unarmed.

I

Parvamplustrum sp.

I' Figures 2.2C, 2.3D

Material Examined. California, Santa Maria Basin, off Point Arguetlo and Point Conception; Sta.

I 61, rep. 1 (I); Sta 86, rep. 2 (1), dissected. Description. Preserved animals minute (Figure 2.2C), less than 1 mm in length. Shell bulloid, often

completely decalcified in formalin fixed material. Foot short, triangular, and acute posteriorly. Tentacles on

I sides of head bifid. Jaw with unarmed polygonal rodlets. Radular formula approximately 20 x 1.0.1. Lateral teeth (Figure 2.3D) curved and broad without denticles. Gizzard plates absent.

Remarks. The genus Parvamplustrum is known only from the type species, P. tenerum Powell,

I 1951, which is known only from the Atlantic (Powell, 1951; Marcus and Marcus, 1969; Gosliner, 1991).

The present species represents the fh-strecord of the genus from the Pacific. It is similar to P. tenerum in all

respects. More material needs to be examined to determine if the present specimens are conspecific with P.

I tenerum, or they represent an species, especially important

whether undescribed This is since the shells of

the present material are completely decalcified.

i The genus Parvamplusturm is placed in the Hydatinidae, though the radular morphology and shell are different from other members of the family. Resolution of the familial status of this genus requires

additional study.

I Distribution. Thus far, this species is known only from the Santa Maria Basin. .I

I Family Cylichnidae

Genus Cylichna Loven, 1846

I Type Species: Cylichna cylindracea (Pennant, 1777).

Diagnosis. Shell bulloid, well calcified. Operculum absent. Jaws with multifid rodlets. Radular

I formula 3-12.1.1.1.3-12. Rachidian teeth bilobed, denticutate. Inner lateral teeth denticulate. Outer lateral teeth smooth. Gizzard plates 3, smooth, equal in size and shape. Reproductive system monaulic. Penis

unarmed.

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Cylichna diegensis (Dall, 1919) :l

Figures 2.4A, 2.5

I

Cylichnella diegensis Dall, 1919:300. i

Cylichna diegensis: Abbott, 1974:315.

Material Examined. California, Santa Mafia Basin, off Point Sal, Sta. 103, rep. 1 (2); Sta R-8, rep. I

1 (5), 1 dissected; Sta PJ-17, rep. 3 (1).

Description. Shell bulloid (Figure 2.4A), external, 3-7 mm in length. Shell with fine spiral striae 1

and brown periostrical lines at anterior and posterior ends, but absent from central region. Preserved animals 1

uniformly light tan.

Jaws with multifid rodlets (Figure 2.5A). Radular formula 13 x 3-4.1.1.1.3-4. Rachidian teeth (Figure 'lli

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2.5C) bifid, with 5-7 denticles on either side of tooth. Inner lateral teeth (Figure 2.5B) with 9-11 denticles on

inner side of hook-shaped primary cusp. Outer lateral teeth curved with numerous fine denticles. Denticles

absent from outermost tooth. Lenticular gizzard plates 3, (Figure 2.5D), of equal size and shape, i

i

Biology. Little is known about the biology of CyIichna diegensis. Other members of the genus are

known to feed on Formaminifera.

Remarks. The anatomy of Cylichna diegensis has not previously been described. Cylichna diegensis II

is similar in appearance to Cylichna attonsa. The two species are known to be sympatric from Santa Monica

south to Baja California. It is appears that C. diegensis has sharper shoulders to its shell than C. attonsa. I

Internally, C. diegensis has 3-4 rows of outer lateral teeth per side while a single specimen of C. attonsa II

from British Columbia examined here has 6-7 rows of outer laterals per side. Both species are unusual in

having denticles on the outer side of the cusp of the outer laterals. More material must be examined to i

ascertain whether Cylichna diegensis and C. attonsa are indeed distinct. 1

Type Locality and Type Specimens. Off Point Loma, San Diego, California, 98-191 fathoms;

(USNM Distribution 209071).

. This species is known from Santa Monica, California to Baja CaLifornia (Abbott, I

1974).

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Genus Acteocina Gray, 1847

Type Species: Acteocina wetherelli (Lea, 1833). 1

Diagnosis. Shell well calcified with elevated or bulloid spire. Operculum absent. Jaw rodlets reduced,

multifid. Radular formula 1.1.1 or 1.0.1. Rachidian teeth bilobed, denticulate when present. Lateral teeth el

denticutate. Gizzard plates 3, unequal in size and shape. Reproductive system monanlic. Penis unarmed or II

armed.

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1 D

I Figure 2.4. A. Cylichna diegensis (Dall, 1919), shell, x 20. B. Acteocina eximia (Baird, 1863), shell, x 10.C., I).

Acteocina harpa (Dall, 1871), C. shell, x 40. D. Gizzard plates of holotype, x 25.

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Figure 2.5. Cylichna diegensis (Dall, 1919). A. Jaw rodlets, × 3000. B. Lateral teeth, × 1000. C. Rachidian teeth, I

× 2000. D. Gizzard plate, × 150.

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I Acteocina harpa (DaU, 1871)

Figures 2.4C-D, 2.6A-B

I Tornatina harpa DaU, 1871:136.

Utriculastra harpa: Thiele, 1925:239.

I Acteocina (Coleophysis) harpa: Abbott, 1974:313.

Material Examined. California, Santa Maria Basin, off Morro Bay, Sta. 6 (1); off Point Conception,

I Sta. BRA 02 (1), dissected.

Description. Shell (Figure 2.4C) external, 1.5 mm in length. Shell with slightly elevated spire.

Sides of shell parallel for most of their length. Aperture wide. Jaws with indistinct rodlets. Radular formula

I x teeth (Figure 2.6A) bifid, with 3-4 denticles on either side of tooth. Inner lateral

12-I7 1.1.1 Rachidian

teeth (Figure 2.6B) with 9-10 coarse, elongate denticles on inner side of hook-shaped primary cusp. Gizzard

plates dissolved in present, formalin fixed material. Gizzard plates of holotype examined in present study

I 2.4D). Plates in size and Dorsal Lateroventral but

(Figure une_qual shape. plate largest. plates narrower,

asymmeWical.

i BRiemark ologys. .Like Theoanat theromembers my ofActeocina of this geharpa nus, Acteocina has not previously harpa likely beenfeeds described. upon fOwing oraminftoerans. this lack

of information, its generic and familial status has remained in question.Acteocina harpa is clearly a member

I ofsmaller comones. plx ofThespecies radularthatteethare ofcharmembers acterizedofbythishaving complex a single havelarge a prominent heart-sharowped ofgizzard rachidian plateteeth and twando

lateral teeth with a short flange containing large denticles. Other members of this complex from the Pacific

I Acoast . harpa of Nhas orth more Americasymmetri a includecalActeoc gizzard ina inc plates ulta,andA. carinata a rachidianand possibly tooth withA, fewmagdalens denticles. is. OfThetheseshellspecies of A,.

harpa is unique in having longitudinal incised lines on the body whorl of the shell. The other species of this

i complex Tappear ype Loctoality be restric and tTyedpeto Sinptecertidal and shallow subtidal waters. imens. Monterey, California; holotype (USNM 56179).

Distribution. Forrester Island, Alaska to San Diego, California (Abbott, 1974).

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Acteocina eximia (Baird, 1863)

I Figures 2.4B, 2.6C-D

Bullina eximia Baird, 1863:67.

I Tornatina eximia: Pilsbry, 1895:189.

Acteocina eximia: Strong, 1921:44.

i Acteoc Utriculastra ina culcculcitella itella exi: mia Marcus, : Willett, 1977:19222,8:37. figs 71-76. Not Gould, 1852.

CyIichnella culcitella: Gosliner, 1979:87, fig.3. Not Gould, 1852.

I Acteocina culcitella intermedia WilieR, 1928: 37. New

synonymy.

Material Examined. California, Santa Maria Basin, off Point Sal, Sta. 31, rep. 1 (1); Stat. PJ-15,

I rep. Conception, 1(1), dissected; off Point San Luis, Sta. R-2, rep. 3 (1); off Purisima Point, Sta. R-4, rep. 2 (1); off Point Sta 85, rep. 1 (1).

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Figure 2.6. Acteoc eximiaina(Baird, harpa1863). (])all,C18. Lateral 71). A. tRaehidian eeth, x 100teeth, 0. D. xGizzard 4000. B.platLateral es, × 20. teeth, x 3000. C., D.Acteocina I

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I Description. Shell (Figure 2.4B) external, well calcified, 4-10 mm in length. Spire slightly elevated

from body whorl. Shell ornamented with numerous brown, spiral periostrical lines. Radular formula 19-28

i x 1.0.1. Rachidian row of teeth absent or consisting of few vestigial teeth in some specimens. Inner lateral teeth (Figure 2.6C) curved with long central denticte. Thirty to 50 short denticles present along almost

entire inner margin of tooth. Gizzard with 3 unequal plates (Figure 2.6D). Two plates large and concave on

i their dorsal Biology. surface. ThisSingle speciesplateappears small,tolaterally be restricted compresse to depths d, concexceeding ave. 22 m. Foraminifera have been

observed in the gizzard of some specimens and appear to be the primary food of this species.

I Remarks. Members of this complex of Acteocina are characterized by having two large and one small gizzard plate. The rachidian row of radular teeth is greatly reduced and rachidians are either vestigial

or entirely absent. The inner lateral teeth have small denticles along most of the inner margin. These denticles

are not on an elevated flange as in species of the Acteocina harpa complex. Considerable confusion has surrounded the taxonomic status of this species and members of this species complex. Willett (1928)

considered three subspecies ofA. culcitella, A. culcitella culcitella (Gould, 1852),A. culcitella eximia (Baird,

I to1863), A. culcitella andA. culcbyitella Marcusintermedia (1977). WiUett, 1928.Acteocina eximia was separated from specimens attributable

Gosliner (1979) considered the material regarded by Marcus asA. eximia andA. culcitella to represent

a single species, based on examination of specimens with intermediate anatomy. Thus, the two species were united under the older name, A. culciteUa. Subsequently, I have discovered that the syntypes ofA. culcitella

contained portions of dried animals. These were examined and it was determined that A. culciteUa is

anatomically distinct from all material considered by either Marcus or Gosliner as A. culcitella orA. eximia. The small gizzard plate of the syntypes of A. culciteUa is dorsally flattened while that of A. eximia is

laterally compressed. Thus, all specimens considered as A. culcitella orA. eximia by Marcus and Gosliner,

I must be regarded as A. eximia. Acteocina culcitella is known from shallow waters from the intertidal zone to approximately 10 m depth from Monterey south to San Diego, while A. eximia is known from depths

exceeding 22 m in the northern part of its range (Vancouver Island) and as shallow as 60 m in southern

I Calif and Ao.rnia. culcitella Concholog has aically strong, Acolumellar . eximia canpleat be sethat paraistedabsent fromAin, Aculcite . eximiaUa.. The The spire anatomy is shoforter A.inculcite A. exUimia a is

identical to that of A. rolleri (Marcus, 1977) and the latter represents a junior synonym of A. culciteUa.

I Type Locality and Type Specimens. Vancouver Island, British Columbia; paratype (BMNH 1869.3.21.78).

Distribution. Vancouver Island, British Columbia to San Diego, California. I

Family Philinidae

I Genus Philine Ascanius, 1772

I Type Species: Bulla aperta Linnaeus, 1767.

Diagnosis. Shell internal bulloid to plate-like. Jaw rodlets absent. Radular formula 0-6.1.0-1.1.0-6.

Gizzard teeth present or absent: when present, 3 in number. Reproductive system monaulic. Penis variable

I in shape, simple or elaborate.

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Philine polystrigma (Dall, 1908), new combination I

Figures 2.7A-C, 2.8A-B

i

Clistaxis ?polystrigma Dall, ] 908:246. I

Material Examined. California, Santa Maria Basin, off Point San Luis, Sta. 25, rep. 2 (1); Sta 25, []

rep. 3 (1), dissected, secondary voucher; Sta. R-3, rep. 3 (1), dissected; off Point Sal, Sta. PJ-13, rep. 3 (1); II

Sta. PJ-]4, rep. 3 (1).

Description. Living animals 0.5-3.0 mm in length. Preserved animals uniformly light tan. Body I

divided into anterior head shield, posterior shield and lateral parapodia (Figure 2.7A). Anterior head shield A

anteriorly andposteriorly rounded. Posterior end of posterior shield roundedandterminating in a blunt lobe

on either side. Shell internal, completely calcified, occupying almost all of posterior shield. Shell narrow •

with broad body whorl on columellar side. Shell with spiral, lightly punctate sculpture. Radular formula 16

x 1.0.1. Lateral teeth (Figure 2.8A) with elongate cusp and 25-30 fine, triangular denticles along the

masticatory margin. Gizzard plates 3 (Figure 2.8B), of equal size and shape. Plates lenticular with thick l[

prominent longitudinal ridge on dorsal side. Central nervous system (Figure 2.7B) euthyneurous. Penis II

(Figure 2.7C) simple with elongate convoluted prostate.

Biology. This species is found only from subtidal fine sediments where it is collected in dredge, _l

core, and trawl samples. Members of this genus feed on hard shelled prey such as bivalves. Smaller species, il

such as P. polystrigma, feed almost exclusively on Foraminifera.

Remarks. Clistaxis polystrigma was described from a single specimen collected from San Diego 1

Harbor (Dall, 1908). It has not been recorded since its original description. Dall was uncertain of the systematic II

status of this species. The dried holotype was rehydrated and examined in this study. The three equal,

lenticular gizzard plates and radula with an inner lateral tooth per side clearly indicate this species is properly 1

placed within the genus PhiIine. All of the anatomy of this holotype, including deeply punctate shell sculpture, inl

shape of gizzard plates, and radula with a formula of 1.0.1, is consistent with specimens examined in this

study. I

Philine bakeri was described from a single shell collected off South Coronado Island. The shell is

similar to the holotype of P. polystrigma. In 1944, Willett described Philine californica from a single shell I

collected off Redondo Beach (LACM 1075). He did not compare this species to other taxa. However, in l_

comparing the holotype shells of P. baker/and P. californica with that ofP. polystr/gma, it appears that they nun

are all similar in shape and size. Philine baker/and P. californica require further study and comparison with

P. polystr/gma. I

Philine polystrigma is one of five philinids known from the temperate Pacific coast of California.

Additional boreal species have been recorded from Alaskan waters (Lee and Foster, 1985). Philine alba i

Mattox, 1958 is readily distinguishable from P. polystrigma by its larger body size, smooth shell devoid of l

any distinct sculpture, proportionately smaller smooth gizzard plates, and simple penis with only a single,

Ill

straight prostate. MacFarland (1966) and Behrens (1991) erroneously identified specimens of P. alba as P.

baker/. I

Philine polystrigma is most similar to the sympatric species Philine sp. 1.The shell ofP. polystrigma

is narrower and more tightly coiled than that of P. sp. 1. Also, the shell of P. sp. 1 has much larger, deeper i

punctations to its sculpture. Internally, the radula of P. polystrigma has only a single row of lateral teeth on |

either side of the radula, while PhiIine sp. 1 has an additional outer row of teeth on either side. The gizzard

plates of the two are similar in shape, though the mid-rib of the plate is thicker and broader inP. polystrigma. I

The penis of P. polystrigma is simple with elongate prostate. In PhiIine sp. 1, the penis is complex, with l

separate ejaculatory and prostatic portions.

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I Figure 2.7. A-C. Philinepolystrigma (Dall, 1908). A. Preserved animal, x 25. B. Cenlral nervous system, x 40. C. Penis, x 40. D-F. Philine sp. 1, D, Preserved animal, x 15. E. Shell, x 20. F. Penis, 40.

I Type Locality and Type Specimens. Off San Diego Harbor entrance, 50 fins; holotype (USNM

110649).

I Distribution. This species is known from Santa Barbara to the Coronado Islands (Abbott, 1974).

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Figure 2.8. A., B. Philine polystrigma. A. Lateral teeth, x 700. B. Gizzard plate, x 50. C., D. Philine sp. 1, C. I Lateral radular teeth, x 1200. D. Gizzard plate, x 70.

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I Philine sp. 1

Figures 2.7D-F, 2.8C-D, 2.9A

l Material Examined. California, Santa Maria Basin, off Point San Luis, Sta. 23, rep. 3 (2), one

dissected, primary vouchers; Sta. 21, rep. 3 (1); Sta. PJ-2 (2); offPoint Sal, Sta. PJ-8 (1); Sta. PJ-11 (1); Sta.

I PJ-15 (1); Sta. PJ-22 (1); Sta. R-8 (1) dissected. Description. Preserved animals 1-5 mm in length, uniformly tan. Body divided into anterior head

shield, posterior shield andlateral parapodia (Figure 2.7C). Anterior head shield anteriorly and posteriorly

I rou Lobes nded. approximately Posterior endequal of posterior in size. shield rounded and terminating in a short, triangular lobe on either side.

Shell (Figure 2.7D) internal, completely calcified, occupying most of posterior shield. Shell flat,

i broad with obvious pitted, punctate sculpture (Figure 2.9A). Posteriormost portion of shell extended into pointed lobe. Columellar portion of shell narrow. Radular formula 15 x 1.1.0.1.1. Inner lateral teeth (Figure

2.8C) large curved, denticulate with 22-34 elongate denticles. Outer lateral teeth shorter and simple, without

I denticles. Gizzard plates (Figure 2.8D) 3, equal in size. Plates lenticular with prominent longitudinal ridge

' on dorsal side. Penis (Figure 2.7F) complex with separate ejaculatory and prostatic ducts. Prostate extremely

elongate and convoluted. Penial papilla simple, conical, unarmed.

I This is known from fine soft sediments in 90-155 of water (Dall, 1908; Biology. species only m

present study). Like Philine polystrigma, Philine sp. 1 probably feeds on Foraminifera.

i Remarks. Philine sp. 1 is morphologically similar to the sympatric species Philine polystrigma.

Specific differences are discussed above, and include different shell shape and sculpture, radular formula,

gizzard plates, and penial morphology.

I Distribution. This species is known only from the Santa Maria Basin (present study).

i FigurPhiline sp. 2 es 2.9B-C, 2.10A

I Ma D terial Examined. California, Santa Maria Basin, off Point San Luis, Sta R-1 (1), dissected. escription. Single specimen (Figure 2.10A) approximately 1 mm in length. Head shield short,

rounded posteriorly. Parapodia short, not extending over head and posterior shields. Posterior shield elongate

I with elongate skirt-like posterior lobes. Shell internal, largely decalcified irt specimen examined. Gizzard plates absent, thin walled crop present. Radular formula 13 x 6.1.0.1.6. Inner lateral teeth (Figure 2.9B)

broad with hook-shaped cusp. No secondary denticles present on inner side of tooth. Outer lateral teeth

I narrower hook-shaped without secondary denticles. Penis with short bulbous prostate. Remarks. This species appears to be undescribed. It is the only species along the Pacific coast with

more than one outer lateral tooth per side of the radula. The only other species of Philine with as many rows

i of radular teeth is P. pruinosa. It differs in having a denticulate inner lateral tooth (Thompson, 1976). Philine sp. 2 is the only species of Philine along the California coast that lacks gizzard plates. Additional

material is required before this species can be described.

I Distribution. Thus far, Philine 2 is known only from the Santa Maria Basin.

sp.

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Figure 2.9. A. Phitine sp. l, shell sculpture, x 300. B. Phifine sp. 2, radular teeth, x 700. C. Gastropteron

pacificum Bergh, 1893, radular teeth, x 300. I

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I Figure 2.10. A. (Bergh, Philine1893). sp. 2,Cpreserve . Preserved d animal, animal, x 80. x 5.]3.D. Aglaja Shellsp., , x pre lO.served E. Penanimal, is, x 10.x 40. C.-E. Aglaja ocelligera

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Family Aglajidae I

Genus Aglaja Renier, 1807

It Type Species: Aglaja tricolorata Renier, 1807.

Diagnosis. Shell an internal, completely calcified plate. Radula absent. Gizzard plates absent.

Reproductive systems monaulic. Penis armed or unarmed. It

Aglaja ocelligera (Bergh, 1893) N

Figures 2.1B, 2.10C-E

Doridium ocelligerum Bergh, 1893a:133.--Bergh, 1894: pl 10, fig. 10; pl. 12, figs. 5,6. I

Aglaja ocelligera: Pilsbry, 1896.--Rudman, 1974: fig. 13.--Gosliner, 1980:330, figs. 2, 14.

Doridium adellae Dall, 1894:73.--Pilsbry, 1896: pl. 9, fig. 22.--Gosliner, 1980: 330. I

,j

Chelidonura phocae Marcus, 1961:8, figs. 18-24.

Aglaja phocae: Rudman, 1974:207.

U

Material Examined. California, Santa Maria Basin, off Point San Luis, Sta 21, rep. 3 (2), primary

vouchers; Sta. PJ-1 rep. 3 (1); off Point Sal, Sta. PJ-15, rep. 3 (1). _1

Description. Living animals (Figure 2.1B, 2.10C) 3-20 mm in length and brownish black with |

small opaque white spots. Body divided into anterior head shield, posterior shield, and lateral parapodia.

Anterior head shield anteriorly lobed with laterally directed projection on either side of body. Posterior end _i

of head shield rounded. Posterior shield terminating in elongate "tail" on left side of body and shorter II

triangular lobe on right side. Lobe on left side may be greatly contracted in preserved material, but still

visible. Shell (Figure 2.10D) internal, completely calcificed, occupying posterior portion of posterior shield.

Radula and gizzard plates absent. Penis (Figure 2.10E) armed with row of cuticular spines. Prostate elongate, I

convoluted.

Biology. This species is common on intertidal mudflats and subtidal habitats consisting of free sand it

or silt. This species is predatory on other invertebrates, though the precise feeding preferences remain !1

unknown. Like other members of the Aglajidae, it ingests its prey whole by means of suction from its

muscular buccal mass. /I

Remarks. This species is one of five aglajids known from the temperate Pacific coast of North

America. It is the only species with a simple lateral lobe of the anterior portion of the head shield, a thin,

completely calcified shell, and a penis with numerous chitinous spines. The three species of Navanax also I

present in the region have an anterior portion of the head shield with four distinct lobes. The posterior ends 1!

of the posterior shield are subequal in length as compared to the elongate left tail present in A. ocelligera.

The three species of Navanax also have different shaped shells. In Navanax aenigmaticus and N. inermis the i

shell has a thin calcified portion and a broad membranous portion. In N. polyalphos, the shell is completely J

calcified, but broader than that of A. ocelligera (Gosliner and Williams, 1972). Melanochlamys diomedea

has a simply rounded anterior portion of the head shield, blunt posterior ends of the posterior shield, and a I

shell that occupies most of the cavity of the posterior shield. The systematics and synonymy of Aglaja !1

ocelligera were discussed by Gosliner (1980).

Type Locality and Type Specimens. Type locality not designated in original description. Type

material unknown.

Distribution. This species is known from the Pacific coast of North America from Sitka, Alaska to

Mission Bay, San Diego (Behrens, 1991). I

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Figure 2.10B

Material Examined. California, off Purisima Point, Sta. R-4, rep. 1 (4); offPoint Sal, Sta. R-8, rep.

2 (2); Sta. PJ-15, rep. 3 (1).

Description. Body uniformly whitish brown, 0.5-1 mm in length. Body divided into anterior head shield, posterior shield, andlateral parapodia (Figure 2.10B). Anterior head shield anteriorly rounded.Posterior

end of head shield rounded. Posterior shield terminating in elongate "tail" on left side of body and shorter

I triangular lobe on right side. Lobe on left side may be greatly contracted in preserved material, but still visible. Shell internal, completely calcificed, occupying most of cavity of posterior shield. Radula and

gizzard plates absent. Penis immature in all material examined.

i Biology. Little is known about the biology of this species. One of the specimens examined had a

large spionid polychaete filling the thin-walled crop.

I Remarks. This species appears to be distinct from A. oceUigera. Although both species have an

elongate ffliform appendage on the left side of the posterior portion of the posterior shield, Aglaja sp. has a

shell that occupies most of the posterior shield cavity. In A. ocelligera, the shell is a thin band that occupies

only the posterior portion of the shield. While the present specimens axe immature, it appears that they

l differ from A. based differences. Further is confirm the

oceUigera,

on conchological study required to

distinctness of this species.

i Distribution. Thus far, this species is known only from the Santa Maria Basin.

i Family Gastropteridae

Genus Gastropteron Meckel, in Kosse, 1813

I Type Species: Gastropteron rubrum Meckel in Kosse, 1813.

Diagnosis. Shell internal reduced to small, flat plate. Parapodia large, natatory. Siphon a simple

l tube. systemRamdular onaulic. formPenis ula 4-6. simple 1.0.1.4-6. to complex. Inner lateral teeth denticulate. Outer laterals smooth. Reproductive

I Gastropteron pacificum Bergh, 1893

Figures 2.1C, 2.9C, 2.11A-B

I Gastropteron pacificum Bergh, 1893b:303, pl. 16 fig. 28, pl. 17, figs. 10-26.

Gastropteron (pacificum Bergh vat. ?) cinereum Dall, 1925: fig. 4.----Gosliner, 1984:243.

i Gastropteron cinereum: Tokioka and Baba, 1964:206.

Material Examined. California, Santa Maria Basin, off Point San Luis and Point Sal, Sta 21, rep.

I 3 (1); StaDes PJ-cri 1 prep. 3 (2); Sta. PJ-6, rep. 1 (2); Sta. R-8, rep. 2 (1). tion. Living animals (Figure 2.1C) 0.5-30 mm in length. Living animals whitish with scattered

red-brown spots. Body divided into anterior head shield, posterior shield, and lateral parapodia. Anterior

I head siphon. shield Posterior roundedshield androunded slightly lposteriorly, obed anteriorly. without Postelateral rior enflagellum. d of head Parapodia shield termilarge, natingmuscular, in simple,usedfoldeford

swimming. Gill situated on right side of posterior shield; consisting of approximately 15 pirmae. Anus

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Figure 2.11. A, B. Gastropteron pacificum Bergh, 1893. A. Shell, after MacFarland, 1966. B. Penis, x 25. C. I

Diaphana californica shell, x 10. D. Retusa xystrum Dall, 1919, shell, x 50. E. Volvulelta californica |

Dall, 1919, shell, x 25. F. Volvulellapanamica Dall, 1919, shell, x 30.

situated near posterior end of gill. Shell (Figure 2.11 A) internal, completely calcificed, occupying posterior i

portion of posterior shield. Radular formula 20-22 × 5-6.1.0.1.5-6. Inner lateral teeth large, denticulate

(Figure 2.9C). Outer lateral teeth smaller, without denticles. Gizzard plates absent. Penis (Figure 2.11B) I

with row of cuticular papillae. Prostate elongate, highly convoluted. 11

Biology. Gastropteron pacificum is common subtidally to depths of 400 m, and occasionally found

on intertidal mudflats. Like other members of the Gastropteridae, it is predatory on other invertebrates, Ill

though the precise food remains unknown. This species is often collected in plankton nets, owing to its J

propensity for swimming through the water column. However, it is primarily a benthic rather than a planktonic

species. I

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I Remarks. This species is the only gastropterid known from the Pacific coast of North America.

Another species, Gastropteron cinereum Dall, 1925, has been shown to be a synonym of G. pacificum

i parapodia. (Gosliner, 1984). It is readily distinguishable by its short, rounded posterior shield and large natatory

Type Locality and Type Material. Aleutian Islands. Type material unknown.

i Distribution. This species is known from the Pacific coast of North America from Alaska to Baja California (Behrens, 1991).

I Family Retusidae

i Genus Retusa Brown, 1827

Type Species: Retusa obtusa (Montagu, 1807).

I Diagnosis. Shell with elevated or bulloid spire. Operculum present or absent. Jaws and radula

absent. Gizzard teeth 3, equal or subequal, tuberculate.Reproductive system monauhc. Penis simple,unarmed.

I, Retusa xystrum Dall, 1919

Figures 2.11D, 2.I2A-B

I Retusa xystrum Dall, 1919:297.

I 2 (20); Sta. Mate21, rialrep.Exa1 (18); minedSta. . Cali R-I,fornia, rep. Santa 1 (2), rep. Mafi3a (4); Basinoff, off Purisima Point San Point, LuiSta. s anR-8, d Point rep.Sal, 1 (2). Sta. 21, rep.

I)eseription. Shell (Figure 2.11D) external, bulloid, with numerous longitudinal striae (Figure 2.12A).

i Shell 1.0-1.5 mm long. Jaws and radula absent. Gizzard plates (Figure 2.12B) unequal; one plate wider than other 2. Plates with rounded tubercles. Largest tubercle situated on inner side of each plate.

Biology. Little is known about the biology ofRetusa xystrum. Other species of Retusa are known to

I feed on foraminiferans.

Remarks. Retusa xystrum its poorly known and its anatomy was undescribed prior to this study. It

has not been adequately compared toR. galapagana Dall, 1919, known only from the Galapagos, R. carpenteri

I 1 known from andR. 1919, known from the Gulf of California Hanley, 859, Mazatlan, Mexico, paz/ana Dall,

from San Luis Gonzaga Bay to La Paz. These species are all very similar conchologically to R. xystrum, but

their anatomy remains unknown. Further investigation is required to establish the systematics of the eastern

Pacific Retusa spp.

Some workers place species with similar shell morphology within the genus Sulcoretusa. At present

i this division is based solely on conchological features. More information regarding retusid anatomy is required before this division can be substantiated.

Type Locality and Type Specimens. San Diego, California; holotype, (USNM 273985).

I Distribution. This species is known from San Pedro to San Diego, California (Dall, 1919).

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Figure 2.12. A.,B.RetusaxystrumDall, 1919. A. Shell sculpture, x 500. B. Gizzard plates, x400. C., D.Diaphana

californica Dall, 1919. C. Rachidian teeth, x 1500. D. Lateral teeth, x 1500. I

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I Genus Volvulella Newton, 1891

Type Species: Volvulella acuminata (Brugi_re, 1792).

I Diagnosis. Shell bulloid, acutely pointed posteriorly. Operculum absent. Jaws, radula and gizzard

plates absent. Reproductive system monaulic. Penis simple.

I Volvulella californica Dall, 1919

I Figure 2.11E

Volvulella californica Dall, 1919:298.

i Material Examined. California, Santa Maria Basin, off Point Estero, Point Sal, and Point Arguello, Sta. 3, rep. 1 (1); Sta. 61, rep. 1 (1); Sta. R-8, rep. 1 (1); Sta. R-8, rep. 2 (1).

i Description. Shell (Figure 2.11E) 1-3 mm in length, smooth, without sculpture, widest in middle, tapered at both ends. Posterior projection acute, elongate. Operculum, jaws, radula, and gizzard plates absent.

Biology. Little is known about the biology of this species. The present material was collected from

I 90-345 mRedepth. marks. Abbott (1974) suggested that VolvuleUa californica may be a "fat" form of V. cylindrica.

Volvulella californica is consistently devoid of sculpture and pyriform in shape, while V. cylindrica has

I sshape, culpturebutanddiffers is farfrommorebothcylindri V. californica cal. Volvuland ella Vpana . cylindrica mica (seein fthat ollowing the posterior descriptiend on) ofisitsalsouter o cylinlipdricbegins al in

at the base of the posterior spine rather than at its apex.

I

211303).

Type Locality and Type Material. Off Santa Rosa Island, California, 53 fins.; holotype (USNM

i Distribution. Santa Cruz, California to Todos Santos Bay, Baja California (Abbott, 1974).

Volvulella panamica Dall, 1919

l Figure 2.11F

VolvuleUa panamica Dall, 1919:298.

I VolvuleIla tenuissima Willett, 1944:71, 14, 1.--Keen, 1971.

pl. fig.

Material Examined. California, Santa Mafia Basin, off Point San Luis, Point Sal, and Point

I Con 8, rep. ception, 2 (4); StSta. a. 80R-8, , rep. rep.2 (31)(3). ; Sta. 86, rep.1 (1); Sta. 79 (1); Sta. R-l, rep. 1 (2); Sta. R-8, rep. 1 (1); Sta. RDescription. Shell (Figure 2.11F) cylindrical, 2.5-5 mm in length, shell smooth, devoid of sculpture,

I with prolonged posterior projection. Posterior end of outer lip originating at base of posterior projection. Jaws, radula and gizzard plates absent.

Biology. Little is known about the biology of this species. The present material is from 90-197 m

_1 depth.

Remarks. Differences between this and other sympatric species are discussed above under Volvulella

I californicaTyp. eTheLocprealsent material represents a northern range extension from Redondo Beach, California. ity and Type Specimens. Panama Bay, 60 m; holotype (USNM 212654).

Distribution. Santa Mafia Basin, California to Panama (Abbott, 1974, present study).

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Family Diaphanidae I

Genus Diaphana Brown, 1837

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Type Species: Diaphana minuta (Brown, 1827).

Diagnosis. Shell thinly calcified. Spire slightly elevated or butloid. Operculum absent. Jaws reduced. I

Radular formula 1.1.1. Rachidian teeth bilobed, denticulate. Lateral teeth elongate with minute denticles. g

Gizzard plates absent. Reproductive system monaulic. Penis simple or complex.

I

Diaphana californica Dall, 1919 I

Figures 2.1F, 2.11C, 2.12C-D I

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Diaphana californica Dull, 1919:299-300.

Material Examined. California, Santa Mafia Basin, off Point San Luis, Sta. 21, rep. 2 (1), secondary I

voucher; Sta. PJ-1, rep. 3 (2); Sta. PJ-19, rep. 2 (2); Sta. PJ-22, rep. 2 (1), dissected.

P

Description. Living animal as in Figure 2.1F. Shell (Figure 2.11C) with slightly elevated spire, 1-5 I

mm in length. Anterior portion of outer lip flared. Jaws reduced, without distinct armature. Radular formula I

13 x 1.1.1. in one small specimen examined. Rachidian teeth (Figure 2.12C) with 2 denticle bearing lobes,

each with approximately 20 denticles. Lateral teeth (Figure 2.12C) curved. Minute denticles present along i

outer edge of tooth. Penis complex, with elongate prostate. 1

Biology. Little is known about the biology of species of Diaphana. They appear to be carnivorous

and are found under rocks with a well developed growth of a variety of colonial animals. Diaphana californica I

is found from tide pools to subtidad depths. Specimens in this study were collected from 49-167 m depth.

Remarks. Species of Diaphana have been recorded as having smooth lateral teeth. This is an

artifact of poor resolution of light microscopy. Scanning electron micrographs clearly show denticulations i

of the outer lateral teeth in all species that have been examined by this means (Gosliner, 1994). Diaphana W

brunnea Dall, 1919, described from Kodiak Island, Alaska, may be a synonym of D. californica. Further

study is required to resolve their taxonomy.

Type Locality and Type Material. Long Beach, California; holotype (USNM 130561). 'm

Distribution. Cape Arago, Oregon to Coronados Islands, Baja California (Behrens, 1991).

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I Family Pleurobranchidae

i Genus BertheUa Blainville, 1825

Type Species: Berthella plumula (Montagu, 1803).

I Diagnosis. Shell internal, flattened, with sculpture. Gill external, without tubercles. Jaw

punctate

elements smooth to multifid. Radula broad with many smooth teeth. Rachidian teeth absent. Reproductive

system diaulic. Penial gland present. Penis unarmed. !

Berthella californica (Dall, 1900)

t Figures 2.1D, 2.13A-B, D, 2.14A

Pleurobranchus californicus Dall, 1900:92.

I Pleurobranchus chacei Burch, 1944.--MacFarland, 1966:84.

Pleurobranchus californicus denticulatus MacFarland, 1966:84, pl. 5. figs. 1-5, pl. 13, figs. 25-34, pl. 16,

fig. 12.

I BertheIla californica: Lance, 1966:71.--Gosliner and Bertsch, 1988:46, figs. 1A, 2,3.

BertheIla denticulatus: Lee and Foster, 1985:442.

I Material Examined. California, Santa Maria Basin, off Point Buchon, Sta. BRA 13 rep. 4, (1),

dissected.

I the dorsalDessurface. cription.Gill Living external, animalssituated (Figureon2.1right D) white sidetoofbrown body.withShescattered ll internal,opaque coveting whitemost spotsofcovering dorsal

_ surface. Shell (Figure 2.14A) with numerous irregular rows of punctate sculpture (Figure 2.13A). Jaws with

t Radular numerousformula rows of34triangular x 26.0.26 rodlets, in specimen withexamined. (Figure 2.13B) Innermost or witeeth thout(Figure a series2.13D) of denticles with bifidalongcusp.margin. Outer

teeth more elongate.

rocky habitats Biologyfrom. Most intertidal speciespoofolsBerthella to about 20feedm ondepthcompound (Bertsch tunicates. and Gosliner, BertheIla 1988).californica The presentisspecimen found in

was dredged from 88.5-100.5 m depth.

i Remarks. Gosliner and Bertsch (1988) have reviewed the morphology and systematics of eastern

Pacific species of Berthella. The only sympatric species, BertheUa strongi, differs in its coloration and

radular and reproductive morphology.

I Type Material and Locality. San Pedro, California, holotype (USNM 107893).

Distribution. Point Craven, Alaska to Coronados Islands, Baja California (Behrens, 1991).

I Genus Pleurobranchaea Leue, 1813

!Ill Type Species: Pleurobranchaea meckelii Leue, 1813.

Diagnosis. Shell absent. Gill situated on right side of body. Anterior margin of velum tuberculate.

Notum and foot broad. Rhinophores well separated. Jaws with muttifid rodlets. Radular teeth elongate with

I acute bifid diaulic. Penis armed unarmed.

cusp. Reproductive system or

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Figure 2.13. A, B. Berthella califomica (Dall, 1900). A, Shell sculpture, x 300. B. Jaw rodlets, x 1500. C. I Pleurobranchaea califomica MacFarland, 1966,jaw rodlets, x 1200. D. Berthella californica (Dall,

1900), inner radular teeth, x 200. E. Pleurobranchaea californica MacFarland, 1966, radular teeth,

x 500. i

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Figure 2.14. A. Berchella caIifornica (Dall, 1900), shell, x 30.B., C. Holoplocamus sp. B° dorsal view of preserved

specimen. C. Ventral view of preserved specimen, x 40. D. Okenia sp., ventral view of preserved

I specimen, x 65. E, Cerberilla mosslandica McDonald and Nybakken, 1975, living animal. F. Tritonia diomedea Bergh, 1894, living animal.

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Pleurobranchaea cah'fornica MacFarland, 1966 I

Figures 2.1E, 2.13C, E

Pleurobranchaea californica MacFarland, 1966:94-101. pl. 15, figs. 16-28, pl. 17, figs. 1-17. '

Material Examined. California, Santa Maria Basin, off Point Sal, Sta PJ-6, rep. 1 (1); Sta. PJ-6, Ill

rep. 2 (1), dissected; Sta. PJ-7, rep. 2 (1); Sta. R-4, rep. 1 (1). I

Description. Living animals (Figure 2.1E) 3-210 mm. Body brown with translucent and opaque

white patches. Rhinophores well separated. Anterior margin of head with numerous papillae. Foot elongate, i

extending well beyond notum posteriorly. Gill pinnate, situated on right side of body. Shell absent. Jaw II

platelets (Figure 2.13C) polygonal with numerous denticles along anterior margin. Radular formula 22-52 x

38-145.0-1.38-145. Lateral teeth (Figure 2.13E) with elongate (injuvenile specimens) or rudimentary cusp i

(in mature specimens). Penis muscular, unarmed. '_l i

Biology. Pleurobranchaea californica is a voracious carnivore and feeds primarily on sea anemones

and other opisthobranchs. It is very opportunistic and feeds on a wide variety of prey. It is found from 1

10-400 meters depth and is one of the most common subtidal opisthobranchs in soft substrate habitats. U

Remarks. The systematics of Pleurobranchaea was reviewed by Marcus and Gosliner (1985). i

Pleurobranchaea californica was placed in its own subgenus, Macfarlandaea, based on the fact that it has l_

lateral teeth with a rudimentary cusp. Willan (1987) stated that this separation was unnecessary. I concur

i

with his rationale for disregarding the subgeneric separation, especially since juvenile specimens have welldeveloped bifid teeth as in other members of the genus. Pleurobranchaea californica is the only member of l[

the genus present along the Pacific coast of North America. It differs from other sympatric pleurobranchids i

in lacking an internal shell.

Type Locality and Type Specimens. San Francisco, outside Golden Gate, southeast of Farallon II

Light, California; syntypes- (USNM 5775225; CASIZ 021594, 024055, 029179, 067460, 067462, microscope

slides 089256-089266). /

Distribution. Port Orford, Oregon to San Diego, California (Behrens, 1991). I

Family Aldisidae I

Genus AMisa Bergh, 1878

Type Species | : Doris zetlandica Alder and Hancock, 1854.

Diagnosis. Notum with low rounded tubercles. Rhinophoral and gill pockets ringed with tubercles. I

Labial tentacles reduced into broad oral pad. Jaws reduced to thin labial cuticle. Radular teeth extremely II

elongate with denticles along margin and at apex. Reproductive system trianlic. Penis lined with rows of

chitinous spines, i

Aldisa sanguinea (Cooper, 1863) ill

Figures 2.15A, 2.16A-B J

Doris (Asteronotus) sanguinea Cooper, 1863:204. Ill

Aldisa sanguinea: MacFarland, 1966:169, pl. 25, fig. 8, pl. 29, fig. 11, pl. 35, figs. 17-22.--MiUen and g

Gosliner, 1985: figs. 2, 4A, 61, 13.

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i Figure 2.15. Li C.vTritoniafe ing animals. A.Aldisa sanguinea (Cooper, 1863),B. Cadlinaflavomaculata MaeFarland, 1905. stiva (Steams, 1873).D. Dendronotusfrondosus (Ascanius, 17741.

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i dissected.Material Examined. California, Santa Maria Basin, off Point Conception, Sta. BRA 2, rep. 1 (1),

Description. Living animals (Figure 2.15A) up to 20 mm in length. Color bright red orange with 2

, large mid-dorsal darker spots, occasionally with tan flecks forming a T-shaped mark anterior to gills. Notum

covered with low rounded tubercles. Oral tentacles reduced, forming a broad area anterior to foot. Jaws

without rodlets. Radular teeth numerous, elongate (Figure 2.16A) with numerous denticles along apical

I margin and along inner side (Figure 2.16B). Penis armed with several rows of hook-shaped barbs.

Biology. This species feeds on sponges of the genera Ophlitaspongia and Hymendesmia (Behrens,

I 1991). It is found from intertidal rock pools to a depth of 80 m.

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Figure 2.16. A, B. Aldisa sanguinea (Cooper, I863], radul_ teeth, x 500. C., D. O_n_ sp. C. Inner t_eral teeth,

x 1500. D, Outer lateral teeth, x 3000. I

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I Remarks. The taxonomy of the eastern Pacific species of Aldisa was reviewed by Millen and

Gosliner (1985). Four species are known from the Pacific coast of North America, yet A. sanguinea is the

I only species found south of Trinidad Head, in extreme northern California. Aldisa cooperi is similar to A.

• sanguinea in having orange body color, but differs in having a series of small dark dots rather than a pair of

large ones. It also has internal differences of the radula and reproductive system (Millen and Gosliner,

i Type Locality and Type Specimens. San Diego Bay, California. Type material unknown.

1985).

Distribution. Aldisa sanguinea is known from Coos Bay Oregon, to the Gulf of California (Behrens,

I 1991).

I Family Chromodorididae

Genus Cadlina Bergh, 1879

mm

I Type Species: Doris laevis Linnaeus, 1767.

Diagnosis. Notum with low rounded tubercles. Labial tentacles auriculate. Defensive glands present

I along margins of notum. Jaw rodlets bifid. Radular teeth cuspidate. Rachidian tooth well developed with multifid cusps. Inner and outer lateral teeth denticulate. Reproductive system u'iaulic. Penis with rows of

chitinous spines.

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Cadlina flavomaculata MaeFarland, 1905

I Figures 2.15B, 2.17A-C

Cadlinaflavomaculata MacFarland, 1905:43-44; 1966: pl. 23, fig. 1; pl. 29, fig. 17, pl. 33, figs. 12-21.

i Material Examined. California: Santa Maria Basin, off Point Arguello and Point Conception, Sta.

BRC 1 (1); Sta. BRA 6, rep. 2 (1), dissected.

I of defensive Descriglands. ption. Rhinophores Living animaldark s (Figure brown 2.15 to black. B) toBody 20 mmovoid. in length. Gills Cwhite, olor white bipinnate. with submarginal Marginal yellowring

ring present. Notum tuberculate. Oral tentacles reduced, auriculate. Jaws present, with numerous bifid rodlets

(Figure with 4 primary 2.17A).cusps. RadulaInner broad. lateral Radular teeth with formulinner a 89andx 1outer 5.1.15. denticles Rachidian on side teethof (Figure primary 2.17 cusp.B)Outer denticlateral ulate

teeth (Figure 2.17C) with 5-12 denticles on outer side of primary cusp. Outer laterals increasingly long and

with moreBi denticles towards outer edge. Penis armed with rows of hooks. ology. Cadlina flavomaculata feeds on sponges of the genus Aplysilla and is found from the

intertidal zone to 200 m depth (Behrens, 1991). Present specimens were dredged from 60-70 m depth.

i Remarks. Cadlinaflavomaculata is one of five sympatric members of the along the

genus present

Pacific coast of North America, It is the only species with dark rhinophores and white gills. The only other

species with dark rhinophores is C. limbaughorum, which also has dark gills. Dark rhinophoral pigment is

I often retained in preserved specimens. Radular differences distinguish the various species of Cadlina (McDonald, 1983).

Type Locality and Type Specimens. Monterey Bay, California; holotype (USNM 181287).

I Distribution. Cadlinaflavomaculata is known from Vancouver Island, British Columbia, to Bahfa

Tortugas, Baja California (Behrens, 1991).

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Figure 2.17. A.-C. Cadlinaflavomaculata MacFarland, 1905. A. Jaw rodlets, x 5000. B. Central portion of

radula, x 1000. C. Outer lateral teeth, x 1000. D° Hotoplocamus sp., radular teeth, x 400. t

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l Family Polyceratidae

i Genus Holoplocamus Odhner, 1926 Type Species: Holoplocamuspapposus Odhner, 1926.

i Radular Diagnosis. Body elongate, limaciform, with scattered notal and velar appendages. Jaws reduced. formula: 7-8.2.1.1.2.7-8. Rachidian consisting of paired papillate plates. Inner lateral teeth

hook-shaped. Outer laterals flattened plates. Reproductive system triaulic. Penis armed with hooks.

I Holoplocamus sp.

i Figures 2.14B-C, 2.17D

Material Examined. California, Santa Mafia Basin, off Morro Bay and Purisima Point, Sta. 6, rep.

; II 1 (1); Sta. BRA-6, rep. 1 (1), dissected.

i Description. Preserved animals 0.5-1.5 mm in length. Body elongate with 11-15 spiculate velar

tentacles and 4-6 lateral appendages per side of body (Figure 2.14B,C). Gills 3, unipinnate to bipinnate.

I Mouth Radularflanked formulaby30pairx of7-8.2.1.1.2.7-8. elongate tentacle Rachis s. Labial (Figure cuticle 2.17D) thin, covered coriaceousby, divided without pair wellofdevepapillate loped rodlets. plates.

Innermost lateral teeth small, hook-shaped. Second lateral teeth larger, with simple hook-shaped cusp.

I Outer lateral B teeth simple, quadrangular. iology. Nothing is known about the biology of this species. It has been collected from both hard

and soft substrate stations.

I Remarks. The material differs from all other polycerids described from the Pacific coast of

present

North America. Two species have a somewhat similar radular formula. Laila cockerelli also has similar

radular teeth, except that the second laterals all have a bifid cusp (McDonald, 1984). Issenapacifica (Bergh,

l 1894) has simple, non-spiculate appendages and a rachis with an undivided central plate. The present material has a radula that is almost identical to Holoplocamus papposus (Odhner, 1926), known from the Subantarctic

Atlantic. Holoplocamus papposus has many more lateral appendages and dorsal papillae than the present

I specimens. More material is required to compare this apparently undescribed species with other taxa.

Distribution. Thus far, this species is known only from the Santa Mafia Basin.

I Family Goniodorididae

t Genus Okenia Menke, 1830

Type Species: Doris elegans Bronn, 1826.

I Diagnosis. Body broad to elongate with noted and velar appendages. Jaws usually with chitinous

elements. Radular formula: 1.1.0.1.1. Inner lateral teeth larger than outer laterals. Reproductive system

i Iriaulic. Penis armed with rows of spines.

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Okenia sp. I

Figures 2.14D, 2.16C-D

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Material Examined. California, Santa Maria Basin, off Point Sal, Sta. 35, rep. 3 (1), dissected. I

Description. Preserved specimen (Figure 2.14D) 1 mm in length. Notum with 3-4 papillae present

along either lateral margin of notum. Gills 4, arranged in semicircular fashion. Labial cuticle without I

prominent platelets. Radular formula 18 x 1.1.0.1.1. Inner lateral teeth (Figure 2.16C) large, curved with I

7-8 denticles on inner side of tooth. Outer lateral teeth (Figure 2.16D) narrow, bicuspid.

Biology. Nothing is known about the biology of this species. The single specimen was found in 548

m depth. w

Remarks. Three other species of Okenia have been recorded from the temperate Pacific coast of t1_

North America. Okenia vancouverensis and the introduced O. p/ana both have a more rounded body and II

more elongate appendages than Okenia sp. and O. angelensis. Okenia angelensis has an inner lateral tooth

with more denticles than O. sp. Also the outer lateral tooth is round and arched with a simple hook-shaped Ill

denticle, while that of O. sp. is narrow and bifid. The radutar teeth of O. sp. axe most similar to O. impexa, ii

known from the western Atlantic (Marcus, 1957).

Distribution. Thus far, this species is known only from the single specimen collected from the m

Santa Mafia Basin. II

Family Dendronotidae ,I

Genus Dendronotus Alder and Hancock, 1845

T Ii ype Species: Amphitritefrondosa Ascanius, 1777.

Diagnosis. Animals elongate with numerous branched gills along margins of notum. Rhinophores I

with cup-shaped sheath with 1 or more branched papillae. Velum with 4-16 branched tentacles. Jaws well tl

developed with several row of denticles along masticatory margin. Radula with triangular, denticulate

rachidian. Lateral teeth elongate, denticulate. Reproductive system diaulic. Penis unarmed. I

Dendronotusfrondosus (Aseanius, 1774) /ll

Figures 2.15D, 2.18A I

Amphitritefrondosa Ascanius, 1774:155-158, pl. 5, fig. 2. I1

Dendronotus venustus MacFarland, 1966:271-280, pl. 40, fig. 2, pl. 46, figs. 9-12, pl. 47, figs. 1, 2, pl. 49, II

fig. 6, pl. 50, fig. 3, pl. 52, figs. 3-6.

Material Examined. California, Santa Mafia Basin, off Point Sal, Sta. PJ-1, rep. 3 (1) dissected; I

Sta. PJ-23, rep. 3 (I).

Description. Animals to 50 mm in length. Living animal (Figure 2.15D) translucent gray, white, t

brown, or yellowish, with opaque white or yellowish spots. Gills 4-7 per side of body, highly ramified. tl

Rhinophores with numerous branches on apex of sheath and atleast 1large branch on side of sheath. Velum

with single row of 3-4 highly branched tentacles per side of body. Jaws strong, chitinous, with series of

dentictes along short masticatory margin. Radular formula 24-49 x 7-15.1.7-15. Rachidian teeth (Figure |

2.18A) broad with 8-10 elongate denticles on either side of triangular cusp. Lateral teeth thin, acutely

pointed with 2-6 denticles along outer edge. I

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I Figure 2.18. Radular teeth. A. Dendronotusfrondosus (Ascanius, 1774), x 700. B. Cuthona rolleri Behrens and

Gosliner, 1988, x 1000. C. Tritoniafestiva (Stearns, 1873), x 1000. D. Tritonia diomedea Bergh,

1894, x 700.
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Biology. Dendronotusfrondosus feeds on a wide variety of thecate and athecate hydroids. It appears I

to prefer species of Tubularia and campanularids.

Remarks. Dendronotusfrondosus and D. subramosus MacFarland, 1966 are the only brownish

dendronotids found south of British Columbia. The two are immediately separable by the degree of branching II

of the gills, appendages on the rhinophore sheaths and velar tentacles. All of these structures axe more

highly branched in D. frondosus than in D. subramosus. I

Type Locality and Type Specimens. Norway, type material unknown. U

Distribution. Circumboreal, Europe, Russia, Japan, Pacific coast of North America (Behrens, 1991). I

Family Tritoniidae

a

Genus Tritonia Cuvier, 1798

Type Species: Tritonia hombergi Cuvier, 1803. I

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Diagnosis. Body broad to slender. Numerous branched gills present along margins of notum.

Rhinophores with cup shaped sheath surrounding series of radial lamellae. Velum with 4-30 simple tentacles.

Jaws well developed with several row of denticles along masticatory margin. Rachidian teeth quadrate with

trifid cusp. Lateral teeth curved to elongate, denticulate or smooth. Reproductive system diaulic. Penis

unarmed or armed.

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TdConiafestiva (Stearns, 1873)

Figures 2.15C, 2.18B I

Lateribranchiaeafestiva Stearns, 1873:77-78, fig. 1.

Tritonia reticulata Bergh, 1882:239-250, p. 8, figs. 7-20, pl. 9, figs. 1-12, pl. 10 figs. 1-10. !

Sphaerostoma undulata O'Donoghue, 1924:3-6, pl. 1, figs. 1-4.

Tritoniafestiva: Johnson and Snook, 1927:491, pl. 7, fig. 5. I

Duvauceliafestiva: MacFarland, 1966:218-226, pl. 39,figs. 1-6, pl. 43, figs. 10-19, pl. 44, fig. 2, pl. 45, figs. 7,8. I!

Material Examined. California, Santa Maria Basin, off Point Sat, Sta. PJ12, rep. 1 (1), dissected.

Description. Living animals (Figure 2.15C) up to 30 mm in length. Body translucent white to II

yellowish, with network and circles of opaque white. Notum smooth. Velum with 4-6 filiform tentacles on

either side of frontal veil. Gills 11-15 per side of body. Jaws elongate, chitinous with elongate masticatory

margin, bearing several rows of denticles. Radular formula 32-57 x 15-49.1.1.1.15-49. Rachidian teeth II

trifid (Figure 2.18C). Inner lateral teeth arched with or without series of f'me denticles along outer margin.

Outer lateral teeth all elongate, narrow, curved. I

Biology. Tritonia festiva feeds on pennatulaceans and gorgonians. U

Remarks. Tritoniafestiva is most similar to T. myrakeenae and T. diomedea. It differs externally

by its translucent white color with opaque white network of lines and circles. It has 4-6 velar tentacles per I_tll

side, while T. diomedea has 5-15 and T. myrakeenae has 3-4. Tritoniafestiva and T. diomedea have far

more ramified gills than T. myrakeenae. The notum of T. myrakeenae and T.festiva are smooth, while that

of T. diomedea is covered with low tubercles.

'W

Type Locality and Type Specimens. Pt. Pifios, Monterey, California. Type material unknown.

Distribution. Japan, Kachemak Bay, Alaska to Coronados Islands, Baja California (Behrens, 1991). i

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Tritonia diomedea Bergh, 1894

Figures 2.14F, 2.18D

I Tritonia diomedea Berg,h, 1894:146-150, pl. 2. figs. 3, 10, 11, pl. 3, figs. 6-10, pl. 4, figs. 1, 5.

Duvaucelia gilberti MacFarland, 1966:235-243, pl. 30, figs. 1, 2, pI. 43, figs. 27-36, pl. 44, fig. 5, pl. 45, fig. 6.

I Tritonia exsulans Bergh, 1894:150-152, pl.3, figs. 11, 12, pl. 4, fig. 6.

Material Examined. California, Santa Maria Basin, off Purisima Point, Sta. BRA 13, rep. 4 (1),

I dissected.

Description. Living animals up to 215 mm in length. Living animals white, pink or red in color.

Notum with numerous low tubercles. Velum (Figure 2.14F) with 5-15 ffliform tentacles on either side of the

I frontal veil. Gills 11-15 side of body. Jaws elongate, chitinous with elongate masticatory margin, bearing

per

several rows of denticles. Radular formula 29-69 x 21-95.1.1.1.21-95. Rachidian teeth trifid (Figure 2.18D).

Inner lateral teeth arched, with or without series of coarse denticles along outer margin. Outer lateral teeth

I short, curved. Inner teeth more elongate towards outer edge.

Biology. Tritonia diomedea appears to feed exclusively on pennatulaceans at subtidal depths.

I from theRemotherarks. largeTrito northniaPacific diomedea tritoniid, growsTochuina much larger tetraquetra than other(Pallas, sympatric 1788),species in having of Triless toniapronounced . It differs

notal tubercles and far fewer gills. There are significant radular and reproductive differences as well.

I specimensTypeunkno Locality wn. and Type Specimens. San Diego Bay and Santa Barbara Island, California. Type

Distribution. Aleutian Islands, Alaska to Bay of Panama; Manatee Bay, Florida (Behrens, 1991).

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Family Arminidae

I Genus Arm/ha Rafmesque, 1814

i Type Species: Armina tigrina Rafinesque, 1814.

Diagnosis. Notum curved with series of longitudinal ridges. Rhinophores close together, anterior to

edge of notum, with radially arranged lamellae. Jaws well developed with several rows of denticles along

I Rachidian teeth Lateral teeth curved, with without masticatory margin. multicuspidate, quadrangular. or

denticles.

I Armina californica (Cooper, 1863)

Figures 2.19A, 2.20A-B

I Pleurophyllidia californica Cooper, 1863:203-204.

Armina californica: MacFarland, 1966:198-206, pl. 38, figs. 1-6, pl. 43, figs. 37-44, pl. 44, figs. 6, 7.

I Material Examined. California, Santa Maria Basin, off Point Sal, Sta. PJ-2, rep. 1 (1), dissected.

Description. Living animals (Figure 2.19A) up to 70 mm. Color dark gray to brown with numerous

pink to white longitudinal ridges. Rhinophores anterior to edge of notum, with longitudinal

folds. Gills

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numerous, ventral, situated between edge of notum and foot. Jaws chitinous, with 4 rows of denticles

(Figure 2.20A) along elongate masticatory margin. Radular formula 25-52 x 33-80.1.1.1.33-80. Rachidian

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Figure 2.19. Living animals. A. Armina californica (Cooper, 1863). B. Cuthona rolleri Behrens and Gosliner, i

1988. C. AeolidielIa chromosoma (Cockerell and Eliot, 1905).

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I teeth (Figure 2.20B) acutely pointed with prominent medial cusp and 3-4 lateral denticles per side. Inner

lateral tooth thick, curved. Outer lateral teeth hook-shaped with small denticle on outer side of inner 21

teeth. Outer Biolo12gy.teeth A rmina simple, californica curved,feeds withoexclusively ut additionalondepennatulaceans nticle. of the generaRenilla and Stylatula.

It is found from the low intertidal to at least 200 m depth.

I Remarks. Armina californica is the only arminid known from the temperate Pacific coast of North America. In the southern part of its range, it is sympatric with the arminid, Histiomena convolvula, which

lacks longitudinal ridges of the notum and is more brightly colored.

I Type Locality and Type Specimens. San Diego Bay, California. Type material unknown.

Distribution. Armina californica is known from Kayak Island, Alaska to the Gulf of Panama

i (Behrens, 1991).

Family Tergipedidae ,!

Genus Cuthona Alder and Hancock, 1855

I Type Species: Aeolis nana (Alder and Hancock, 1842)

Diagnosis. Body slender to broad. Anterior end of foot generally rounded, but rarely with elongate

I tentacles. anterodorsaltyOraltotentacles first post-cardiac short. Rhinophores ceratal row. commonly Jaws well developed smooth butwithrarely singleannulate. row of denticles Anus situated along

masticatory margin. Radular uniseriate with cuspidate teeth. Reproductive system diaulic. Penial gland

i present. Penis unarmed or armed with tubular stylet.

Cuthona rolleri Behrens and Gosliner, 1988

I Figures 2.18B, 2.19B

Cuthona rolleri Behrens and Gosliner, 1988:262-266, figs. 1-4.

I Material Examined. California, Santa Maria Basin, off Point Sal, Sta. R-7, rep. 1 (1), dissected.

Description. Living animals (Figure 2.19B) up to 15 mm in length. Color translucent white. Ceratal

I cores pink. Body dorsoventrally Rhinophores smooth, separated.

salmon flattened. Foot wide. well Cerata

large, arranged in longitudinal rows well elevated from notum. Jaws thick, chitinous with single row of

denticles on the masticatory margin. Radular formula 21-31 x 0.1.0. Radular teeth (Figure 2.18B) with

I large, triangular primary and 5-7 denticles flanking either side.

cusp

Biology. Little is known about the biology of this species. Most specimens have been found on

intertidal mud flats. The present specimen, collected from 575 m depth, represents the first subtidal record

t of this species.

Remarks. Cuthona rolleri is immediately identifiable by its dorso-ventrally flattened body with

I elongateTcerata. ype Locality and Type Specimens. Morro Bay, California; holotype, (CASIZ 064894), paratypes

(CASIZ 064895, 064896).

I Distribution. Cuthona rolleri was previously known from Piedras Blancas to Morro Bay, San Luis Obispo County, California. The present specimen extends the range southward to Point Sal, Santa Barbara

County. It also considerably extends the bathymetric range of the species to 575 m depth.

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Figure 2.20. A., B. Armina californica (Cooper, 1863). A. Jaw rodlets, x 1000. B. Central radular teeth, x 700. C. I

AeolidieIla chromosoma (Cockerell and Eliot, 1905), radular teeth, x 500. D. Cerberilla mosslandica

McDonald and Nybakken, 1975, radular teeth, x 700.

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I Family Aeolidiidae

I Genus Aeolidiella Bergh, 1867 Type Species: AeoIis glauca Alder and Hancock, 1848.

I Diagnosis. Body broad. Anterior end of foot rounded or with elongate tentacles. Oral tentacles long. Rhinophores commonly smooth but rarely annulate or perfoliate. Anus situated posterior to first

post-cardiac ceratal row. Jaws well developed with single row of denticles along masticatory margin or with

i smooth system diaulic. masficatory Penismargi unarmed. n. Radula uniseriate with pectinate teeth, with medial indentation. Reproductive

I (Cockerell Eliot, 1905)

Aeolidiella chrolfto$oma and

Figures 2.19C, 2.20C

I Spurilla chromosoma Cockerell and Eliot, 1905:51-52.

AeolidieUa chromosoma: Gosliner, 1985:257, fig. 15B.

I Material Examined. California, Santa Mafia Basin, off Purisima Point, Sta. BRA 16, rep. 1 (1),

dissected.

I • yellowishDescription with orange. Land ivingopaque animalswhite(Figure markings 2.19C)on upheadtoand25 notum. mm in length. Cerata gray Colotor translucent brown, withwhit opaque e to

white spots. Rhinophores bulbous, with diagonally slanted lamellae. Foot broad. Anterior comers of foot

I elsingle ongatspecimen e, tentacular examined, . Jaws thi21-24 ck chitin in previously ous, with smrecorded ooth masti matecato rial.ryRadular border. teeth Radular (Figure formula 2.20C)19 xpectinate, 0.1.0 in

arched with up to 30 denticles on either side of triangular medial denticle.

i identified.Biology. Aeolidiella chromosoma feeds on sea anemones, though specific prey have not been

Remarks. Aeolidiella chromosoma is most similar to A. olivae MacFarland, 1966. The two differ

I in their color. Aeolidiella olivae has reddish rhinophores and cerata, while those ofA. chromosoma are drab

gray or brown. The two differ slightly in their radular morphology with A. olivae having fewer denticles on

the teeth.

! Type Locality and Type Specimens. San Pedro, California. Type material unknown.

Distribution. This species is found from Morro Bay to the Gulf of California (Behrens, 1991).

I Genus Cerberilla Bergh, 1873

I Type Species: CerberiUa longicirra Bergh, 1873.

Diagnosis. Body broad. Anterior end of foot with elongate tentacles. Oral tentacles long. Rhinophores

t smooth, short. Anus situated posterior to first post-cardiac ceratal row. Jaws well developed with single row

of denticles along masticatory margin or with smooth masticatory margin. Radula uniseriate with pectinate

teeth often with unevenly elongate denticles. Reproductive system diaulic. Penis unarmed.

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Cerberilla mosslandica McDonald and Nybakken, 1975 I

Figures 2.14E, 2.20D

B

Cerberilla mosslandica McDonald and Nybakken, 1975:378-382, figs. 1, 2. 1

Material Examined. California, Santa Mafia Basin, off Point San Luis and Point Sal, Sta. R-3, rep. ,I

1 (1); Sta. R-8, rep. 3 (2), one dissected. I

Description. Living animals (Figure 2.14E) up to 9 mm in length. Body translucent white, encrusted

with reddish brown pigment. Cerata brownish with white apices. Cerata congested in many linear rows. i

Rhinophores smooth. Anterior margins of foot elongate. Jaws thin, coriaceous, with smooth masticatory I

border. Radular formula 10 x 0.1.0. Radular teeth (Figure 2.201)) evenly curved with elongate, pectinate

denticles.Biology I . Members of the genus Cerberilla feed on cerianthid anemones and other sand and mud

dwelling coelenterates. They burrow below the surface of sand and mud. McDonald and Nybakken (1975)

found this species at 17-70 m depth. The present material was found in slightly deeper water (90 m depth). I

I

Remarks. Two other eastern Pacific Aeolidiidae are similar to Cerberilla mosslandica in having

smooth rhinophores. Both AeoIidieIla indica and Aeolidia papillosa are known only from shallow rocky

habitats. CerberiIla mosslandica differs from both of these species in having elongate anterior margins of I

the foot. II

Type Locality and Type Specimens. Off Moss Landing, Monterey Bay, California; holotype, It,

(CASIZ 020365); paratypes (CASIZ 020366). I

Distribution. Monterey Bay to La Jolla Submarine Canyon, California (Behrens, 1991).

Literature Cited I

Ill

Abbott, R. T. 1974. American Seashells. Van Norstrand Reinhold, New York, NY, 663 pp. I

Ascanius, P. 1774. Beskrivelse over en Norske sneppe og et sodyr. Det Kongelige Norske Videnskabers im

Selskabs Skrifter 5: 153-158. |

Baird, W. 1863. Descriptions of some new species of shells, collected at Vancouver Island, and in British

Columbia by J. K. Lord, esq., naturalist to the British North-American Boundary commission, in I_

the years 1858-1862. Proceedings of the Zoological Society of London 1863: 66-70. I

Behrens, D. 1991. Pacific coast nudibranchs. Second edition. Sea Challengers: Monterey, CA, 107pp. I

Bergh, R. 1882. Beitr_ige zur Kennmiss der Aeolidiaden. 7. Verhandlungen der k. k. Zoolisch-botanischen II

Gesellschaft in Wien 32:7-84.

Bergh, R. 1893a. Die Gruppe der Doridiiden. Mittheilungen aus der Zoologischen Station zu Neapel zugleich I

ein Repertorium fur Mittelmeerkunde 11:107-135.

Bergh, R. 1893b. Die Gattung Gastropteron. Zootogische Jahrbucher. Abtheilung ffirAnatomie und Ontogenie i

der Thiere 7:281-308. I

Bergh, R. 1894. Die Opisthobranchien. Reports on the dredging operations off the west coast of Cen_al Im

America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of I

Alexander Agassiz, carried on by the U.S. Fish Commission steamer "Albatross", during 1891,

Lieut. Commander Z. L. Tanner, U.S.N. commanding. Bulletin of the Museum of Comparative ml

Zoology Harvard University 25 (1):125-235. I

210 I
!

I Burch, J. Q. 1944. Minutes of the Conchological Club of Southern California 37:17-18

Carpenter, P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca

oScience f the west 1863:coas 517-686. t of North America. Report of the British Association for the Advancement of

I Cooper, J. G. 1863. New genera and species of California Mollusca. Proceedings of the California Academy of Sciences 2:202-207.

Chamberlain, R. and Behrens, D. 1980. Deep water collections of opisthobranchs in central California.

I Veliger 22: 282-285.

Cockerell, T. D. and C. Eliot 1905. Notes on a collection of California nudibranches. Journal of Malacology

i 12:31-53. Dall, W. H. 1871. Descriptions of sixty new forms of mollusks from the west coast of North America and

the North Pacific Ocean, with notes on others already described. American Journal of Conchology

I 7: 93-160.

Dall, W. H. 1894. Description of a new species of Doridium from Puget Sound. Nautilus 8 (7):73-74.

I Dall, W. H. 1900. A new species of Pleurobranchus from California. Nautilus 14(8):92-93.

Dall, W. H. 1908. Descriptions of new species of mollusks from the Pacific coast of the United States, with

notes on other mollusks from the same region. Proceedings of the United States National Museum

I 34 (1610): 245-257.

Dall, W. H. 1919. Descriptions of new species of Mollusca from the North Pacific Ocean in the collection of

I the United States National Museum. Proceedings of the United States National Museum 56:293-371.

Dall, W. H. 1925. A new species of Acteocina fxom British Columbia. Nautilus 39(1):25-26.

I Gosliner,Nautilu T. M. 1979. A review of the systematics of Cylichnella Gabb (Opisthobranchia: Scaphandridae). s 93: 85-92.

i Gosliner, T. M. 1980. Systematics and phylogeny of the Aglajidae (Opisthobranchia: Mollusca) Zoological Journal of the Lirmean Society 68: 325-360.

Gosliner, T. M. 1981. Origins and relationships of primitive members of the Opisthobranchia (Mollusca:

I Gastropoda). Biological Journal of the Linnean Society 16:197-225.

Gosliner, T. M. 1984. Two new species of Gastropteron (Gastropoda: Opisthobranchia) from southern

I Gosliner,Africa T . Publications of the Seto Marine Biological Laboratory 29: 231-247.

. M. 1985. The aeolid nudibranch family Aeolidiidae (Gastropoda: Opisthobranchia) from tropical

southern Africa. Annals of the South African museum 95(6):233-267.

I Gosliner, T. M. 1991. Morphological parallelism in opisthobranch gastropods. Malacologia 32:313-327.

Gosliner, T. M. 1994. Gastropoda: Opsithobranchia, pp. 253-355. In F. Harrison and A. Kohn (eds.),

i Microscopic Anatomy of Invertebrates. Volume 5. Mollusca. John Wiley and Sons, New York.

Gosliner, T. M. and H. Bertsch. 1988. A review of the genus Berthella (Opisthobranchia: Notaspidea) from

the Pacific coast of North America. Veliger 31: 46-67.

I Gosliner, T. M. and G. C. Williams, 1972. A new species of Chelidonura from Bahia San Carlos, Gulf of

California, with a synonymy in the family Aglajidae. Veliger 14: 424-436.

!

i 211
!

Gould, A. A. 1852. Mollusca and shells. United States Exploring Expedition during the years 1838, 1839, I

1840, 1841, 1842, under the command of Charles Wilkes, U.S.N. 12 (Atlas): 1-510.

Haszprunar, G. 1988. On the origins and evolution of major gastropod groups, with special reference to the I

Streptoneura. Journal of Molluscan Studies 54: 367-441. J

Johnson, M. E. and H. J. Snook. 1927. Seashore animals of the Pacific coast. Dover Publishers, New York,

N.Y., 659 pp. il

Keen, A. M. 1971. Sea shells of tropical West America. Stanford University Press, Stanford, CA, 1064 pp.

n

Lance, J. R. 1966. New distributional records of some northeastern Pacific Opisthobranchiata (Mollusca: I

Gastropoda), with descriptions of two new species. Veliger 9:69-81.

Lee, R. S. and N. R. Foster. 1985. A distributional list with range extensions of the opisthobranch gastropods I

of Alaska. Veliger 27: 440-448.

MacFarland, F. M. 1966. Studies of opisthobranchiate mollusks of the Pacific coast of North America. I

Memoirs of the California Academy of Sciences 6: 1-546. II

Marcus, Er. 1957. On Opisthobranchia from Brazil. 2. Journal of the Linnean Society of London 43 (292):

390-486. I

Marcus, Er. 1961. Opisthobranch mollusks from California. Veliger 3 (supplement): 1-85.

Marcus, Ev. 1972. Acteonidae. On some Acteonidae (Gastropoda, Opisthobranchia). Papeis Avulsos I

Zoologica S_o Paulo 25: 167-188.

Marcus, Ev. 1977. On the genus Tornatina and related forms. Journal of Molluscan Studies (suppl. 2): 1-35. I

II Marcus, Ev. and Er. Marcus. 1969. Opisthobranchian and lamellarian gastropods collected by the Vema.

American Museum Novitates 2368: 1-33.

m

McDonald, G. R. 1983. A review of the nudibranchs of the California coast. Malacologia 24:1 I4-276. I

McDonald, G. R. and J. W. Nybakken. 1975. Cerberilla mosslandica, a new eolid nudibranch from Monterey

Bay, California. Veliger 17: 378-382. I

Millen, S. V. and T. M. Gosliner. 1985. Four new species of the dorid nudibranchs (Mollusca:

Opisthobranchia) belonging to the genusAldisa, with a revision of the genus. Zoological Journal of I

the Lirmean Society 84:195-233. II

Odhner, N. 1926. Die Opisthobmnchien. Further results of the Swedish Antarctic Expedition 1901-1903

under the direction of Dr. Otto Nordenskjold. 2(1): 1-100. Iw

O' Donoghue, C. 1924. Notes on the nudibranciate Mollusca from the Vancouver Island region. 4. Transactions

of the Royal Canadian Institute 15:1-33.

II Pilsbry, H. A. 1895. Manual of Conchology. Acanthochitidae, Cryptoplacidue and appendix. Tectibranchiata.

15(60): 181-436.

Pilbsry, H. A. 1896. Manual of Conchology. Philinidae, Gastropteridae, Aglajidae 16(61): 1-48. I

Powell, 1951. Antarctic and subantaretic Mollusca, Pelecypoda and Gastropoda. Discovery Reports 26:

47-196. I'

Rudman, W. 1974. A comparison of Chelidonura, Navanax and Aglaja with other genera of the Aglajidae

(Opisthobranchia: Gastropoda). Zoological Journal of the Linnean Society 54:185-212. I

212 I
!

i Steams, R. 1873. Descriptions of a new genus and two new species of nudibranchiate mollusks from the

coast of California. Proceedings of the California Academy of Sciences 5:77-78.

I Strong, A. M. 1921. Notes on the genus Acteocina. Nautilus 35: 44-48.

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee Expedition. part 2. Wissenshaftliche ergebnissee der

i Deutschen Tiefsee Expedition auf dem Dampfer "Valdivia" 1898-1899. 17:38-382. Thompson, T. E. 1976. Biology of opisthobranch molluscs. Volume 1. Ray Society: London.

I Tokioka,Publications T. and Baba,ofK. 1964. Four new species and a new genus of the family Gastropteridae from Japan. the Seto Marine Biological Laboratory 12:201-229.

Willan, R. 1987. Phylogenetic systematics of the Notaspidea (Opisthobranchia) with reappraisal of families

n and genera. American Malacological Bulletin. 5: 215-241.

Willett, G. 1928. Notes on some Pacific coast Acteocinas, with description of one new subspecies. Nautilus

i Willett, 42: 37-38. G. 1944. New species of mollusks from Redondo, California. Bulletin Southern California Academy

of Sciences 43: 71-73.

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i 213
Appendix I

Lists and Maps of Stations 1

_ble A.1. Position of soft-subs_ stations Ken d_g _e Phase I Reconn_ssance. I

Station Latitude Longitude Depth

(m) I

1 35°27.86'N 121°05.33_ 98

2 35°27.70'N 121°06.52_ 2_ J

II 3 35°27.07_ 121°10.2_ 291

4 35%6.56_ 121°14.93_ 393

5 35°25.77_ 121°21.69_ 585

6 35°20.88_ 120°59.62_ 109 J

II 7 35°20.65_ 121°02.57_ 197

8 35°20._ 121°06.58_ 308

9 35°19.48_ 121°I0.06"W 398 I1

10 35°18.28"N 121°18.65_ 591 1

11 35°17.8_N 121°22.13_ 690 U

12 35°t5.03'N 1_°57.3 I'W 98

13 35°14.54'N 1_°59.77_ 197 I

14 35° l4.15'N 121°02._ 299 •

Ill 15 35° 13.98'N 121°_.54_ 393

16 35°12.23'N 121°16.29_ 591

17 35°ll.61'N 121°22.55_ 654

II

18 35°_.08'N 120°56.55_ 197

19 35°08.93_ 120°59.66_ 296

20 35° 15.72_ 121°_.68_ 396

21 35°06.11'N 120°_.82_ 49 •

22 35°05.85_ 120°50.23_ n

23 35°05.6_ 120°55.18_ 195

25 35°05.07"N 121°_.75_ 390

26 35°_.38_ 121°15.9_ 590 •

27 35°_.3_ 121°19.27_ 611

28 35°_.22_ 121°19.65_ 603

30 M°54.1 _N 1_°47.07_ 98

II 31 34°53.76'N 1_°52.96'W 200

32 34°53.56'N 1_°56.8 I'W 297

33 34°53.43'N 1_°59.66'W 396

34 34°53.15'N 121°_.4_ 492 I

II 35 34°52.96_ 121°10.3_ 548

36 34°52.77_ 121°15.37_ 492

38 34°49.81_ 1_°52.66_ 197

39 34°49.53_ 120°56.85_ 294 J

40 34°49.24_ 121°_.81_ 392 l

41 34°48.35_ 121°19.14_ 495

42 34°48._ 120°47.5_ 1 i

43 34°46.59_ 1_°52.92_ 197 •

45 34°_.91_ 120°59.59_ 395

46 M°41.22_ 121°13.56_ 597

47 M°41.9_ 121°10.81_ 378 I

I

48 M°45.11 _ 120°52.85_ 196

49 34°45.03"N 120°56.31 _ 290

50 34°37.8_ 121°01.66_ 591

52 34°39.56'N 120°47.64_ 98

II 53 34°37.6_N 120°50.38_ 196

54 34°36.57'N 120°52.02_ 396

55 34°33.66'N 120°56.31_ 590

56 34°30.32'N 121°01.02_ _ 1

214 I
Table A.1 (Continued)

Station Latitude Longitude Depth

(m)

58 34°34.35'N 120°45.18'W 99

59 34°33.65'N 120°47.18'W 216

60 34°33.25'N 120°48.34"W 275

61 34o33.01'N 120°48.89"W 345

62 34°30A6'N 120°52.13'W 582

63 34o26.29'N 120°58.08"W 930

64 34o33A5'N 120°40.90"W 59

65 34°31.27'N 120°43.27"W 107

66 34o30.46'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 34o22.88'N 120°54.20'W 927

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 34o28.21"N 120°36.80"W 98

74 34°26.84"N 120°38.6 I'W 201

75 34o26.08"N 120°39.65"W 293

76 34o25.59'N 120°40.98"W 387

77 34°22.62"N 120°44.02'W 578

78 34° 18.78'N 120°49.30qN 762

79 34°24.12"N 120°28.32'W 98

80 34o22.86"N 120°28.34"W 196

81 34°21.26"N 120%8.83'W 294

82 34°18.7 I'N 120°29.55'W 394

83 34o17.20"N 120o30.20'W 444

84 34° 13.54"N 120°31.19"W 394

85 34°25.88"N 120° 16.3I'W 113

86 34°24.45'N 120° 17.02'W 197

87 34°21.60'N 120° 17.11'W 299

88 34o17.89"N 120°16.86'W 393

89 34o13.79"N 120°16.56"W 471

90 34°09.44'N 120°16.30'W 375

91 34°11.73'N 120O07.43'W 540

92 34O08.70'N 120O07.50'W 444

93 34°07.63'N 120°07.5 I'W 357

96 34°22.91'N 120°05.42'W 296

94 34°24.54'N 120O05.47"W 96

95 34°23.70'N 120°05.47"W 198

97 34°22.28'N 120°05.49'W 393

98 34°12.87'N 120°05.59"W 561

99 34°11.22"N 120°05.86'W 540

100 34O08.67'N 120°05.50"W 443

101 34°07.5 I'N 120°05.65'W 357

102 34o59.71'N 120o48.22'W 99

103 34o59.63"N 120°53.56"W 197

104 34o59.45"N 120°56.49'W 294

105 34o59.23'N 120°59.60"W 392

106 34°58.95"N 121°04.42"W 492

107 34o58.65'N 121°15.08'W 573

108 34o58.21'N 121°17.88'W 492

Note: Sample labels from the SoR-subslrate stations have several identification codes which include a station number, sample

type, replicate number, and analysis type. These are as follows: 001 to 200 = the range of station numbers; BSS = Benthic

Sediment Single (i.e., a non-replicated station); BSR = Benthic Sediment Replicate (three replicates taken at this station); BSV =

Benthic Sediment Variance (subsamples); 01-09 = replicate numbers; TX = a taxonomy sample. Sample labels having the

designation BRA, represents a sample from rocks taken as part of the hard bottom survey.

215
1

",2 , I

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I ,7 6

9 • O •

10 • ,, m

II • ", 1

• 14

• •

17 16 15 " 1

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19 "IS 1

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l''[ _ 22 21 l

27 26 _- "• • •

@ O _ iLl l

2,8 104 103 ! I_2

108 107 106 • • • © • I

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36 35 34 R9 • 32 31:: o o • ,i O •

• • g 33 o

40 39 3S 0

41 • • g

• _• •

45• 49• O_• S oR4 1

7.000 46 47 :l, _

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'", • _i.59_ • 1

'"--, 56 61 • ;' 65

"'................... • ..... 67 :" • 1

:' 68 _•170 73

" ........ . ......... • .-,.-... •- 94 n '. . . . 1

.... l O _ 097

i o82 088 1

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089

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91 •

O_

-- 34° 10'N

. KEY o_ 92 • olCX)

93 • • 101 l

I

Figure A.1. Map showing location of softsubstrate stationsfrom Phase I and Phase II MMS Reconnaissance 1

and Monitoring Programs.

I

I

216 l
!

I Table A.2. Location of soft-subslrate stations taken during the Phase lI Monitoring Program.

I Station Latitude Longitude Depth (m)

I R- R-21 35°05 35°05..8350'

'NN 112020o°53.4( 49.16Y'WW 161 91

R-3 35°05.30"N 121°00.90'W 409

R-4 34°43.01'N 120°47.39"W 92

I R-5 R-6 3434°°42.69 41.40''NN 120 120°°50.83"W 57.90"W 4154 10

R-7 34°52.90'N 121o10.30"W 565

I R-8 34°55.30"N 120°45.87"W 90

R-9 34053.68"N 120059.12"W 410

PJ- 1 34°55.79"N 120049.9 I'W 145

PJ-2 34°55.32'N 120°49.59"W 142

I PJ-3 34°56.26'N 120°49.58'W 138 ! PJ-4 34°56.26'N 120°50.24'W 150

PJ-5 34°55.32'N 120°50.24'W I52

PJ-6 34°54.71"N 120°49.91'W 148

I PPJ-7 J-8 34°55.7 34°56.879"

'NN 12120°4 0°48.6 9.90'IW'W 123 142

PJ-9 34°55.79"N 120°51.23'W 169

PJ- 10 34°53.63'N 120°49.9 I'W 147

I PPJ-J- 1121 34°57 34°55.58 .95'

'NN 12120°49.9 0°49.9 II'

'WW 136 145

PJ-13 34°56.01'N 120°49.91'W 144

I P J- 14 34°55.79'N 120°49.26'W 134

PJ-15 34°55.79"N 120°50.57'W 155

PJ- 16 34°55.03'N 120°48.99"W 130

PJ-17 34°56.56'N 120°48.98"W 126

J PJ-18 34°56.56'N 120°50.84'W 158

II 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°01.23'N 120°51.15'W 143

I PPJ-22 J-23 34°55.25 34°56.33'

'NN 120°49.93 120°49.90""WW 143 143

I Table A.3. Sampling dales of MMS Phase II Monitoring Program.

i Cruise Date 1-I October 1986

1-2 June 1987

I 1-3 May 1987

1-4 July 1987

2-1 October 1987

2-2 January 1988

. 2-3 May 1988

3-1 October 1988

3-2 May 1988

!

i 217
1

Table A_4. MMS Phase I - Locations of hard-subs_a_ _s_ l 1

Station Berg End Depth •

Latitude Longitude Latitude Longitude (m) 1

1

1 A/B 34°24.454"N 120°01.876"W 34°24.464'N 1200001878"W 69-73.5 1

1

1 CJD 34°24,076_N 120°00.443%V 34°24. 184'N 120°01,480_ 73.5-78

2 A/B 34° 11.3772q 120029.318_W 34° 11.28fin 120°28,774%V 110-126

2 C_ 34° 10.984"N 120°28.094W¢ 34010.780'N I20°27.554_ 120-123 1

4 A/B 34°27.539'N 120°40.364%V 34°28.162'N 120°40l 189%V 168-237

6 A/B 34°30.246'N 120°35.555"W 54-63 1

6 C_ 34°30".421 q_ 120°341315_W 54-63

13 A/B 34°42'.570'N 120°47.899'W 34°42,107q_ 120°48,253%V 92-100 ll

]3 CJD 34°42,556'N 120°48.147W¢ 34°421974'N 120°47,424W¢ 88.5-100.5 •

14 A/B 34°43.589'N 120°49.093'W 34°42.826'N 120°48,370%V 96-105 I

14 C/D 34°43,244'N t20°49,406'W 34°42,893'N 120°48.822%V 105-I 17

16 A/B 34°46,544'N I20°50,197'W 34°45.912'N 120°49,726W¢ 91.5-123 Ill

17 A/B 34°49,382'N 120°50.768%V 34°49.600q_ 120°50,688%V 160.5-168 II,

19 AfB 34°47,833'N 120°51,425W¢ 34°47.0972q 120°50,793%V 148.5- I77 1

20 A/B 34°46,470'N 120°50,289%V 34°46.1402q 120°49.885%V 90-130.5

21 AJB 34°47,335'N 120°45,903'W 34°47.5482q 120°46.123%V 75-90

22 A/B 34°50.365'N 120°48.22l'W 34°50.990"N 120°48.365'W 114-115.5 1

23 AJB 34°49,868'N 120°47,393'W 34°50.0032q 120°47.480%V 93-102 1 25 AIB 35°05,662'N 120°47,562%V 35°06.0362q 120°47.652%V 64.5-72

26 C_ 35°11,586'N 120°55.556%V 35°11.555'N 120°55.233%V 108-111

27 A/B 35°20.906'N 120°591657_q¢ 35°21.0352q 120°59.603%V 96-126 •

28 A/B 35°21.539'N 120°591641_ 35°21.8672q 120°59.299%V 96-105 l

29 A/B 35°27,864_ 121°05,331%V 35°27.805_N 121°05.277W¢ 102-106.5

I

1

Table A_, MMS Phase H - Locations of hard-subs_a_ photosurvey stations.

I

Station Latitude Longitude Depth

(m)

I

PH-E 34°30.26'N 120°42.76"W 119 _

PH-F 34°30.81'N 120°42.36'W 11)5

PH-I 34°29.96'N 120°41.68'W 107 I

PH-J 34°29.82'N 120041.82"W 117

PH-K 34°29.37'N 120°42.26'W 160

PH-N 34°29.21'N 120°42.05W¢ 166 •

PH-R 34o29.11 _q 120°42.67W¢ 213

PH-U 34°31.48'N 120°43.51W¢ 113

PH-W 34o31.52'N 120°45.86W¢ 195

I

I

1

218 I
I ., •

26

I ". 25

I

,' 17

i • 22

!16

eO • 21

| i,_ ,+. ,.o

:: • 14

" • 13

i 34o32, N "\_

I +°'+ : l, _

• • • ..... o;

I P++-w _. .....,, • 2 .

...................... 200 ................................

PH'EPH°? Ill PH-,

.._o_o,N

I ,3.7.I,,H.,

i 34o28,N 120 °44'WI

I

I Figure A.2 Map showing locations of hard substrate stations from Phase I and Phase I/MMS Reconnaissance

and Monitoring Programs.

I

i 219
IlIIIlIllIIIlillIIl
I

I

!

I

I

I

The Department of the Interior Mission

I As the Nation's conservation the Department of the Interior has responsibility

principal agency,

for most of our nationally owned public lands and natural resources. This includes fostering

sound use of our land and water resources; protecting our fish, wildlife, and biological diversity;

i preserving the environmental and cultural values of our national parks and historical places;

and providing for the enjoyment of life through outdoor recreation. The Department assesses

our energy and mineral resources and works to ensure that their development is in the best

interests of all our people by encouraging stewardship and citizen participation in their care.

The Department also has a major responsibility for American Indian reservation communities

I and for people who live in island territories under U.S. administration.

The Minerals Management Service Mission

I _,__'_ As primaarybureau respoofnsithebilitiDeespartm are toentmanage of the Interior, the mineral the resourc MineralsesManagement located on theService's Nation's(MMS) Outer

Continental Shelf (OCS), collect revenue from the Federal OCS and onshore Federal and Indian

lands, and distribute those revenues. I

_I Moreover, in working to meet its responsibilities, the Offshore Minerals Management Program

administers the OCS competitive Ieasing program and oversees the safe and environmentally

sound exploration and production of our Nation's offshore natural gas, oil and other mineral

I resources. The MMS Royalty Management Program meets its responsibilities by ensuring the efficient, timely and accurate collection and disbursement of revenue from mineral leasing and

production due to Indiantribes and allottees, States and the U.S. Treasury.

I The MMS strives to fulfill its responsibilities through the general guiding principles of: (1) being responsive to the public's concerns and interests by maintaining a dialogue with all potentially

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 deve|opment and environmental protection.

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