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•I OCS Study

i MMS 96-0044

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I TAXONOMIC ATLAS OF THE BENTmC FAUNA OF

THE SANTA MARIA BASIN AND

] WESTERN SANTA BARBARA CHANNEL

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1 FINAL REPORT

Volume 3 of 14

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The Cnidaria

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I Edited by: James A. Blake

Paul H. Scott

I Submitted by:

i Science 10260 Campus ApplicatPoint ions InDrive ternational Corporation

San Diego, California 92121

I For:

U.S. Department of the Interior

/ Minerals Management Service

= Pacific OCS Region

770 Paseo Camarillo

Camarillo, California 93010

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

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Minerals Management Service

I Pacific 0CS Region July 1996
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DISCLAIMER I

TShis e reporthas beenreviewedby the PacificOuterContinentalShelfRegion,MineralsMangement I rvice, U.S. Departmentof the Interiorand approvedfor publication.The opinions,findings,

conclusionsorrecommendationsexpressedin this reportare thoseof the authors,and do not

nnot ececonsitiut ssarilyreefleendorsement ctthe view oforthreecomm Mineraelsndation Managfor ement use.SeTrvi hiscerepo . Menrtthas iononot f trabdeeenan mes editeddoes for I

conformitywith MineralsManagementServiceeditorialstandards.

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TAXONOMICDISCLAIMER

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

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

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i PROJECT ORGANIZATION

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i PROGRAM MANAGER

DR. ANDREWL. LIss_wa_

i Science Applications International Corporation

10260 Campus Point Drive

I San Diego, California 92121

I DEPUTrPROGRAM ANAGER

I DR.JAMESA. BLAKE

ENSR, 89 Water Street

I Woods Hole, Massachusetts 02543

I PROJECTTECHNICALOFFICER

I MR. F_r_x J. MANAGO

U.S. Department of the Interior

Minerals Management 770 PaseoServiCamarillo ce, Pacific OCS Region

I Camarillo, California 93010

i CONTgmtrrlNGAtrrHORS

DR. DAFm,m FAUTIN

I MP.. DR. FJoItN . G. I_mmENK HoctmFa_OV

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ACCESS NUMBER: 30484.3

TECHNICAL SUMMARY i

STUDY TITLE: TaxonomicAtlasof the SantaMaria BasinandWestem SantaBarbaraChannel. d

I REPORT TITLE: TaxonomicAtlasofthe Santa MariaBasinand WesternSanta BarbaraChannel.Volume

3. The Cnidaria.

I

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

SPONSORINGOCS REGION: Pacific. i

i

APPLICABLEPLANNINGAREA: SouthernCalifornia.

FISCALYEARS OF PROJECTFUNDING: 1990; 1991; 1992;1993; 1994; 1995. i

COMPLETION DATEOF REPORT: April 1996. m

COSTS: FY 1989,$229,013; FY1990,$150,000; FY 1991$150,000;FY 19935150,000; FY 1995$45,371. i

CUMULATIVEPROJECT COST: $724,384. B

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PROJECT MANAGER: Andrew L. Lissner.

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AFFILIATION: ScienceApplicationsInternationalCorporation.

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

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PRINCIPALINVESTIGATORS" JamesA. Blake, AndrewL. Lissner. i

KEYWORDS: MarineinvertebratesCn, idaria,Hydrozoa,Anthozoa,Octocorallia,Stolonifera,Pennatulacea, []

Actinaria,Ceriantharia,Zoanthinaria,California,Santa Maria Basin,Santa Barbara Channel,continental 11 shelf.

BACKGROUND: The TaxonomicAtlasofthe Santa MariaBasinandwesternSantaBarbaraChannelisan R

extensionof the benthicreconnaissance(PhaseI) andmonitoringprograms(PhaseII) thatwereconducted W

bythe MMS since1983. The organismsthatwere collectedas partof thoseprogramsprovidethe material

onwhichthe Atlasisdeveloped.Inorderto fullydocumentthefaunacollectedbythoseprograms,a series m

of14 volumeswillbe preparedthat providekeys,descriptions,and illustrationsofthe benthicfaunaofthe

hardandsoftsubstrateenvironments.A team of40 expertsonthefaunahas beenassembledto carryout

this workand their contributionsare distributedamongthe 14volumes, i

M OBJECTIVES: Theobjectives ofVolume 3 are to describe the Cnidariathat occurin the Santa Maria Basin

and Western Santa Barbara Channel. Three chapters covering the Hyrezoa,the Octocorallia (Anthozoa),

and threeorderof anemones(Anthozoa). i

DESCRIPTION:Volume3 treatsthe speciescomprisingthe PhylumCnidaria. This phylumincludesthe

hydroids, octocorallians,and so-calledsea anemones. The text is organizedinto 3 chapterschapters:

Hydrozoa;Anthozoa, Subclass Octocorallina;and Anthozoa, the Orders Actinaria,Ceriantharia,and =

Zoanthinaria. These chaptersare organizedinto sectionsthat includethe morphology,taxonomichistory,

biology,keysof species,anddescriptionsof generaandspecies. Eachspeciesisfullyillustrated,oftenwith tii

relevantcharacteristicslabeledand identified. =

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I

ACCESS NUMBER: 30484.3

I SIGNIFICANT CONCLUSIONS: The keys, detaileddescriptions,and illustrationsto thevarioushyrdoids,

octocoraLsa,ndanemonespresentedhererepresentanimportantcontributiontounderstandingthedistribution

iI of cnidariansin the offshorewatersof California.Manypoorlyknownspeciesare newlydefinedwithnew

I

illustrations.

STUDY RESULTS: The fauna ofthe Santa MariaBasinandwesternSanta BarbaraChannelis evaluated

I from samplesthat were taken from depths of about50 to 1,000 m. Organismswere collectedfromsoft sediments using box cores and from rocks using manipulatorarms of submersibles and remotelyoperated

vehicles. The collections are organizedinto sets of vouchers from Phase I and II that have been made

I avNatur ailaablleHistory to theMuseum team ofofinvestiga LosAngtoerless.AdCounty ditional isalso matberiaeinglfrexamined. omthe bulkThecoltotal lectionunsmnber owofarchspeivciedewstritheated the

intheentireTaxonomicAtlas(14volumes)mayexceed1,000species.

I Volume3, representingthe PhylumCnidariaincludesthe hydroids,octocorals,and 3 orders ofanemonies:

Actinaria,Ceriantharia,and Zoanhinaria. A total of 40 generaand 49 species are included. None are new

to science, several of the anemones are likely new species, but are poorly representedin the collections.

i Numerousspeciesareredescribedand illustrated.

Among the Hydrozoa, 24 species and 17 genera are treated. Each species is illustrated and keys are

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forreeach sentedspteocialels. geThe nera1and 2species specieosf

ioctocorals ncluded. Where arenewly availaillust ble

ra,ateddiscandussio describ nofreed,leva provid ntbioinglogy reaindersincwiluded tha

usefulguideto assistintheiridentification.Mostoftheanemonesbelongto theOrderActinadaandinclude

manyof thecommonseaanemonesfoundinoffshorewaters. In contrast,theavailablespecimensofthe

I OrderCefianthariawere mostlytoosmall to be identifiedand the Order wasnotadequatelytreated. Onlya

singlespecies of Zoanthinariawas found in the collections.

i STUDY SantaMaPRODUCT: fia BasinandBlaWeste ke,J.Arn.anSanta d P.Barbara H.Scott (EChannel. ditors).Volume 1996. T3ax. The onomCnida icAtlasria.oAfthfienal Bereport nthicFprepared aunaoftheby

ScienceApplicationsInternationalCorporation,fortheU.S. DepartmentoftheInterior,MineralsManagement

t Service, PacificOCS Region,Camanllo, CA. OCS Study MMS 96-0044. ContractNo.14-35-0001-30484.

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i "P.l'saffiliationmaybedifferentht anthatlistedforProjectManager(s).

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ACCESS NUMBER: 30484.3

I_MeI. P_ llil _ I17 e I

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LOS ANGELES

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KEY MAP

PACflC REG_N _

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SouthernCaliforniaPlanningAreaIndicatingtheTaxonomicAtlasStudyRegion.

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REPORTDOCUMENTATIONs.n[eowrNO. MMS 96-0044 2" =. R=¢i_= _=¢e._, No.

PAGE

4. _ end Subtitle S. Report I)ate

TaxonomicAliasofthe Santa MariaBasinandWastemSantaBarbaraChannel.Volume July 1996

3. The Cnidaria. ¢_

7. Au_hor(a) 8. Performir_ Orl_nization RepL No.

James A. Blakeand PaulH. Scott(editors)

9. Pen_ormintOq_nizMion Name and Acldms 10. Projectrrnk/Work Unit No,

ScienceApplicationsInternationalCorporation lz. Co,t_,_ orGre,,(mNo.

10260 CampusPointDrive

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

(G)

12. SI=onsorinl Orlanlzetlon Name and Address 13. Type of _eport & PeriOdCovered

Final 3 of 14volumes,

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

Camarillo,California93010 :4.

ZS. SUpl_amentary Notes

Contributingauthors: Dr. DaphneFautin,Dr.F. G. Hochberg,Mr.JohnLjubenkov.

16, Abstm_"t(Umft: 200 words)

Volume 3, representingthe PhylumCnidariaincludesthe hydroids,octocorals,and3 orders of anemonies:Actinaria,

Ceriantharia,andZoanhinaria. A totalof40 generaand 49 species are included. None are new to science,severalof

the anemonesare likelynew species,but are poorly representedin thecollections. Numerous speciesare redascribed

and illustrated.

Among the Hydrozoa,24 species and 17generaare treated. Each species is illustratedand keys are presentedto all

generaand species included. Where available,a discussion of relevantbiology isincludedfor each species. The 12

species of octocorals are newly illustrated and described, providing readers with a useful guide to assist in their

identification. Most of the anemones belongto the Order Actinaria and include many of the common seaanemones

found in offshore waters. Incontrast, the availablespecimens ofthe Order Cerianthariawere mostly too small to be

identifiedandthe Order was notadequatelytreated. Onlya singlespecies of Zcanthinariawas found in thecollections.

17. Dacume_t Analysis s. D*sc_l_rs

Marineinvertebrates,Cnidada, Hydrozoa,Anthozoa,Octocorallia,Stolonffera,Pennatulacea,Actinaria,Ceriantharia,

Zoanthinada,taxonomy,California,SantaMariaBasin,Santa BarbaraChannel,continentalshelf.

b. IdentlflemlOp4m-Ended Terms

TaxonomicAtlas,Keysto species,faunal guide,marineecology,Cnidaria,hydroids,anemones,sea pens,marine

benthos.

¢, COSATI Fi_dlGroup

ZL AveUIllbil_tyStlltimlnt |g. Security CIISS {ThisReport) 2/. No. of Pallas

Unlimited Unclassified 116

_. S,curayCh"==(ThisPqSe) _.

ANSI-Z39.I_r) Sea lnstrtmtk)ns cm Reverie OPTIONAL FORM 272 (4--7/)

(Formerly NT1S-35)

I _rt_ of Coeumerce
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Table of Contents w

1. CLASSHYDROZOA(by F. G. Hochberg and John Ljubenkov) I

Inlxoduction ..................................................................................................................................... I _'

Glossary of Morphological Terms .................................................................................................. 2 n

Abbreviations Used in the Figures .................................................................................................. 4

List of Species ................................................................................................................................. 5 Ii

Key to Families of Attached Polypoid Stages ................................................................................. 6 l

Description of Species ..................................................................................................................... 7

Family Bougainvilliidae Lutken, 1850 .............................................................................. 7 Ii

Key to Species of Bougainvilliidae ....................................................................... 7 l

Rhizorhagiumformosum (Fewkes, 1889) ................................................ 7

Perigonimus repens (Wright, 1858) ......................................................... 9

Family Corymorphidae Altman, 1872 ............................................................................. 10 l

Key to Species of Corymorphidae ................................................................................................ 10

Corymorpha palma Torrey, 1902 .......................................................... 11 1

Euphysa sp. A ......................................................................................... 13 II

Euphysora bigelowi Maas, 1905 ............................................................. 14

Family Olindiidae ............................................................................................................. 16 11

Monobrachium parasimm Mereschkowsky, 1877 ................................. 16 I

Family Campanulariidae Johnston, 1836 ......................................................................... 18

Clytia denticulata (Clark, 1876) ............................................................ 19 al

Laomedea altitheca (Fraser, 1948) ........................................................ 20 I!

Family Campanulinidae Hincks, 1868 ............................................................................. 21

Key to Species Campanulinidae .......................................................................... 21 Im

Calycella syringa (Linnaeus, 1767) ....................................................... 21 l

Oplorhiza gracilis (Stechow, 1921) ....................................................... 23

Family Lafoeidae A. Agassiz, 1865 ................................................................................. 24 _.

Acryptolaria pulchella (Allman, 1888) .................................................. 25 |

Family Lovinellidae I-Iincks, 1868 ................................................................................... 26

Lovenella producta (G.O. Sars, 1873) ................................................... 26 /

Family Aglaopheniidae .................................................................................................... 27 i

Key to Species of Aglaopheniidae ...................................................................... 27

Cladocarpus gracilis Fraser, 1948 ......................................................... 28 lira

Cladocarpus vancouverensis Fraser, 1914 ............................................ 29 i

Family Plumulariidae L. Agassiz, 1862 ........................................................................... 30 m

Key to Species of Plumutariidae ......................................................................... 30

Plumularia corrugata Nutting 1900 ...................................................... 31 i

Plumularia exilis Fraser, 1948 ............................................................... 32 11

Family Sertulariidae l_,amouroux, 1812 ........................................................................... 33

Key to Species of Sertulariidae ........................................................................... 33 _l

Abietinaria amphora Nutting, 1904 ....................................................... 34 II

Abietinaria pacifica Stechow, 1923 ....................................................... 35

Abietinaria traski (Torrey, 1902) ........................................................... 36 1

Abietinaria variabilis (Clark, 1876) ...................................................... 38 I1

Dynamena pumila (Linnaeus, 1758) ...................................................... 39

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I Family Thyroscyphidae Stechow, 1920 ........................................................................... 41

Key to Species of Thyroscyphidae ...................................................................... 41

I SympIectoscy mplectoscyphus ped tricuspidata rensis (Torrey, (Alder, 1904) 1856).......................................... ....................................... 413

Symplectoscyphus turgidus (Trask, 1857) ............................................. 44

I Literature Cited ............................................................................................................................. 46

2. CLASS ANTHOZOA: SUBCLASS OCTOCORALLIA, ORDERS STOLONIFERA AND PENNATULACFA

i (by Introduction F. G. Hochbe................................................................................................................................... rg and John Ljubenkov) 53

Collection and Preservation .......................................................................................................... 54

i Labo Glossary ratory......................................................................................................................................... Study ........................................................................................................................... 556

List of Species ............................................................................................................................... 57

I DescriptiFamily on of Spe Telestidae cies ................................................................................................................... ............................................................................................................. 58

Telesto californica Kiikenthal, 1913 ...................................................... 58

I Telesto Telestulanuttingi ambiguaKiikenthal, (Nutting, 1913 1909)........................................................... ........................................................ 5961

Key to Families of Pennatulaceans ..................................................................... 62

i Suborder Sessiliflorae Kiikenthal, 1915 ........................................................................................ 65 Family Stachyptilidae K611iker, 1880 .............................................................................. 65

StachyptiIum superbum Studer, 1894 ..................................................... 65

i Family Ombellulidae Williams, 1995 .............................................................................. 66 Ombellula magniflora K611iker 1880 ..................................................... 66

Suborder Subselliflorae Kiikenthal, 1915 ..................................................................................... 68

i Family Halipteridae Williams, 1995 ................................................................................ 68 Halipterus californica (Moroff, 1902) ................................................... 68

Family Pennatulidae Ehrenberg, 1834 ............................................................................. 70

Pennatula californica Kiikenthal, 1913 ................................................. 70

I Ptilosarcus gurneyi (Gray, 1860) .......................................................... 71

Family Virgulariidae Verrill, 1868 ................................................................................. 73

Virgularia agassizii Studer, 1894 .......................................................... 73

I Acanthoptilum Nutting, ......................................................

album 1909 74

Acanthoptilura gracile (Gabb, 1863) ..................................................... 76

Stylatula elongata Gabb, 1863 ............................................................... 78

t Acknowledgments 80 Literature Cited ............................................................................................................................. 80

i 3. CLASSANTHOZOA:OnVEItSA_ C_mANTHAR_ANDZOANTH_AmA(by Daphne Gail Fautin)

Introduction ................................................................................................................................... 83

Biology .......................................................................................................................................... 84

i Glossary of Morphological Terms 84 Key to the Orders of Non-Skeletalized Hexacorallia .................................................................... 86

Order Actiniaria ............................................................................................................................. 86

\ Key to the Species of Actiniaria 87

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Description of Species ................................................................................................................... 88 II

Hormathia digitata (Miiller, 1776) ........................................................ 88

Actinostola cf. carlgreni Wassilieff, 1908 ............................................. 90 i

Metridium giganteum Fautin, Bucklin, and Hand, 1990 ........................ 91 II

Anemonactis mazeli (Jourdan, 1880) ..................................................... 93

Halcampa decemtentaculata Hand, 1955 .............................................. 95 i

Peachia quinquecapitata McMurrich, 1913b ........................................ 96 l

Pentactinia californica Carlgren, 1900 .................................................. 98

Edwardsia californica (McMurrich, 1913a) .......................................... 99 _1

Scolanthus sp........................................................................................ 101 l

Halcampoides purpurea (Studer, 1879) ............................................... 101

Calamactinia sp................................................................................... 103 i

Halcampella sp..................................................................................... 103 II

Order Ceriantharia ....................................................................................................................... 104

Order Zoanthinaria ...................................................................................................................... 105 E

Description of Species of Zoanthinaria ............................................................. 106 n

Epizoanthus induratum Cutress and Pequegnat, 1960 ......................... 106

Literature Cited .......................................................................................................................... 107 am

APPENDIX..................................................................................................................................... 111 i

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I List of Figures

i Figure 1.1. Rhizorhagiumformosum (Fewkes, 1889): A. entire colony on gastropod shell, natural size;

B-C. zooids; D. gonophore [after Fraser, 1937] ................................................................. 8

I Figure 1.2. Perigonimus repens (Wright, 1858): A. entire colony on shell of snail, Nassarius, natural

size; B. hydranths on terminal portion of stem; C-D. hydranths with gonophores [after

I Fraser, 1937] ..................................................................................................................... 10

Figure 1.3. Corymorphapalma Torrey, 1902: A. entire polyp, natural size; B. enlarged zooids showing

gonophores; C. male medusoid; D. female medusoid [Figure A. after Ricketts et al., 1968;

i B. after MacGinitie and MacGinitie, 1968; C, D. after Torrey, 1907] 12

Figure 1.4. Euphysa sp. A: A. entire polyps, natural size; B. adult specimen with developing medusa

I buds; C. young specimen; D. details of a tentacle showing rings of nematocysts [original illustrations] ...................................................................................................................... 13

Figure 1.5. Euphysora bigelowi Maas, 1905: A. composite drawing of mature polyp; B. newly released

I medusa; C. adult medusa; D-H: cnidome, undischarged nematocysts, D. small stenotele

(aboral tentacle, polyp); E. large stenotele (aboral tentacle); F. des moneme (aboral tentacle);

G. microbasic mastigophore (oral tentacle, polyp); H. anisorhiza (secondary tentacle, adult

I medusa) [after Sassaman and Rees, 1978]. Scales in mm .............................................. 15

Figure 1.6. Monobrachium parasitum Mereschkowsky, 1877: A. colony on shell of bivalve mollusk,

i Axinopsida serricata, note presence of dactylozooids at the margin of the shell; B. enlarged • section of colony showing cluster of feeding and reproductive zooids; C. gonozoids [Figure

A. after Hand, 1957; B. after Naumov, 1960; C. after Fraser, 1937]. Scale in mm ........ 17

I Figure 1.7. Clytia denticulata (Clark, 1876): A. entire colony, natural size; B. hydrotheca and pedicel showing characteristic annulations; C. detail of hydrotheca to show diaphragm [Figures AB. after Fraser, 1937; C. after Torrey, 1902] ................................................................... 19

I Figure 1.8. Laomedea altitheca (Fraser, 1948): A. entire colony, natural size; B. three hydrothecae

with long pedicels; C. fascicled stem hydrocaulus with hydrddaeca and gonangia; D. strongly

i corrugated gonangium [after Fraser, 1948] ...................................................................... 20 Figure 1.9. Calycella syringa (Linnaeus, 1767): A. entire colony, natural size; B. enlarged aspect of

colony; C. single hydrotheca showing characteristic short, annulated pedicel; D. stolon

t with after hydrothe Fraser, 1911; cae andD. after gonopNaumov, hores [Figure 1969]A...................................................................... after Fraser, 1937; B. after Calkin, 1899; 22C.

i Figure 1.10. Oplorhiza gracilis (Steehow, 1921): A. entire colony, natural size; B. enlarged section of

colony; C. nematophore [after Fraser, 1937]. Scale in mm ............................................. 24

Figure 1.11. Acryptolariapulchella (Allman, 1888): A. entire colony, natural size; B. portion of fascicled

I main stem; C. portion of simple stem or branch [after Fraser, 1937] .............................. 25

Figure 1.12. Lovenellaproducta (G.O. Sars, 1873): A. entire colony, natural size; B-D. single zooids to

show variation in sizes, note characteristic annulations only at base of pedicels; D. branched

I colony [Figures A, C, D, Fraser, 1937; Fraser, 1] ................................

after B. after 191 27

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Figure 1.13. CIadocarpus gracilis Fraser, 1948: A. entire colony, natural size; B. first hydortheca and

portion of proximal stem, lateral view; C. medium portion of stem, lateral view; D. two

hydrothecae, lateral view; E. arrangement of hydrothecae and nematophores on lower stem,

stylized, enface view [Figures A-D. after Fraser 1948; E. original] ................................ 28 II

Figure 1.14. Cladocarpus vancouverensis Fraser, 1914: A. entire colony, natural size; B. portion of lIB

hydrocladium, lateral view; C. hydrotheca, enface view; D. gonophores and phylactogonia;

E. arrangement of nematophores on lower stem, note absence of hyrotheca, stylized, enface m

view [Figures A-D. after Fraser, 1937; E. original] ......................................................... 30 III

Figure 1.15. Plumularia corrugata Nutting, 1900:A. entire colony, natural size; B. portion of main stem and B

hydrocladia; C. male gonphores; D. female gonophores [after Fraser, 1937] ...................... 31

Figure 1.16. Plumularia exilis Fraser, 1948: A. entire colony, natural size; B. portion of colony showing I

branching pattern and distribution of hydrothecae and nematophores; C. gonophores [after i

Fraser, 1948] ..................................................................................................................... 33 II

Figure 1.17. Abietinaria amphora Nutting, 1904: A. entire colony, natural size; B. portion of branch I

with hydrothecae, lateral view; C. gonophores [after Fraser, 1937] ................................ 35

Figure 1.18. Abietinaria pacifica Stechow, 1923: A. entire colony, natural size; B. portion of stem I

showing origin of branch; C. portion of stem showing hydrothecae and gonphores [Figures

A, C. after Fraser, 1948; B. after Fraser 1937 from original in Stechow, 1923] ............. 36

m

Figure 1.19. Abietinaria traski (Torey, 1902): A. Portion of colony, natural size; B. portion of branch i

with hydrothecae and gonophores, lateral view [after Fraser, 1937] ............................... 37

Figure 1.20. Abietinaria variabilis(Clark, 1876): A. Portion of colony, natural size; B. portion of branch B

with hydrothecae; C. branchlet with gonotheca [Figures A-B. after Fraser, 1937; C. after I

Naumov, 1969] ................................................................................................................. 38 in

Figure 1.21. Dynamena pumiIa (Lirmeus, 1758)."A. portion of colony near base showing arrangement i

of hydrothecae; B. two pairs of hydrothecae; C. gonotheca and pair of hydrothecae [after

Naunov, 1969]. Scales in mm .......................................................................................... 40 t

II Figure 1.22. Symplectoscyphus pedrensis (Torrey, 1904): A. entire colony, natural size; B. portion of

stem and smooth hydrothecae; C-D. details of sculptured hydrothecae; E. portion of stem

with gonotheca; F. detail of spinouse texture of gonotheca [Figures A, B, E. after Fraser,

1937; C, D, F. after Torrey, 1905] ................................................................................... 42 i

Figure 1.23. Symplectoscyphus tricuspidata (Alder, 1856): A. entire colony, natural size; B-C. portion ii

of stem showing branching and hydrothecae; C. branch with attached gonotheca; D. gontheca |

[Figures A, B, D. after Fraser, 1937; C. after Naumov, 1969] ........................................ 44

Figure 1.24. Symplectoscyphus turgidus (Trask, 1857): A. portion of colony, natural size; B. portion of

branch showing hydrothecae and gonothecae; C-D. details of gonotheca sculpture [Figures IJ

A-B. after Fraser, 1937; C-D. after Torrey, 1904] ........................................................... 45

Figure 2.1 Generalized illustration of an octocoral ........................................................................... 53 I

Figure 2.2 Telesto californica, views of colony: A, SBMMH 01-65-32; B, USC 1948-50; C, SBMNH

51530 ................................................................................................................................ 58 R

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Figure 2.3 Telesto nuttingi, views of colony: A, SBMNH; B, SBMNH 1348-4 ............................... 60

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I Figure 2.4. Telesto ambigua: A, views of colony, Holotype, USNM 25421; B, sclerites from the body

...................................... ......................... ........... ............................ ........................ ............ 61

i 2.5. Genera of sessiliflorae entire colonies: A. Renilla 10-25 Figure pennatulaceans, amethystina: In,

Redondo Canyon, CA (SBMNH 51257), 60 mm RL; B. Kophobelemnon affine: 2430-2710

m, Monterey Bay, CA (SBMNH), 100 mm TL; C. Ombellula lindhali: 1005-1060 m, off

I Avila, Banks, (SBMNH 54), mmTL; Anthoptilum grandiflorum:

Santa Lucia CA 491 350 D.

457-512 rn, off Avila, Santa Lucia Banks, CA (SBMNH 49152), 380 mm TL; E. Stachyptilum

superbum: 181 m, off Avila, Santa Lucia Banks, CA (SBMNH 49156), 100 mm TL; F.

I Distichoptilum gracile: 1260-1277 off Avila, Santa Lucia Banks, CA (SBMNH 49155),

m,

1100 mm TL ..................................................................................................................... 63

I Figure 2.6. Genera Santa Barbara of subselliflorae Channel, CApennatulaceans, (SBMNH 45568), entire950colonies: mm TL;A,B,Halipteris Stylatula elongata californica: (modified : 475 m,

from MacGinitie & MacGinitie, 1968), 280 mm TL; C, Virgularia agassizi: 14 m, off

I Santa Santa Barbara, Catalina CA (SBMNH 49487), 300 mm TL; D, Acanthoptilum gracile: 15-35 m, off Island, CA (SBMNH 49691), 650 mm TL; E, Ptilosarcus gurneyi: 15-35 m,

off Santa Catalina Island, CA (SBMNH 49688), 225 mm TL; F, Pennatula califomica: 475

i m, Santa Barbara Channel, CA (SBMNH 45509), 90 mm TL ........................................ 64 Figure 2.7. Stachyptilum superbum: A, B, views of colony; C, sclerites from the body ................... 66

] II Figure 2.8. Ombellula magniflora: view of colony ............................................................................ 67

I

I E Figure 2.9. Halipterus californica :A, B, C views of colony; D, sclerites from the body ................ 69

I Figure 2.10. Pennatula californica: A, view of colony; B, sclerites from the body ............................ 71 Figure 2.11. PtiIosarcus gurneyi: sclerites from the body ................................................................... 72

I Figure 2.12. Virgularia agassizii: views of colony .............................................................................. 74 Figure 2.13. Acanthoptilum album: A, B, C views of colony; D, sclerites from the body .................. 75

I Figure 2.14. Acanthoptilum gracile: A, B, C, D views of colony; E, sclerites from the body ............. 77 Figure 2.15. Stylatula elongata: A, B, C, views of colony; D, sclerites from the body ...................... 79

I Figure 3.I. BHormathia , specimen digitata with less: A,promine specimen nt protuberances with pronouncedattached protuberances to crustacean(fromHyas Grebel'nyi, (from Carlgre 1980); n,

1942) ................................................................................................................................. 89

I Figure 3.2. Actinostola groenlandica (from Carlgren, 1921; slightly larger than life): the numerous,

short tentacles and wrinkled column are typical of preserved specimens of all species of the

genus ................................................................................................................................. 91

i Figure 3.3. Metridium giganteum in strong current (from Koehl, 1976): A, side view; B, column and

underside of oral disc. Note raised collar near oral end of column, and bracing of oral disc

i lobes ................................................................................................................................. 92 Figure 3.4. Anemonactis mazeli (from Uchida, 1938): A, whole specimen; B, cross-section .......... 94

i Figure 3.5. HaIcampa decemtentaculata: cross-section from Hand (1955) illustrating distinctive arrangement of mesenteries .............................................................................................. 95

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Figure 3.6. Peachia quinquecapitata (from McMurrich, 1913b): A, whole animal; B, oral disc and I

tentacles ............................................................................................................................ 97

Figure 3.7. Pentactinia californica: A, cross-section, illustrating distinctive arrangement of mesenteries I

(from Carlgren, 1900); B, whole animal (from Carlgren, 1931) ..................................... 99

Figure 3.8. Di expanded agrammatic animal, sketchillustrating of Edwardsia basal (from physa,Williams long scapus , 1981): withA,nemathybomes, oral disc and tentacles; and distal B, I

scapus ............................................................................................................................. 100

Figure 3.9. Halcampoides purpurea (from Manuel, 1981a): distal end of expanded animal in situ .... 102 I

Figure 3.10. Ceriantharian (after Hyman, 1940) ................................................................................ 104

Figure 3.11. A colony of zoanthids (after Hyman, 1940) ................................................................... 105 I

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I Acknowledgments

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 Marie Murphy of the Santa Barbara Museum of Natural History provided graphic design and production assistance for this volume.

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i List of Acronyms

i AHF Allan Hancock Foundation, University of Southern California, Los Angeles, California. BLM Bureau of Land Management.

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

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,

i LACM CNatural aliforniHistory a, USA.Museum of Los Angeles County, Los Angeles, California, USA.

i MC MECZ Mu Ma seum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA. rine Ecological Consultants, Carlsbad, California.

i MMS Minerals Management Service. MNHNT Museum of Natural History Society, Newcastle-upon-Tyne, England.

i MSUI Museum of the State University, Iowa.

• OCSD Orange County Sanitation District, California.

i RCBM Royal British Columbia Museum, Victoria, British Columbia, Canada. SCAMIT Southern California Association of Marine Invertebrate Taxonomists.

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

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

I SMNH Swedish Museum of Natural History, Stockholm, Sweden. USNM United States National Museum. A historical designation for the National Museum of

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

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I 1. CLASSHYDROZOA

by

I F.G. Hochberg I and John Ljubenkov 2

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Introduction

I The Class Hydrozoa contains a large number of species, the majority of which measure less than an

few centimeters in height. Alteration of generations occurs in most genera. In classic life cycles this involves

I an asexual benthic polypoid stage alternating with a sexual planktonic medusoid stage. One or the other generation may be suppressed or lacking. The polyp stages treated here are typically the dominant stages in

the life cycle. Hydroid polyps are usually colonial and often polymorphic. Over 2,500 species have been

I dtreated escribed, here.theBmajority enthic siphonophores of which are ma (e.g., rine.Dromalia The class) and is divided stylasterine into sehydrocorals ven suborders(e.g., of wAllopora hich only) occur two arein

the depth range of the study area, but they have not been collected.

i Hydroids occur at all depths, but are most common in the littoral or shallow subtidal zones. The

• deep-water fauna off California is less diverse and more poorly understood than adjacent shallow-water

faunas.

I Our knowledge of the systematics of hydroids of the eastern pacific is in need of revision. In many

cases the two stages in the life cycle have been studied and described by separate authorities which has

resulted in a morass of conflicting names for what are different life history morphs of the same species.

I reevaluations of in Vervoort and in the Western North Monographic hydroids Europe by Cornelius, others;

Atlantic by Calder; and in the Western North Pacific by Naumov and others have resulted in a number of

recent changes in the definition of families and genera, and clarification of species names. New collection

i and examination of live material is essential in such revisionary work.

The treatment in this Atlas is taken principally from the works of Fraser (see literature cited; see

i also Schmitt, 1948). Many of his names refer to European species which will eventually need to be reevaluated.

A number of Fraser's species appear to be identical to those described by Nutting and need to be critically

reevaluated by a study of types and other identified material in museum collections. Much of material

i studied or identified by Fraser does not appear to be extant. The specimens and types which document Fraser's 1948 paper were transferred from the Allan Hancock Foundation at the University of Southern

California to the Santa Barbara Museum of Natural History and are now available for study.

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_12_artment of Invertebrate Zoology, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road,

I Santa Barbara, CA 93105-2936 USA

2MEC Analytical Corp., 2433 Impala Drive, Carlsbad, CA 92009 USA

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II

Glossary of Morphological Terms •

Abcauline. Off, from or away from the stem; used Cnidae. IntraceUular "stinging capsules" that are 1

in reference to the side of a hydrotheca, used in offense anddefense (also termed

nematocysts).

Aboral. Pertaining to the region away from or II

most distant from the mouth. Cnidome. The collective types of cnidae II

Adeauline. Next to or toward the stem; used in possessed by a cnidarian or a part thereof.

reference to the side of a hydrotheca. Coenosarc. The cellular and intercellular i

substance of a zooid or whole colony.

i

Adnate. Congenitally joined together; said of

unlike parts (e.g., one side of hydrotheca Colony. Genetically identical individuals formed t

growing attached to stem ). by asexual reproduction that live as a group II

of individuals or zooids physically

Alternate. A branching pattern in which the connected by a common stem or stolon. It

branchlets of a hydroid colony come off the []

stem at alternate levels. Pattern of branching Daetylozooid. A small mouthless zooid armed U

may be regular or irregular, with nematocysts, specialized for protective

functions; tentacles and gastrovascular H

Anchor Filaments. Filaments at base of stem of cavity lacking.

solitary polyps (e.g., Corymorpha) which

serve as a holdfast to anchor polyp in soft Diaphragm. A transverse, perforated partition or In

substrates (also termed frustules), thin shelf located at the base of the l

hydrotheca; forms a support for the

Annulated. Possessing a ringed appearance; hydranth.

usually refers to the series of rings on the

pedicel at the base of the hydrotheca or Distal. Toward the oral region or tip of a branch, U

adjacent to the attachment with the stem. away from the base or stem.

Athecate. Hydroids which lack a protective, Fascicled. A bundle or association of two or more B

chitinous covering or hydrotheca around the stems; varying degrees of intimacy, either in

hydranth; also used to refer to intemodes in loose contact or tightly associated, with few m

some hydroids that lock hydrotheca or many cross communication (also termed I

(opposite of thecate), polysiphonic).

Blastostyle. Modified zooid (gonozooid) from Filiform. Long, threadlike tentacle over which the i

which medusoids are budded; axial portion nematocysts are evenly distributed; tapering il

of a gonangium; may be surrounded by a gradually but slightly from base to tip.

protecting theca (gonotheca or gonangium). Frustules. see anchor filaments. O

W

Calyeine. Location in relation to the hydrotheca Gastrozooid. Hydroid polyp specialized for

(e.g., subcalycine, below the hydrotheca;

feeding; hydranth. m

supracalycine, above the hydrotheca). •

Capitate. Knobbed at the tip, as in tentacle; Gonangium. Asexual reproductive polyp in

ml

certain hydrozoans; consists of an axial

distinct terminal knob studded with blastostyle and surrounding gonotheca. B

nematocysts. m

Cauline. Having or growing on a stem (e.g., Gonophore. The reproductive structures on a

abcauline, adjacent to the stem; adcauline, hdirectly ydrozoanto the colosexual ny which products 1) gives(gametes); rise or i

away from the stem). 2) gives rise to an attached or free medusa

which in turn releases gametes; or 3) the []

sexual medusa itself. II
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I Gonosome. All specialized generative zooids of a Monosiphonic. Simple, not fascicled or

colony and associated perisarcal structures polysiphonic; in reference to the stem.

wsexual hich which reproduction are directly (i.e., involved produce medusae with Nematoeyst. A type of cnida.

or gametes). Nematophore. The small, highly modified zooid

I Gonotheca. The peridermal cup- or vaselike containing protective function; nematocysts characteristic which are ofusedthefor

structure surrounding the reproductive polyp

or blastostyle of thecate hydroids. Family Plumulariidae.

I Gonozooid. Hydrozoan zooid specialized for

Nematotheea. The chitinous, peridermal, cuplike

structure surrounding the nematophores or

sexual reproduction, nematocyst containing defensive polyps of

I Hydrant&. The feeding polyp consisting of the thecate hydroids. body, hypostome, mouth and tentacles, but

not including the hydrotheca or pedicel. Nematozooid. Zooid specialized for defense;

I Hydroeadus. Branched, upright, axis connecting bears (see dadense ctylozooi concentration d). of nematocysts

and supporting the polyps of an erect

hydrozoan colony; or the single pedicel in Opereulum. Chitinous flap or lid of one or more

I segments that closes the aperture of the solitary forms (alsotermed stem).

hydrotheca or gonotheca when the polyp is

Hydroeladium. Lateral branch(es) growing from retracted.

I Opposite. In reference to a branching pattern in

hydrocaulus.

Hydrorhiza. Branched, rootlike structure which the branchlets of a hydroid colony

i attaching colony to the substrate (also come off the stem at the same level termed stolon). (compare with alternate).

Hydrotheca. The chitinous, peridermal, cuplike Oral. Adjacent to the mouth, as in tentacles.

i structure surrounding and protecting the feeding polyp of most thecate hydroids; Pediele. A small stalk or stalklike support that

hydranth may be completely drawn in when bears the hydrotheca.

I disturbed (also termed theca). Peduncle. In hydromedusae, the cone of jelly

Hypostome. Conical raised area surrounding the which hangs down into the subumbrellar

cavity and on which the manubrium and

oral region in some hydrozoans; bearing the stomach are situated.

I tentacles (also termed manubrium,

proboscis). Pelliele. Thin, translucent covering the stalk of

some athecate hydroids.

t lnternobetwee de. Thn atnoticea portionbleofjointhetsstemor node or sbranch ; Periderm. see perisarc.

characteristic of the Plumulariidae.

i Perisare. Thin, chitinous, noncellular, nonliving manubrium. A tubular structure bearing the protective covering surrounding the stolons

mouth at its free end (see also hypostome), and upright branched portions of some

I Margin.termEdge ed rim. of hydrotheca aperature; also hydrozoan cells of the cocoenosa lonies;rc.secreted by ectodermal

Phylaetogonium. Protective branch or leaf of

' I/ Medusoid. Medusa stage which is sessile or

remains attached to the colony, hydroid colony surrounding the gonangia; II containing nematophores.

mesial. In, near or toward the middle.

Planula. Ciliated free-swimming larval stage with

I a solid, cylindrical body.
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Polymorphism. Polyps modified to perform Stolon. see hydrorhiza. 1

specific functions - for example, feeding

gastrozooids, reproductive gonozooids, and Subealyeine. Situated below a hydrotheca. _l

defensive dzctylozooids. Supraealycine. Situated above a hydrotheca. J

Polyp. see zooid. Tentacle. Long, unsegmented, cylindrical In

protruberance from the oral region of the l_

Polysiphonlc. see fascicled.

polyp; used for food gathering (see

Proximal. Toward base or stem, away from oral modifiers: capitate, filiform, moniliform; ab

region or tip of branch, aboral, oral; primary, secondary). I

Pseudohydrotheea. False theca which partially Theca. see hydrotheca.

surrounds hydranth in some athecate _l

hydroids. Theeate. Hydroids with a chitinous (perisarcal) It

covering surrounding the hydranth.

Radial canal. In medusae, the canals extending Opposite of athecate.

from the stomach to the ring canal. Trophosome. All specialized portions of a I

Ring canal. In medusae, the canal extending hydroid colony or zooid and associated

around the margin of the bell. perisarcal structures which are directly il

Sessile. Hydrotheca or other structures attached involved with feeding and defense;

U

structures not directly involved with sexual

directly to stem or branch of a colony; reproduction (see gonosome), m

lacking a pedicel.

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Tubular. Sides parallel or nearly so; used to

Solitary. Individual polyps not physically attached describe a hydrotheca.

to others. I

Sporosae. Reduced gonophore that does not Whorl. Tentacles arranged in a circle on the circumference of the hydranth.

develop into a free medusa, but remains ill

attached and produces gametes. Zooid. The individual member of a colonial I

organism; may be nutritive, generative, m

Stem. see hydrocaulus, defensive, sensory or a combination of the

above (also termed polyp). I

Abbreviations Used in the Figures ][

iN

ab, adaxial nematocyst bulb ec, endodermal canal nt, nematotheca

ac, acrocyst ga, gastrozooid op, operculum j

III

ad, axial canal go, gonozooid ot, oral tentacle

af, anchor filaments gp, gonophore pb, pedicel branch (first) II

ar, annular rings (annulation) gt, gonotheca pd, pedicel N

at, aboral tentacle hr, hydrorhiza pr, perisarc

bp, bifurcate phylactogonium hs, hypostome pt, primary tentacle I

Ill

bs, bivalve shell ht, hydrotheca (in, intemode) (n,node) rc, radial canal

c, chevron (=oblique internode) hy, hydrocaulus s, stolon 1

da, dactylozooid m, manubrium st, secondary tentacle |

di, diaphragm np, nematophore tb, terminal bulb

te, tentacle I

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i List of Species

I Order Hydroida

Suborder Anthomedusae [= Athecata, Gymnoblastea]

I Family Bougain Rhizorhagiu villiidae m formos Lutken, um 1850 (Fewkes, 1889)

Perigonimus repens (Wright, 1858)

i Family CCorymorpha orymorphidae Altman, 1872 palma Torrey, 1902

Euphysa sp. A

I Euphysora bigelowi Maas, 1905 Family Monobrachiidae Mereschkowsky, 1877

I Monobrachium parasimm Mereschkowsky, 1877

Suborder Leptomedusae [--Thecata, Calyptoblastea]

I Family Campan Clytia denticula ulariidaetaJohns (Clark, ton, 1876) 1836

Laomedea altitheca (Fraser, 1948)

I Family Campanulinidae CalyceU Hincks, 1868 a syringa (Linnaeus, t767)

Oplorhiza gracilis (Stechow, 192I)

"l Family Lafoeidae A. Agassiz, 1865

Acryptolaria pulcheUa (Allman, 1888)

i Family Lovenellidae Lovenella producta (G.O. Sam, 1873)

Family Aglaopheniidae

I Cladocarpus CIadocarpus gvancouverensis racilis Fraser, 1Fr9a48ser, 1914

Family Plumulariidae L. Agassiz, 1862

i Plumu Plumularia laria corrugata exilis FraseNr,utting, 1948 1900

Family Sertulariidae Lamouroux, 1812

I Abietinaria amphora Nutting, 1904 Abietinaria pacifica Stechow, 1923

Abietinaria traski (Torrey, 1902)

i Abietinaria variabilis (Clark, 1876) Dynamena pum//a (Linneus, 1758)

i Family Thyroscyphidae Stechow, 1920 Symplectoscyphus pedrensis (Torrey, 1904)

Symplectoscyphus tricuspidata (Alder, 1856)

i Symplectoscyphus turgidus (Trask, 1857)

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Key to Families of Attached Polypoid Stages •

Hydranths unprotected by hydrothecac or pseudotheca; gonophores unprotected by gonangia ...... l

1A.

....................................................................................................... Suborder Anthomedusae ......... 2

lB. Hydranths protected by hydrothecae; gonophores protected by gonangia or similar structures...... _1

....................................................................................................... Suborder Leptomedusae ........ 4 II

2A. Hydranths with a single f'fliform tentacle ................................................................ Monobrachiidae i

2B. Hydranths with numerous f-fliform tentacles; typically in 1 or 2 whorls ........................................ 3

m

3A. Hydranths with single basal (aboral) whorl of tentacles .......................................... Bouganvilliidae I

3B. Hydranths with 2 whorls (oral and aboral) of tentacles ........................................... Corymorphidae B

II

4A. Hydrotheca free from stem or branches (may be adnate in some Lafoeidae) ................................. 5

4B. Hydrotheca sessile and more or less adnate to stem or branches .................................................... 7 I

5A. Hydrotheca without operculum ....................................................................................................... 6 i

5B. Hydrotheca with operculum and with diaphragm ................................................... Campanulinidae II

6A. Hydrotheca tubular, with or without diaphragm ............................................................... Lafoeidae i

6B. Hydrotheca campanulate, with diaphragm ............................................................. Campanulariidae

Im

7A. Nematophores present on stem or branches or both; hydrotheca without operculum .................... 8 I

7B. Nematophore absent; hydrotheca with operculum .......................................................................... 9

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8A. Gonophores without protection ................................................................................... Plumulariidae

8B. Gonophores protected by phylactogonia or corbulae ............................................... Aglaopheniidae 1

9A. Hydrotheca margin entire (smooth), operculum with 1 or 2 valves; gonophores smooth or with

longitudinal ridges ......................................................................................................... Sertulariidae /

I

9B. Hydrotheca margin with 3 teeth; operculum with more than 2 valves .................... Thyroscyphidae

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Description of Species

I Family Bougainvilliidae Lutken, 1850

i• proboscis.DiaGonophores gnosis. Perisarc producing well desporosacs veloped; hydrants or free medusae. with single basal whorl of filiform tentacles and conical

i Key to Species of Bougainvilliidae

I 1A. Gonophores .................................................................................................................... producing sporosacs; sporosacs not permanently surrounded Rhizorhagium by perisarc ................... formosum

lB. Gonophores producing free medusae; medusae with tentacles arranged singly ..... Perigonimus repens

I Genus Rhizorhagium Sars, 1874

I as thin pseudohydrotheca. Diagnosis. Colony bGonophores ranchedor unfixed brancsporosacs; hed. Hydranths bornefusiform; on distinct perisarc branchexten -likedspedicels. up aroundSporosa hydranthcs

initially enclosed in thin perisarc; bursts in later stages and then confined to pedicels.

I Remarks. The hypstomes of bougainvilliid hydranths generally are conical with a single, regular (or irregular) circlet of tentacles. The polyps of Rhizorhagium are very similar to the polyps of Perigonimus

but the latter have a more distinct pseudohydrothecal structure and the perisarc does not appear silty.

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Rhizorhagiumformosum 0Fewkes, 1889)

I Figure 1.1

Perigonimusformosus Fewkes, 1889: 6.

I Bimeriaformosa: Torrey, 1902: 8.

Garveiaformosa: Fraser, 1911: 23; 1937: 35, pl. 5(figs. 20a-d); 1948: 195.

i Rhizorhagiumformosum: Cairns et al., 1991:14

Material Examined. California: Santa Mafia Basin, off Pt. Buchon, 35°05.8'N, 120°49.2%V,91 m;

I coll. Battelle, Additional M/V Aloha material , cruise examined:-- 1-1, Sta. R-l(rep OCSD 8609, 3), OctoSta.berCON1986 (SBMNH). (rep 4), 57 m, 21 Apr 1986.-- OCSD

8503, Sta. 0 (rep 4), 57 m, 23 Oct 1985.

l Description. Trophosome: Stolons grow in cracks and sutures of gastropod shells andother suitable hard substrates; occasional branching, but without obvious reticulated structure. Individual zooids arise

separately on pedicels; height variable, up to 4 mm. Perisarc smooth or wrinkled; with texture produced by

I incorporation of veryfine silty material; terminates above level of tentacles and basal to hypostome. Hydranth with 10-16 tentacles in single whorl.

Gonosome: Gonophores arise singly from stolon; borne on short pedicels. Perisarc thick in early

I stages.

Nematocysts: Not described.

i Type Locality and Type Specimens. Not searched. Off the coast of California.

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Figure 1.1. Rhizorhagiumformosum (Fewkes, 1889): A. entire colony on gastropod shell, natural size; B-C.

zooids; D. gonophore (afterFraser, 1937). I

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Distribution. EasternNorthPacific Ocean, recordedfrom SanFrancisco Bay, California to Natividad w

Island, Baja California, Mexico. Depth range, 10-550 m.

Biology. The biology of this species has not been elucidated. I

Genus Perigonimus M. Sars, 1846 i

!1 Diagnosis. Colonies stolonal on shells of bivalves; with monosiphonic or polysiphonic hydrocaulus.

Perisarc soft, forming a pseudohydrotheca. Hydranth with single distal whorl of filiform tentacles; hypostome

conical. Gonophores free medusae, arising singly or in clusters from hydrorhiza, hydrocaulus or hydrocladia. I

Medusae with short marmbrium; oral tentacles I per radial canal, usually branched. Radial canals 4; ring

J

canal present. Marginal tentacles all alike in structure, arising in clusters from 4 tentacle bulbs. Ocelli

usually present. Gonads on manubrium. Medusa buds occasionally produced. I

Remarks. There is considerable confusion with regards to the status of Perigonimus. The genus

was reviewed by Rees (1956). The medusa stage needs to be recovered in order for the identification of the

genus and species to be clarified. •

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I Perigonimus repens (Wright, 1858)

i Figure 1.2

Eudendriumpusillum Wright, 1857: 231; 1858: 448.

I Atractyl Perigonimus is repens Wright, 1858: 450. repens: Allman, 1864: 365.--Calkins, 1899: 339.--Torrey, 1902: 29-30.--Fraser, 1911" 24;

1913: 150; 1914: 120; 1936: 123; 1937: 38, pl. 6(figs. a-d); 1938a: 17; 1938c: 131; 1948: 195.--

i McCormick, 1965: 141-144.--Lees, 1986: 105-106.

Material Examined. California: Santa Maria Basin, off Pt. Sal, Sta. BSS-40, colony on Saxicavella

i pacifica colony ,on(USNM); off Pt. Conception, colony on Nuculana leonina, Sta. BSS-78 (SBMNH); off Pt. Sal, snail, 123 m; cruise 1-1, Sta. PJ-7, (rep 1), Box Corer, October 1986; (SBMNH).

Additional material examined: OCSD Voucher: colony on Nassariusperpinguis, Survey 8948, Sta.

I C-2 2 (rep(repB),2),25055ftm,[7622m], Jul71989, Dec 1986, MEC MEC A106.A69; CSAcolony Voucher: on Nassariusperpinguis colony onBittium sp., , OCSD panel 1,Survey survey8718, 2, Sta.St1-a.

C-2 (rep 3), 56 m, 12 Jan 1987, OCSD Survey 8948, Sta. C-2 (rep 2), 55 m, 22 Jul 1989; OCSD Survey

I 8948, Sta. 5 (rep 2), 60 m, 22 Jul 1989. Description. Trophosome: Colonies typically grow on shells of gastropods and ostracods. Individual

polyps arise directly from reticulated stolon which grows in grooves and sutures of shell substrate. Polyps

I single, height fromsimple,1.5-2occmm. asionally Perisarcbranched typicallyto produce with same2 polyps; diameterwith throughout thin, clear butpseudohydrotheca; with occasional irregula maximum rities

or constrictions; appears silty. Hydranths with about 10 tentacles; tentacle bases scattered, mainly falling in

'1 one cycte.Gonosome: Gonophores stalked; shape roughly spherical; arise singly or in pairs, arise either directly

from hydrorhiza or from lower parts of stem. Perisarc clear.

I Medusae: Free; with two well developed and 2 rudimentary tentacles.

Nematocysts: Not described.

I Type Locality and Type Specimens. Not searched; possibly not extant. Not traced. Distribution. Eastern North Pacific Ocean, Admiralty Islands, Alaska to Santa Elena Bay, Ecuador

and the Galapagos Islands. Depth range, 18-105 m. Also recorded from the Western North Atlantic Ocean,

Great Britain.

Biology. Off southern California this species is the main hydrozoan epibiont on: 1) gastropod mollusks

(Bittium quadrifilatum, Colus halIi, Nassa mendica, Nassarius fossatus, N. perpinguis, Turbonilla spp.); 2)

I bivalve mollusks casternsis, Nuculana leonina, and 3) ostracods. Colonies first

(Acila Saxicavella pacifica )

appear near the apex of the shell and eventually grow to cover the entire shell. Medusae are ready for

liberation in March.

i In southern California the greatest densities are found at about 90 m on the shelf edge whereNassarius

perpinguis is the prefered substrate. McCormick (1965) reported the presence ofPerignimus repens on 74%

i of the shel Remarks ls of the. bivalve, Fraser (1937) Acila careported stemsis,onlycollected two specie off Orego s in this n. genus Perigonimus on the west coast,

namely P. repens and P. serpens both Allman, 1863. We suspect they should be synonymized in the future

I asAtlantic the morphological and perhaps more differences thoughtareshould very sbelight. giveHowever, to a critical theseredescription species are ofboththefromformthefromEastern the Eastern North

North Pacific.

i
I

|

Figure 1.2. Perigonimusrepens (Wright, 1858): A. entire colony on shell of snail, Nassarius, natural size; B.

hydranths on terminal portion of stem; C-D. hydranths with gonophores (afterFraser, 1937).

I

At least two other species of Perigonimus occur in the region: P. yoldia-arcticae Birula, 1897

which lives exclusively on the shells of bivalve mollusks and an undescribed species, P. sp. A. which lives

on the introvert of the sipunculid Nephasoma diaphanes. Both of these species have been collected in the II

Santa Maria Basin and western Santa Barbara Channel.

J

Family Corymorphidae Altman, 1872 I

Diagnosis. Zooids solitary, large. Hydrants with proximal (oral) and distal (aboral)rings of filfform I

tentacles. Gonophores producing free medusae with 4 radial canals; 3 of the 4 tentacles very much reduced.

me

Key to Species of Corymorphidae II

1A. Oral tentacles few, capitate; polyp size minute ................................................................ Euphysa sp. A I

lB. Oral tentacles many, filiform; polyp size medium to large ................................................................... 2

2A. Aboral tentacles numerous, 50-70 in number .......................................................... Corymorpha palma I

2B. Aboral tentacles 15-20 in number ............................................................................ Euphysora bigelowi

i

l0 !
!

I Genus Corymorpha M. Sars, 1835

Diagnosis. Pedicel with thin perisarc; tubular, fleshy processes (anchor filaments) arising from

i pedicel near the base. Hydranth abruptly distinct from pedicel; with 2 distinct sets of f'diform tentacles; proximal tentacles longer than distal tentacles; distal set in several contiguous rows. Gonophores borne on

branched pedicels between 2 whorls of tentacles; producing free medusae with 4 radial canals; 3 of 4 tentacles

I aborted or much reduced.

very

Remarks. The polyp genus Corymorpha includes a heterogeneous mixture of several separte

evolutionary lines (see Sassaman and Rees, 1978). One line is associated with the medusa genus Euphysa

I Forbes, 1848 (see comments for that genus) and the second with the medusa genus Steenstrupia Forbes,

1846. Despite differences of nomenclatural opinion, the separation of the family Corymorphidae into several

distinct lines is generally accepted as elucidated by Rees (1957) !

Corymorphapalma Torrey, 1902

I 1.3

Figure

i Tub Corymorpha ularia ? Hargitt, palma Torrey, 1902. 1902: 37-43, pl. 2(fig. 21).--Torry, 1902: 987.--Torrey, 1904:9-10. Fraser,

1911: 27; 1937: 49, pl. 9(fig. 39a-b).--Lees, 1986: 105-114.

i Material Examined. California--1 specimen; 34°56.9'N, 120_49.9"W, 142 m; coll. Battelle, M/V

A/oha, cruise 1-1, Sta. PJ-8 (rep 2), Box Corer, October 1986 (SBMNH).

II Additional material examined: Anaheim Bay Voucher: survey 0990, Sta. R (rep. 1); beam trawl;

II MEC A400.

Description. Trophosome: Polyps large, solitary; length up to 14 cm. Hypostome with 2 cycles of

I tentacles Aboral . Oral tentacles villiform; 18-30 in number, irregularily arranged in several whorls around mouth. tentacles viUiform; numerous, 50-70. Perisarc with numerous anchoring filaments at base of polyp;

filaments forming rootlike mass which attaches polyp to substrate.

I distal tentacles. Gonosome: Gonophores borne on peduncles attached to body of hydranth between proximal and

Medusae: Attached medusoids, with 4 radial canals; manubrium, often several times longer than

bell.

Nematocysts: Not described.

I Type Locality and Type Specimens. Not traced. San Pedro, California. Distla'bullon. Eastern North Pacific Ocean: central California to northern Baja California andpossibly

the Gulf of California, Mexico (JL, pers. obs.). Depth range from low intertidal, 0-60 m.

i Common Name. hydroid. Falrypalm

Biology. The species typically lives in bays and estuaries where the water is turbid from moderate

currentvelocities or offshore in soft mud habitats where the sediment is likewise stirredup. Eggs are laidin

I May to June.

The large size and relative ease of collecting and maintaining Corymorpha palma in the laboratory

I hhasas fbeen acilitated an extensive literature dealing with the biology of this species. The behavior of the species treated in papers by Torrey (1904a, 1905), Parker (1917), and more recently by Wyman (1965),

Ball (1973) and Ball and Case (1973). Development, regeneration, budding, and fission have been extensively

investigated by Torrey (1907, 1910a-c). Details of the aggregation of dissociated cells is discussed in Child

|
I

I

i

8 I

at A I

I

I

ot II

af ii

C D

I

Figure 1.3. Corymorpha palma Torrey, 1902: A. entire polyp, natural size; B. enlarged zooids showing

gonophores;C. male medusoid;D. female medusoid[Figure A. after Ricketts et al., 1968; B. after 1

MacGinitie and MacGinitie, 1968;C, D. after Torrey, 1907]. II

(1928). The movement of chemicals through the tissues is addressed in Chapman and Pardy (1972) and I

algae associated with Corymorpha studied by Pardy and Rahat (1972). Details of holdfast movement and of

the statocyst were elucidated by Campbell (1968, 1969). i

!11

Remarks. Tubularine polyps and corymorphine polyps cannot be easily differentiated except for

their method of attachment to the substrate (anchored to hard substrate vs. "rooted" in mud). Fraser (1937)

placed this species in the family Corymorphidae, whereas Naumov (1969) considered Corymorpha palma 1

to belong in the Family Tubuladidae. Following Sassaman and Rees (1978) we have left this species in the II

Corymorphidae. i

Genus Euphysa Forbes, 1848

Diagnosis. Polyps solitary, small; stalk enveloped by thin perisarc. Hydranths with conical hypostome I

and 2 cycles of tentacles; oral or distal tentacles few in number, capitate; proximal tentacles numerous, II

moniliform. Gonophores borne on short stalks, arising from hypostome between tentacle cycles.

Medusae free; umbrella with or without apical projection; apex rounded, dome-like; with 4 radial _

canals, with 1-4 moniliform marginal tentacles often unequally developed; stomach not extending beyond

umbrella margin. Without ocelli. Gonads completely surrounding stomach, i
!

I Remarks. The genera Corymorpha and Euphysa have been def'med in a number of different ways.

Naumov (1969) placed both Euphysa and Steenstrupia (a hydromedusa name) in Corymorpha M. Sars. In

i tentacles Corymorphaare moniliform the tentacleswhereas are villiftheormoralortentacles moniliformare incapitate. both cycles. Most recent However, authorsin Euphysa place Euphysa the aboinraal

family distinct from Corymorpha on this basis.

I Euphysa sp. A

I Figure 1.4

Corymorpha aurata (non Forbes): Fauchald and Jones, 1976.

U Euphysa sp. Lees, 1986:106-114.

Material Examined. California: BLM-AHF specimens.--1 specimen; OCSD 8606, Sta. 5 (rep 3),

I 595 (repm, January 3); OCSD1986, Voucher; 1 specimen; surveyOC9382, SD 85Sta. 01,37Sta. (rep211), (rep551), m;4620m,Jan211993. Aug 1985, 1 specimen; OCSD, Sta.

Description. Trophosome: Polyps solitary, small. Hypostome with two whorls or cycles of tentacles.

I Aboral tentacles moniliform; about 10 in single whorl. Oral tentacles capitate; at least 4 in number. Growth buds present on underside of tentacular area; appear as swollen bulges directed downward. Perisarc thin,

often with pattern of tings formed by small pits; continuing past end of body to form long thin tendrils or

i anchor filaments; tendril ends often wraping around sand grains. Gonosome: Medusoid buds develop on small stalks on hypostome between oral and aboral rings of

tentacles.

!

I h'

n } A

| ::.:

Ii

!1 o .

!

Figure 1.4. Euphysa young specimen; sp. A: A.D.endetails tire polyps of a ,tentacle naturalshowing size; B. adrinugsltsopecim f nematocysts en with de(ovrigin elopainglillustrations). medusa buds; C.

I 13
!

n

Medusae: Free medusae unknown, i

Nematocysts: unknown.

lie

Distribution. Eastern North Pacific Ocean, central and southern California. Depth range from 4- i

100 m. Recorded in both Los Angeles Harbor and Marina del Rey, and from about 400 m depth on the Santa

w

Rosa Plateau adjacent to the Santa Cruz Basin. Is

Biology. Widely distributed on soft sediment bottoms off central and southern California. Presumed I

to feed on either suspended particulate matter or meiofauna.

Remarks. Ljubenkov (inFauchald and Jones, 1976) previously identified this species as Corymorpha i

aurata because it appeared similiar to the European species. The species does not belong in the genus

Corymorpha and asit lives in southern California, we are treating it as a distinct species until more information

is available. Norenburg and Morse (1983) recently reviewed the genus Euphyoa. I

It

Euphysora bigelowi Maas, 1905

Figure 1.5 I

Euphysora bigelowi Maas, 1905: 6-7; pl. l(figs. 1-3).--Vanhoffen, 1911: 197; 1913: 7; pl. l(fig. 3).-- i

Browne, 1916: 173-174.--Kramp, 1928: 34-35, figs. 8-12; 1948: 20; 1953: 262; 1958: 340; 1961: U

39.--Chiu, 1954: 51-52.---Chow and Huang, 1958: 174, 189; pl. l(fig. 3).--Ganapati and

Nagabhushanam, 1958: 92-94.---Cairns et a/.,1991:20, i

Corymorpha bigelowi: Hartlaub, 1907: 80.---Sassaman and Rees, 1978: 485-495; figs. l(a-d), 2(a-d), 3(a-e). Hi

Steenstrupia bigelowi: Mayer, 1910: 36; fig. 9.--Lele and Gae, 1935: 91.--Bal and Pradhan, 1952: 76. I

Euphysa bigelowi: Uchida, 1927: 188-189; text fig. 28, pl. 10 (fig.3).--Uchida, 1947: 300.--Yamazi, 1958: .i

135.

Material Examined. California: Santa Maria Basin, off Purisima Point, 2 specimens; 34°41.4'N, I

120°57.9'W, 410 m; coll. Battelle, MtVAloha, cruise 1-3, Sta. R-6 (rep 2), Box Corer, May 1987 (SBMNH).

w

Description. Trophosome: Solitary, erect, up to 13 mm high by 1mm wide. Hypostome terminal, 3 t

mm high, flask-shaped, with 2 whorls of tentacles; distinctly demarcated from hydrocaulus by circular I

groove or diaphragm. Aboral tentacles up to 5 mm long, filiform, 15-20 in number in single whorl; oral

tentacles much shorter, slightly thickened at tips, 30-35 in number in several irregular rows; in life tentacles

not very contractile. Hydrocaulus thick and parenchymatous, more or less cylindrical; papillae absent. Perisarc i

thin, flexible, enclosing hydrocaulus from thickened annular or ectodermal ring just below diaphragm to

base; often extending beyond base of hydranth in form of thin tube. Endodermal canals well developed and n

easily visable through hydrocaulus. Base slightly swollen, width 1.5 mm; anchoring rootlets or filaments •

present, arising from endodermal canals.

aim

Gonosome: Gonophores produce free medusae. Pedicels (blastostyles) short, 1-2 mm long, inflated, /s

not highly branched; arising just distal to aboral tentacles. Clusters of medusae buds present, a single bud at I

the terminus of each branch.

Medusae: Elongate, bell-shaped, up to 5 mm high. Nematocysts absent on exumbrellar surface. •

Apical projection conical with small papillae on surface. Apical canal well developed, variable in length, II

extending about 65% way to tip. Radial canals 4 in number. Manubrium tubular, extending to velar opening;

mouth simple, circular, armed with nematocysts. Marginal tentacles 4 in number, allhollow. Single primary I

tentacle as long as bell, unbranched, with club-shaped terminal nematocyst bulb plus 8-9 subterminal, adaxial •

nematocyst bulbs. Secondary tentacles, short, simple, without nematocyst bulbs but covered with scattered

!

14 !
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i D --,,-m

di _ rc

iii ( pt

,42x

E (_ G :-a_ -st

I •

ee C

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I l "

!

F_'e 1.5. Euphysora bigelowiMaas, 1905:A. composite drawingof mature polyp;B. newly released medusa;

I E. C.adultmedusa;D-H:cnidome,undischargednematocysts,D.small stenotele(aboraltentacle,polyp);

large stenotele (aboral tentacle); F. desmoneme (aboral tentacle); G. microbasic mastigophore

(oral tentacle, polyp); H. anisorhiza (secondarytentacle, adult medusa) (after Sassaman and Rees,

i 1978).

I nematocysts; Ocelli tentacles lateral to primary tentacle twice as long as tentacle opposite to primary tentacle. absent. Gonads surrounding manubrium. Medusae colorless except for pale yellow nematocyst bulbs

on primary tentacle.

I Nematocysts: Hydroids, Desmonemes 4-6 x 3.5-4.5 microbasic mastigophores 7.5-8.5 x 3-4

Fan;

lain;stenoteles (small) 7-8 x 4.5-5 _un;stenoteles (large) 13-16 x 8.5-10 pm.--Medusae, Desmonemes 6.5-

9 x 3.5-5.5 pin; microbasic mastigophores 3-4.5 x 8-9.5 btm;stenoteles (small) 7-10 x 7-9 pro; stenoteles

I (large) 11-15 x 9-13 p_m;?anisorhizas 9-11 x 10-12 pan.

Type Locality and Type Specimens. Types not searched. Malay Archipelago.

Vouchers. Deposited by Sassaman and Rees (1978), polyps, USNM 56762 and medusae, USNM

I 56760.

Distribution. Eastern North Pacific Ocean, California to Chile; precise distribution not known.

I Depth Ocean, range Australia; not known. China; andAlsJapan. o recorded Indo-We fromst Pacific the IndiOcean, an Ocean, MalayIndiaArchipelago; , Alphonse Hong Island.Kong; Western Philippines; Pacific

Nicobar Islands; and Palau Islands. (see Kramp, 1961 for review of distribution).

!

I 15
!

II

Biology. Asexual reproduction was observed by Sassaman and Rees (1978). Termed "frustulation" •

by Kramp (1948) the process is similar to transverse fission in which the terminal portion of the base of the

polyp detaches from the hydrocaulus, develops tentacles and within a few days breaks free of the parental am

perisarc. I

Gonosome development occurs about two weeks after larval metamorphosis and medusae are released

about six weeks later (Sassaman and Rees, 1978). The asynchronous development of gonosomes results in I

a prolonged period of medusae liberation. One polyp observed by Sassaman and Rees (1978) produced 34 i

medusae over a period of three weeks. Newly released medusae lived for only a few days in the laboratory.

!

Family Olindiidae

IL

Diagnosis. Zooids growing individually from a stolon. Hydranth with only 1 tentacle. Gonophores I

producing free medusae.

Remarks. This family variously has been designated as the Monobrachiidae or the Oliniadiidae. H

The latter is a variant spelling of the family name.

B

Monobrachium Mereschkowsky, 1877 1

Diagnosis. Colonies reptant; epibiotic on living bivalves and foraminiferans; polyps arising singly

from reticulate stolon. Hydranth without perisarc; with conical hypostome and only 1 tentacle; constricted

at base; stalks not pronounced. Gonophores arising from stolon; medusoids sessile, with 4 radial canals and i

up to 16 tentacles.

II

Monobrachium parasitum Mereselakowsky, 1877 HI

Figure 1.6 I

Monobrachiumparasitum Mereschkowsky, 1877: 226-228; pl. 5(figs. 1-6).--Levinsen, 1893:151.--Stafford,

1912: 73.--Broch, 1916: 42.--Fraser, 1918a: 337; 1918b: 131-138; 1921: 141; 1927: 326; 1937: I

24-25; pl. 2 (fig. 7); 1927: 326; 1937: 24-25; pl. 2(fig. 7); 1927: 326; 1937: 24-25; pl. 2(fig. 7)..-- i

Rylov, 1923: 156.--Vut'fius, 1937: 69.--Naumov, 1957:168-169; text figs. 1-2; 1960 [in 1969]:

547-549; text figs. 394-396---Hand, 1957: 84- 87; figs. 1-2.--Rees, 1967: 219..--Lees, 1986: 106- i

113, 117. III

Monobrachiumparasiticum [sic]: Vagner, 1889: 1-3; 1890: 273-307; pls. 8-9.---Bonnevie, 1899: 51.

Material Examined. California: off Point Arguello, 34_34.35'N, 120°45.18_, 99 m; BSS-58, 5 _•

Jan 1984, USNM; off Point Buchon, 35°05.8'N, 120°42.9'W, 91m; coll. Battelle, M/V Aloha, cruise 1-1,

Sta. R-1 (rep 1), H-S Box Corer, Oct 1986 (SBMNH). D

i

Description. Trophosome: Colonies stolonal with creeping reticular hydrorhiza bearing numerous

(20 or more) sessile hydranths. Hydranths concentrated in posterior region of shell around hinge. Hydrorhiza I

filiform, smooth, extending over surface of shell to margin. In small bivalve species stolonial network

plate-like covering entire shell. Hydrorhiza and part of hydrocaulus enclosed in thin, wrinkled perisarc; gO

terminal portion of hydrorhiza which projects beyond margin of shell without perisarc. Gastrozooids small,

1-3mm high, elongate and fusiform. Pedicel reduced or absent. Hypostome 25-50% as long as hydranth; •

separated from hydranth by deep constriction. Mouth terminally located, surrounded by densely arranged I!

!

,6 !
I

I

i l

I - hy

I

I Figure 1.6. Monobrachiumparasitum Mereschkowsky, 1877:A. colonyon shellof bivalve mollusk,Ax/nops/da

serricata, note presence of dactylozooids at the margin of the shell; B. enlarged section of colony

showing cluster of feeding and reproductive zooids; C. gonozoids (Figure A. after Hand, 1957; B.

I after Nanmov, 1969;C. afterFraser, 1937).

I

nematocysts. Single filiform tentacle present, projects at 90° to body; length when extended exceeding

i length of hydranth; entire surface covered with scattered nematocysts. Capitate dactylozooids present where hydrorhiza projects beyond margin of shell; appearing as globular concentrations of large nematocysts.

Gonosome: Gonophores arising singly fl'om hydrorhiza on very short stalks; completely invested

I with medusopedsarc. id produced Numberperofgonophore; gonophores shape limited, globular, typically3 mm2, buthighnever by more 1.5 mmthanwide; 4 pervelum colony.well Onlydeveloped; 1 sessile

radial canals 4 in number; manubrium rudimentary, mouth absent; tentacles short, 16 in number, located

between velum and ectodermal fold. Gonads extending as 2 longitudinal folds along sides of each radial

i canal.

Nematocysts: Not described.

I Type Locality and Type Specimens. Unknown.

Distribution. Eastern North Pacific Ocean, Alaska to Baja California, Mexico. Depth range 30-

200 m. Also recorded from the Western North Pacific Ocean; Japan, Okhotsk Sea; North Atlantic Ocean;

I Canada, and Greenland. Arctic Ocean; Spitsbergen, Barents Sea, Chuckchi Sea, Kara Sea, and the White Sea.

!
!

!1

Biology, Monobrachium is an epibiont that is restricted in its occurrence to the shells of live, deep- I

water bivalve mollusks and arenaceous foraminiferans. In the Eastern North Pacific Ocean it has been

found on the shells of the following bivalves: Thyasiridae, Axinopsida serricata (Carpenter) [=A. viridus i

(Dall)]; Lasaeidae, Axinodon redondoensis (Butch) [=Tomburchus or Aligena redondoensis]; Astartidae - II

Astarte crenata (Gray); Tellinidae, Macoma baItica (Litre.) [=T. solidula Pulteney],M. calcarea (Chemnitz)

[=T. lata Gmelin], and M. moesta (Deshayes). It also has been reported to live on the test of the foraminiferan i

Halophragmoides planissimum. The above clam species are small and have short siphons. They live near II

the sediment surface in free sand and soft silt. Although the hydroids have easy access to the shells of these

surface-dwelling clams they are poorly protected from abrasion. In life the polyps are very mobile as are the am

tentacles. Given the location on the shell they can intercept food in the clams respiratory curents. II

Rees (1967) postulated that Monobrachium may be similar to other commensal hydroids such as

Proboscidactyla. When the host dies the hydrorhiza become activated and the club-like dactylozooids at •

the margin of the shell elongate and increase in number. In Proboscidactyla the free ends of the hydrorhiza iN

are delicate and sticky and easily torn off even in weak currents of water. When broken off they are capable

of attaching to a new host and thus may serve to propogate the species asexually. HI

i On the Hueneme Shelf, Monobrachium has been observed in moderate densities in spring, winter

and summer (Lees, 1986). Lees (1986) also noted in his study that although the clam Axinopsida serricata

occurred in depths as shallow as 27 m, the hydroid only was found at depths greater than 50 m. He suggested

that the upper limits of the hydroid distribution must be controlled by some factor other than substrate. i

In BLM-OCS samples, populations of Monobrachium parasitum were concentrated at outer

continental shelf stations between 31-491 m where the substrate is fairly coarse with sand, shell fragments •

and pebbles. The highest densities were at 150 m in depth. At stations close to the Patton Escarpment nearly i

100% ofAxinodon redondoensis were infected whereas at stations in the San Pedro Basin the host clam was

present but no Monobrachium were observed. At deeper stations there was a tendency for the clams to

become overgrown with a rusty coating which apparently inhibits growth of the hydroid. Although UP

Monobrachium appears to be present at all times of the year adult colonies in a reproductive state were

noted only in summer samples. In the first year of the BLM-OCS study of the bivalves collected 35 of 81 I

(43 %) specimens of A. redondoensis and 22 of 173 (13 %)Axinopsida serricata were infested with colonies U

of M. parasitum.

!

Family Campanulariidae Johnston, 1836

!11

Diagnosis. Hydrotheca campanulate, never sessile, never adnate to or immersed in the stem or i

branches; diaphragm always present; hydranth with trumpet-shaped proboscis. Gonophores producing

sporosacs or free medusae; when produced medusae usually have Iithocysts; gonads located on radial canals. I

Genus Clytia Lamouroux, 1812 II

Diagnosis. Colonies arising from an unbranched or irregularly branched stem. Gonophores producing

free medusae, somewhat spherical, with 4 tentacles at time of liberation.

!

!

!
!

I Clytia denticulata (Clark, 1876)

Figure 1.7

I Campanularia denticulam Clark, 1876: 213, pl. l(fig. 4).--Nutting, 1901: 189; 1915: 36.--Torrey, 1902a:

51, pl. 4 (fig. 34).--Fraser, 1911: 29-30; 1913: 150; 1914: 32; 1937: 60, pl. 12 (fig. 50); 1948:

I 203.---Stechow, 1913: 122.

Clytia denticulata: Cairns et al., 1991:23.

I Material Examined. California: Western Santa Barbara Channel, Sta. BRC-02 (USNM).

Description. Trophosome: Polyps small; arising singly from reptant stolon, occasionally branched.

Pedicel of varying lengths; annulated at base and just below hydrotheca. Hydrotheca deep bodied, with

I about 15 sharp teeth.

Gonosome: Gonotheca arising directly from stolon; opening as wide as gonotheca; collar absent.

I Pedicles Nematocysts: short, annulated. Not described.

Type Locality and Type Specimens. Presumed to be in the collections of YPM, not searched. Port

I Etches, Alaska, 10-18 fm [18-33 m].

Distribution. Eastern North Pacific Ocean, Alaska to Tenacatita Bay, Mexico. Depth range, 15-75 m.

i Biology. Biology information not reported in the literature.

I

I

--at"

I ,tat/.. di

i

!

I

i Figure 1.7. Clytia denticulata (Clark, 1876): A. entire colony, naturalsize; B. hydrotheca andpedicel showing

characteristic annulations; C. detail of hydrotheca to show diaphragm (Figures A-B. after Fraser,

1937;C. after Torrey, 1902).

I
!

II

Genus Laomedea Lamouroux, 1812 •

Diagnosis. Colonies typically stolonal, seldom erect and branched; stolons of hydrorhiza not

anastomosing. Hydrotheca campanulate, walls with unthickened perisarc, not abruptly everted distally; il

demarcated from pedicel by annular perisarc thickening; diaphragm absent. Gonophores fixed sporosacs.

g

Laomedea altitheca (Fraser, 1948) I

Figure 1.8 l

Campanularia altitheca Fraser, 1948: 202, pl. 23(figs. a-d). i

Laomedea altitheca: Cairns et al., 1991:23. 1

Material Examined. California: holotype (SBMNH) (see below); California Sta. BRC-01 (USNM).

Description. Trophosome: Colonies small; height about 2 cm; stem fascicled, unbranched. •

Hydrothecae large; with 10 castellate teeth. Pedicels long arising from fascicled stem; 2-3 annulations Ii

below base of each hydrotheca.

!

!

I

I

ar he i

11 ,

Figure 1.8. Laomedea altitheca (Fraser, 1948): A. entire colony, natural size; B. three hydrothecae with long I

pedicels; C. fascicled stem hydrocaulus with hydrotheca and gonangia; D. strongly corrugated

gonangium(afterFraser, 1948). i

20 !
!

I Gonosome: Gonangia tubular, truncate at free end, tapering at base; arise directly from stem; walls

vary from slightly wavy to strongly annulated.

I Nematocysts: Not described.

Type locality and Type Speeimeus. Holotype SBMNH [AHF 33]; San Clemente Island, California,

south of Pyramid Cove, 55-69 fms [100-126 m], coll. R/V Velero 111,Sta. 1012-39.

i Paratypes not located.

Distribution. Eastern North Pacific Ocean, Santa Maria Basin, California to Cabo San Lucas, Baja

I California del Sur, Mexico. Depth range, 70-165 m. Biology. Nothing is known about the biology of this species.

Remarks. This genus as well as the entire family are critically in need of revision. I

I Family Campanulinidae Hincks, 1868

Diagnosis. Colonies stolonal or erect, branched or unbranched. Hydrothecae pedicellate or sessile;

always operculate; opercutum formed of converging segments. Hydrants with conical proboscis.

I Gonophores producing sporosacs or free medusae.

I Key to Species Campanulinidae

I 1A. Nematophores absent; operculum with 8-9 segments ........................................... CalyceUa syringa lB. Nematophores present; operculum with 12 segments .......................................... Oplorhiza gracilis

I Calycella Hincks, 1864

Diagnosis, Colonies stolonal, not erect nor branching. Hydrotheca arising directly from stolon,

I walls Gonophores of unthickened fLXedspomsacs; perisarc;arising margin directly distinfromct; separated stolon; extruded from pintoedicacrocysts. el by thick, annulated perisarc.

Remarks. In some treatments this genus is placed in its own family, the Calyeellidae.

I

Calycella syringa (Linnaeus, 1767)

I Figure 1.9

Sertularia syringa Linnaeus, 1767:1311.

I Calycella syringa: Clark, 1876a: 217, pl. 6(fig. 25).---Calkins, 1899: 358, pl. 4(fig. 20A-C), 6(fig. 20D).--

Nutting, 1889: 741; 1901: 194-195.--Torrey, 1904b: 20.--Fraser, 1911: 42, pl. 3(fig. 6); 1913:

152; 1914: 156; 1935: 144; 1936: 124; 1937: 91-92; 1940: 40; 1948: 216.--Bmch, 1918: 32-34.--

et al.,1991

Cairns :21.

Calicella syringa: Stechow, 1923: 134.--Naumov, 1960 [in 1969]: 332, text fig. 198.--Vervoort, 1946:

i 216, text-figs. 92, 93; 1972: 36.

Material Examined, California: Western Santa Barbara Channel: Sta. BRC-01(USNM).

!

m 21
!

II

Description. Trophosome: Colony composed of solitary polyps, arising directly from creeping •

hydrorhiza; hydrorhiza smooth, not reticulated. Hydrotheca tubular; margin distinct, forming rim,

reduplicated; operculum composed of 8-9 segments. Pedicels long or short, always armulated.

It Gonosome: Gonophores oval or obovate; borne on stolon; pedicel with 2-3 annulations; sporosacs

extruded into acrocyst. lira

Nematocysts: Not described, l

Type Locality and Type Specimens. Unknown, probably not extant.

Distribution. EasternNorth Pacific Ocean, Alaska to Cedros Island, Baja California, Mexico. Depth []

range, intertidal, 0-300 m. Also reported from the Western North Pacific Ocean, Northern Asia and Kara

Sea. Eastern North Atlantic Ocean, Great Britain, Norway, Iceland, and Greenland. Western North Atlantic

Ocean, Canada (Labrador).

It Common Name. Creeping bell hydroid.

Biology. Reported to be an epibiont which typically grows attached to the stems of other hydroids n

(e.g., Tubularia, Hydrallmania, Lafoea), algae and other similar living substrates (Clark, 1876c; Calkins, 1

1899). In

Remarks. A small species often confused with Oplorhiza gracilis. Cornelius (1978) reviewed the III

I

genus.

!

_.._jop m

gp/

!

!

B C

!

Figure 1.9. Catycella syringa (Linnaeus, 1767):A. entirecolony, natural size; B. enlarged aspectof colony; C. I

single hydrotheca showing characteristicshort, annulated pedicel; D. stolon with hydrothecae and

gonophores (Figure A. after Fraser, 1937; B. after Calkin, 1899; C. after Fraser, 1911; D. after

Nanmov, 1960 [in 1969]). I
!

!

Genus Oplorhiza Allman, 1877

I Diagnosis. Colonies usually stolonal, occasionally erect polysiphonic stem.

and branched with

Hydrotheca pedicellate, radially symetrical, deeply campanulate, cone-shaped, widest at distal end; thin

diaphragm present. Operculum cone-shaped, distinctly separated from hydrotheca. Nematophores solitary

I or aggregated, occuring on hydrorhiza or on hydrorhiza pedicels. Gonophores

both and unknown.

Remarks. In the past this genus variously has been placed in the Family Lovenellidae or

i Campanulinidae gonophores. In Cairns becauseetitalcould . (1991)notitbehasestablished not been whether allocatedspecies to a family produceandfreeitsmedu classification sae or havstatus e fixedis

indicated as uncertain. The genus resembles Egmundella Stechow, 1921 but differs in having fusiform

i instead of cone-shaped hydrotheca.

Oplorhiza gracilis (Stechow, 1921)

I Figure 1.10

i Egmundella 101).--McCormick, gracilis Stechow,1965: 1921:142.--Lees, 226; 1923: 124.--Fraser, 1986: 106.---Cairns 1935: etal 144; 1936: 124; 1937: 95; pl. 19 (fig. ., 1991: 22.

Oplorhiza gracilis: Naumov, 1960 [in 1969]: 339.

I Material Examined. California: Santa Maria Basin, off Purisima Pt., 34°41.4q_1,120°57.9'N, 410

m, coil Battelle, M/V A/oha, cruise 1-3, Sta. R-6 (rep 2), Box Corer, May 1987 (SBMNH).

I branchedDescri hydrorhiza. ption. Trophosome: Pedicels unbranched Colonies small, or withzooids 1-2 1-2 branches; mm high; annutated arising fromat base; creeping, passingirregularly almost

imperceptibly into hydrotheca. Perisarc moderately thin with irregular wrinkles. Hydrothecae turbinate

I whydrorhiza; ith sinuouscontracted margins; opercul at base utomformwithshort 12 segstalk. ments. Nematophores spherical; present on pedicels and/or

Gonosome: Gonophores unknown.

I Medusae: Unknown.

Nematocysts: Not described.

I Type Di Locality and Type Specimens. Not searched. Not traced. stribution. Eastern North Pacific Ocean, Queen Charlotte Islands, British Columbia, Canada to

Santa Maria Basin, California. Depth range 20-100 m.

I Biology. Very rare in its occurence. The few reports of this species have noted its occurrence on

shells of gastropod mollusks (Neptunia lyrata).

Remarks. Some investigators have placed this species in the genus Egmundella which it closely

i resembles. Except for the presence of nematophores the species also closely resembles Lovenella producta.

!

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i

i

--o0 |

I

o I

I

I

I

Figure 1.10. Oplorhiza gracilis(Stechow, 1921):A. entire colony,naturalsize; B. enlarged sectionof colony; C. I

nematophore (after Fraser, 1937).

I

Family Lafoeidae A. Agassiz, 1865

Diagnosis. Colony stolonal or erect and branching. Hydrothecae tubular, even rimmed; without i

operculum; with or without diaphragm; pedicellate or sessile, adnate in some species. Hydranth with conical

hypostome. Gonothecae aggregated, as scapus or coppinia. Sexual generation a fixed sporosac. I

Genus Acryptolaria Norman, 1875

Diagnosis. Colonies erect, more or less alternately branched; hydrocaulus andbranches polysiphonic i

with 2 longitudinal rows of alternate hydrothecae. Hydrothecae sessile, tubular, adnate to axial tube or

ihymmersed d in accessory tubes basally, curving outwards and becoming free distally; diaphragm absent; I

rothecal base indistinctly demarcated by ring of small desmocytes. Reduced hydtothecae occasionally

present on accessory tubes. Nematotheca absent. Gonophores fixed sporosacs. Gonothecae aggregated;

coppiniaeRemarks. with or wSiteehourece t mntodifi disecdusshyiondroofthethecalgtuenubes. s in Calder (1991). I

i
I

I Acryptolaria pulchella (Allman, 1888)

I Figure 1.11

Cryptolaria pulchella Allman, 1888: 40.--Fraser, 1925: t72.

I Acryptolariapulchella: Nutting, 1927: 210.--Fraser, 1937: 116, pl. 24 (figs. 31a-c); 1938c: 134.

Material Examined. California: Sta. BRA-02 (USNM).

I fascicled;Descr nodesiption. absent; Trophprimary osome:branches Colony 5-6 pinnate, cm tall; irregularly with stouarranged t, fascicledor stem; looselybases alternate; of bransecondary ches also

branching regularly alternate. Hydrothecae curved outward, bases more or less adnate; same size throughout;

i alternate Gonosome: and regularlyUnknown. placed.

Nematocysts: Not described.

I Type Locality and Type Specimens. Not searched. Pacific Ocean (Challenger Reports).

Distribution. Eastern North Pacific Ocean, San Francisco Bay, California to the Galapagos Islands,

i Ecuador.BioDepth logy.range, Nothing10-200 has been m. reported in the literature about the biology of this species.

i Remarks. Fraser (1944: 213, pl. 42(figs. 193a-c)) reported this species in the Western North Atlantic

I Ocean but the drawings do not match the description provided in his 1937 book.

I

I

I

!

I

I

!

Figure 1.11. Acryptolariapulchella (Allman, 1888): A. entire colony, naturalsize; B. portion of fascicled main

I stem; C. portionof simple stem or branch (afterFraser, 1937).

!iI 25
!

n

Family Lovinellidae Hincks, 1868 •

Genus Lovenella Hincks, 1868 i

Diagnosis. Colony branched or unbranched; hydrotheca mrbinate; operculum sharply def'med by II

sinuous margin on tube of hydrotheca; nematophores absent. Gonophores borne on stems, producing free, l

bell-shaped medusae with 8 tentacles in 2 sets and 4 lithocysts.

LoveneUa producta (G.O. Sars, 1873) I

Figure 1.12 I

CaIycellaproducta G. O.Sars, 1873: 30.--Hincks, 1874:134

Lovenellaproducta: Fraser, 1911: 44-45, pl. 3(figs. 7-10); 1913: 152; 1914: 159; 1937: 96, pl. 19 (figs. 102a- i

c); 1938a: 40; 1938b: 109; 1948: 219.

Material Examined, California: Santa Maria Basin, off Pt. Buchon, 1colony, 35°05.5q_1,120°53.4'W, []

161m; coll. Battelle, M/V Aloha, cruise 1-1, Sta. R-2 (rep 2), Box Corer, Oct 1986 (SBMNH). |

Description. Trophosome: Zooids typically arising singly from hydrorhiza, occasionally one or

more zooids branching off another pedicel. Hydrotheca turbinate; margin appears distinctly scalloped or am

sinuous due to curved bases of opercular segments. Operculum with 12 or more pie-shaped segments which n

interlock to form cone; often inverted. Pedicels long, length to 6 ram; always annulated at base. Demarcation

between pedicel andhydrotheca gradual, not distinct; in region of transition hydrotheca often slightly turbinate •

or twisted. Nematophores absent. il

Gonosome: Unknown.

Nematocysts: Not described, i

m

Type Locality and Type Specimens. Not searched. Not traced.

Distribution. Eastern NorthPacific Ocean - Vancouver Island, British Columbia, Canada to Anacapa II

Island, California. Depth range 90-700 m. Also reported from the Western North Atlantic Ocean - Maine, il

USA.

Biology. Fraser (1913) recorded this species growing on tubes of serpulid annelids, i

Remarks. This species is nearly indistinguishable from Oplorhiza gracilis from which it differs in

the complete absence of nematophores. Fraser (1937) feels that many of the reported instances of this

species are actually O. gracilis, especially from the Vancouver Island region. Lovenella nodosa, reported by i

Lees (1986) from the Hueneme Shelf may actually be L. producta.

II

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26 !
I

I

I

I Figure 1.12. variation Lovenellainproducta sizes,note(Gcharacteristic .O. Sars, 1873): annulations A. entireonlycolony, atbasenatural of pedicels; size;B-D D. branched . single colony zooidsto[Figures show

A, C, D, after Fraser, 1937; B. after Fraser, 1911].

!

Family Aglaopheniidae

I Key to Species of Aglaopheniidae

I 1A. Colony small, slender, height to 1 cm; hydrotheca present on stem ..................... Cladocarpus gracilis

lB. Colony large, plumose, height to 12 cm; hydrotheca absent on stem ....... Cladocarpus vancouverensis

I Genus Cladocarpus Allman, 1874

I nematophores Diagnosis. short.Hydr Gonophores othecae deepnotwisituated th smoothwithin margincorbulae; or crenulated associated with low, withblslender unt teeth.branches Mesial

(phylactogonia) which arise from stem at base of hydrocladia. Phylactogonia bearing nematophores but

I lacking hydrothecae.

i
I

Cladocarpus _cilis Fraser, 1948 I

Figure 1.13

Cladocarpus gracilis Fraser, 1948: 269-270, pl. 36(figs. 39a-d),

Material Examined. California: holotype (SBMNH, see below). Additional material examined: n

BLM/OCS Baseline Survey--specimens from the following BLM station localities: 143, 145, 148, 152, U

155, 164, 175, 220.

Description. Trophosome: Colony delicate, small, 10 mm high; simple, slender, arising directly I

from creeping stolon; holotype unbranched but other colonies examined alternate and pinnately branched.

U

Distal half of hydrocaulus thecate. Nematophores in single wandering row on face of hydrocaulus;

nematophores originating below insertion of first chevron-shaped segment. Hydrothecae long, slender, length i

more than twice width; diameter increasing from base to margin; prominent medial tooth on face of margin,

m

remainder of margin irregularly waved or slightly toothed. Up to 9 hydrothecae present on face of stem n

between distal chevron and origin of branch pedicels; hydrothecae absent on stem face distal to origin of I[

first branch pedicel. Supracalycine nematophores much smaller than length of hydrothecae; mesial I

but

nematophores not adherent to face of hydrothecae, attached to internode below base of hydrotheca. !

-P |

,

i o'

!

D E I

1

Figure 1.13. Ctadocarpus gracilis Fraser, 1948:A. entire colony,naturalsize; B. first hydortheca andportion of m

proximal stem, lateral view; C. medium portion of stem, lateral view; D. two hydrothecae, lateral

view; E. arrangementofhydrothecae andnematophoresonlowerstem,stylized, enfaceview (Figures 1

A-D. after Fraser 1948;E. original).

28 I
!

I Gonosome: Gonophores and phylactocarps located on distal parts of stem.

Nematocysts: Not described.

I Type Locality and Type Specimens: California, holotype - SBMNH [AHF 131]. California, Santa Catalina Island, bearing 2.8 mi WNW of Long Point, 64-88 fm [117-161m]; coll. R/V Velero II1,Sta. 1307-

41.

I Distribution. Eastern North Pacific Ocean: Santa Rosa Ridge and Santa Catalina Island shelf

California. Depth range, 90-268 m.

I Biology. R Information on the biology of this species is not available in the literature. emarks. Fraser (1948) described this species from an incomplete single specimen which lacked

the stem above the hydrothecae. He concluded that the species should perhaps be placed in the genus

I Antene enoughUaspecimens , in which aretheavailable hydrocladiato seearisethatdirectly a pinnatefromcolony a creeping is formed stolon.similiar In thetomaterial other species recentlyinexamined, the genus

Cladocarpus. Cladocarpus gracilis is the only species in the genus that has hydrothecae on the stem.

I Cladocarpus vancouverensis Fraser, 1914

l Figure 1.14

Cladocarp_ vancouverensis Fraser, 1914: 204; 1937: 182-183, pl. 41(figs. 221a-d); 1946: 93, 408; 1948: 273.

I Material Examined. California, offMorro Bay, Sta. BRA-027 (USNM).

Additional material examined: Several specimens from near San Miguel and Santa Rosa Islands;

I BLM/OCS Baseline Survey (1975-77), Stas. 28 (129 m), 38 (133 m), and 155 (163 m). Description. Trophosome: Colony large, plumose, height to 12 cm; stem simple, unbranched.

Hydrocladia regularly alternate, branches on 2 sides not in same plane; divided into regular internodes.

I Hydrothecae adnate throughout; deep-bodied, deeper than wide; distinct large medial tooth on face of margin, remainder of margin weakly crenulated with small, wavy teeth. Two supracalycine nematophores, much

smaller than hydrotheca length; mesial nematophore projected outward, base adnate, distal portion free.

I Nematophores on front of stem changing from widely spaced pairs to very narrowly spaced pairs below origin of first chevron-shaped segment. Chevron-shaped segments present, 2-4 below origin of branch

pedicels. Septal ridge present at base of supracalycine nematophore, at base of hydrotheca and 2 others

I regularly placed between these.

Gonosome: Gonophores borne on front of stem; ovoid with rounded distal end. Protected by 2-

pronged phylactogonia, tips of each prong typically bifurcate.

I Nematocysts: Not described.

Type Locality and Type Specimen. Presumed to be in collections of BCPM (Arai, 1977); not

I searched.Dis[Canada. British Columbia], Vancouver Island region. tribution. Eastern North Pacific Ocean, Queen Charlotte Island, British Columbia, Canada to

San Benito Islands, Baja California, Mexico. Depth range, 25-400 m.

I Biology. Unknown.

Remarks. The colonies of Cladocarpus vancouverensis resemble those of the genus Aglaophenia

but axesmaller, more delicate andhave fewer hydrothecae onthe branches. Three other species of Cladocarpus

I are reported to occur in southern California: C. pinguis is shorter and stouter; gracilis (incompletely by

C.

Fraser from an incomplete specimen); and an undescribed species (sp. A sensu Ljubenkov) which has been

collected so far only from the Encina Ouffall. !

i I 29
I

Figure 1.14, Cladocarpus vancouverensisFraser, 1914: A. entire colony, natural size; B. portion of hydrocladium, I

lateralview; C. hydrotheca,enfaceview; D. gonophores and phylactogorda;E. arrangementof

nematophores on lower stem, note absence of hyrotheca, stylized, enface view (Figures A-D. after I

Fraser, 1937;E. original). I

I

Family Plumulariidae L. Agassiz, 1862

ml

Diagnosis. Hydrothecae growing only on one side of the branches (hydrocladia); sessile, more or I

less adnate; operculum absent; nematophores always present. Gonophores producing fixed sporosacs; often

protected by special modifications of the branches. I

Key to Species of Plumulariidae III

1A. One mesial nematophore on each thecate and athecate internode; intemodes with conspicuous septal

ridges .............................................................................................................. Plumularia corrugata I

1B. Two mesial nematophores on each thecate and a/hecate intemode; intemodes smooth, without sepatal

ridges ...................................................................................................................... Plumularia exilis

!

I

30 I
I

I Genus Plumularia Lamarck, 1815

Diagnosis. Colonies erect; hydrocaulus typically unbranched; hydrocladia unbranched, pinnately

I arranged; each branch with hydrothecae on only 1 side. Hydrothecae sessile, more or less adnate; margin entire. Nematophores present, movable. Gonophores fixed sporosacs; without protection.

i North Remarks. This is a large and confusing genus with well over 30 species described in the Eastern Pacific. The genus is badly in need of revision because the extent of variability of characters such as

the number of nematophores per internode is not known.

I Plumularia eorrugata Nutting 1900

Figure 1.15

! .

PlumuIaria corrugata Nutting, 1900: 64, pl. 6(figs. 1-3).--Fraser, 1911: 82; 1914: 205; 1935: 145; 1937:

i M186-187, aterial pt. Exam42 i(figs. ned. 225a- California: d; 1938a:San63;Diego, 1938b:OCSD 111; 1938c: 8503, 136; Sta. 371948: (rep276. 5), 61 m; 22 Oct 1985

[several colonies].---Oregon: USNM BLM-OCS Voucher, Tanner Bank; Sta. 816.

I "

!

I gP

, /

I "f\n

| Bc D

I Figure 1.15. Plumularia corrugata Nutting, 1900: A. entire colony, natural size; B. portion of main stem and

I hydrocladia; C. male gonphores;D. female gonophores(after Fraser, 1937).

!

I 31
!

i

Description. Trophosome: Colonies medium size, height to 2 cm. Stem simple, arising directly •

from rootlike hydrorhiza; divided into regular intemodes; branches arising from small process atdistal end

of each internode. Branches alternate, all in the same plane; with 5-10 polyps per branch. Cauline intemodes •

marked by constriction; septum not obvious. Thecate and athecate internodes alternate on hydrocladia; first I

intemode after branch arises very short and non-hydrothecate. Two supracalycine nematophores and one

subcalycine mesial nematophore on each thecate intemode; single mesial nematophore on all but first athecate •

intemodes; single mesial nematophore on each cauline internode on side opposite branch pedicel; single

nematophore in each branch axil.

Gonosome: Gonophores attached to branch pedicels. Female gonagium elongate, three times as I_

long as broad; ovoid, distal end truncate. Male gonangium elongate; distal end often constricted in bottleneck

shape.

Nematocysts: Not described. 1

ID

Type Locality and Type Specimens. Syntypes - USNM 18609; USNM 18610; MSUI 11721;

MSUI 11722. 10 mi east of Petros Island.

Distribution. Eastern North Pacific Ocean - San Juan Archipelago, Washington to the Galapagos i

Islands, Ecuador. Depth range 60-90 rn.

Biology. Information on the biology of this species not reported in the literature. I

Plumularia exilis Fraser, 1948

Figure 1.16 i

Plumularia exilis Fraser, 1948: 277-278, pl. 38(figs. 43a-c). 1

Material Examined. Mexico: holotype (SBMNH see below).-- California: off Pt. Arguello, Sta.

BRA-06.D I escription. Trophosome: Colony medium size, clustered, height to 4 cm. Hydrocaulus thin, sinuous;

internodes elongate with branch pedicel located at distal end. Branching pattern alternate. Proximal internode

of each branch very short, lacks both a nematophore and a hydrotheca; thecate and athecate internodes II

alternate distal to first naked internode. Two nematophores present on branch face of each athecate

hydrocladial intemode; two supracalycine nematophores present on each thecate intemode. Each hydrocauline

intemode with up to 4 nematophores on side opposite of hydrocladia; single nematophore in each branch i

axil.

Gonosome: Gonophores numerous, large, length to 1.25 mm; shape, elongate obovate, tapering at

proximal end, truncate with small opening at distal end; attached to face of branches or in axil of branches. I

Nematocysts: Not described.

Type Locality and Type Specimens. Holotype - SBMNH [AHF 140]; Mexico, Lower California •

[Baja California del Sur], off Cape San Lucas, San Jaime Bank, 75 fm [137 m]; coll. R/V Velerolll, station II

618-37.

Distribution. Eastern North Pacific Ocean, Santa Maria Basin, California to Cabo San Lucas, Baja •

California del Sur, Mexico and into the Gulf of California. Depth range, 10-275 m.

Biology. Information on the biology of this species has not been reported in the literature.

I

!
I Figure 1.16. PlumuIariaexilis Fraser,1948:A. entirecolony,naturalsize; B.portionof colonyshowingbranching

patternanddistributionof hydrothecaeandnematophores;C. gonophores(afterFraser,1948).

I

i Family Sertulariidae Lamouroux, 1812

Diagnosis. Hydrothecae sessile, usually arrangedon bothsides of stem andbranches; either alternate

or opposite; more or less adnate; operculum present. Nemotaphores absent. Gonophores producing fixed

I sporosacs.

I to of Sertulariidae Key Species

I 1A. lB. Hydrothecae Hydrothecae oalternate, pposite, margins margins tridentate; entire or smooth; operculumoperculum with 2 valves with single .................. flap ................................ Dynamena pumila

.................................................................................................................... Abietinaria ................. 2

I

2A. Stem relatively slender, hydrothecae short and thick ............................................ Abietinaria traski

i 2B. Stem stout ....................................................................................................................................... 3

I

I 33
!

|

3A. Cauline nodes rarely, if at all, with an annulus; hydrothecae elongate; gonangia smooth ............... •

........................................................................................................................... Abietinaria pacifica

3B. Cauline nodes with single annulus .................................................................................................. 4 i

4A. Gonangia with strong longitudinal crests or ridges .......................................... Abietinaria amphora •

4B. Gonangia smooth or transversly annulated ..................................................... Abietinaria variabilis l

m

Genus Abietinaria Kirchenpauer, 1884 i

i

Diagnosis. Colony erect, branched or unbranched; hydrocaulus monosiphonic. Hydrocaulus and

hydrocladia with hydrothecae in 2 longitudinal rows. Hydrothecae sessile, arranged in alternate o_ subopposite •

pairs; broad proximally, tapering toward aperature. Hydrotheca margin smooth. Operculum a single flap II

attached to adcauline side of hydrotheca aperature. Gonophores fixed sporosacs.

m

Abietinaria amphora Nutting, 1904 1

Figure 1.17 i

J

Abietinaria amphora Nutting, 1904:119, pl. 34(figs. 2-4).--Fraser, 1911: 58; 1913: 153; 1914: 179; 1935:

144; 1936: 125; 1937: 127; 1948: 235. i

Material Examined. California: off Pt. Conception, 2 reproductive colonies, Sta. BRS-073; San

Onofre, Barn Kelp Bed, 40 ft; Apr 1973 (USNM).

Description. Colony large, height to about 4 cm. Hydrocaulus straight, with regularly alternate II

branching pattern; branches and hydrothecae absent on lower stem; 3 hydrothecae present on same side of

stem between 2 succesive branch pedicels; secondary branching absent; branch nodes and septa very weak. •

Hydrothecae nearly opposite; base and about 50% adjacent side adnate. I

Gonosome: Gonophores typically arranged in 2 longitudinal rows on face of stem or proximal

portions of branches. Gontheca large, fusiform or amphora-shaped, with 4 or 5 longitudinal crests or ridges, i

Nematocysts: Not described.

Type Locality and Type Specimens. Hototype - USNM 19821. Pacific Ocean, 54°18'N, 165°55'W, m

56 fm [102 m]; coll. R/V Albatross, station 2866. i

i

Paratypes- USNM 19900, MSUI 18745.

Distribution. EasternNorth Pacific Ocean, Alaska to Tanner Bank, California. Depth range,intertidal •

to 275 m.

Biology. Information on the biology of this species not found in the literature.

Remarks. The speciesAbietinaria costata (Nutting, 1910) from Alaska closely resemblesA, amphora 1

i

and may be a junior synonym.

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34 !
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i ? !

i ?

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i Figure 1.17. Abietinaria amphora Nutting, 1904: A. entire colony, natural size; B. portion of branch with hydrothecae,lateral view; C. gonophores (after Fraser, 1937).

I Abietinaria pacifica Stechow, 1923

i Figure 1.18

Abietinariapacifica Stechow, 1923: 197. --Fraser, 1937: 134, pl. 29(fig. 154); 1948: 237, pl. 27(figs. 16a-c).

I Material Examined. California: Western Santa Barbara Channel, Sta. BRC-01 (USNM).

Additional material examined: MMS/MEC CARP; south of Point Arena, 1150 ft [350 m]; cruiseTi 11, Sta. SB43 Descri(preptionA), . Trophosome: 29 NovemberColonies 1987. large, height to 9-10 cm. Hydrocaulus simple (monosiphonic)

with very weak nodes and septa. Branching pattern alternate; length of branches to 12 mm; 3 widely spaced

I hydrothecae in branch axil;presesingle nt between joint present 2 succeonsiveeacbran h branch ch pediatcels junction on sameof branch side of colony and branch ; singlepedicel. hydrotheHydrothecae ca present

regularly alternate, not closely packed; elongate shape; base and about 75-85% of adjacent side adnate,

I convex; Gonosome: free side sligGonotheca htly concave; neck absent; margin positioned at right angle to stem.

shape obovate, sharply truncated at distal end, collar absent, surface smooth.

Nematocysts: Not described.

I Type Locality and Type Specimens. Not searched. Not traced.

Distribution. Eastern North Pacific Ocean - Point Arena, California to San Benito Islands, Baja

I California, Mexico. Depth range 50-350 m.

I 35
Figure 1.18. Abietinaria pacifica Stechow, 1923: A. entire colony, natural size; B. portion of stem showing i

origin of branch; C. portion of stemshowing hydrothecae and gonphores (Figures A, C. afterFraser,

1948; B. after Fraser 1937 from originalin Stechow, 1923).

!

Biology. The original fragment which Stechow (1923) described came from the back of a decorator

crab. m Remarks. This species bears a great similarity to Abietinaria traski (Torrey, 1902) and is probably

closely related if not the same. The principal differences are the shape of the hydrotheca, the distance

between successive hydrothecae, and the distance to the first hydrotheca on the branch, m

Abietinaria traski (Torrey, 1902)

Figure 1.19 m

Sertularia traski Torrey, 1902a: 69-70, pl. 9(fig. 83). 1

m

Abietinaria traski (Torrey).--Nutting, 1904:118, pl. 33(figs. 6-11).--Fraser, 1911: 63; 1913: 153; 1914:

182; 1935: 145; 1937: 135-136, pl. 29(figs.156a-b); 1948: 238.--McCormick, 1965: 142-143.

Material Examined. California: off Morro Bay, BRA-27 (USNM). m

Description. Stem long, straight, lower portion free of branches; simple (monosiphortic) with poorly m

developed nodes. Branching pattern alternate; nodes absent on branches. Hydrothecae alternately arranged, 1

rather widely spaced; base and about 50% of hydrotheca adnate, convex; free side very slightly concave.

m

Stem light horn color; branches white. ..

|

36 !
!

! B

I Figure 1.19. Abietinaria traski (Torey, 1902): A. Portionof colony, naturalsize; B. portionof branchwith

hydrothecaeandgonophores,lateralview (afterFraser,1937).

I

Gonosome: Gonophores arising in rows on faces of branches, inserted near bases of hydrothecae;

I gonotheca shape elongate-ovoid, without distinct neck or collar, smooth.

Nematocysts: Not described.

Type Locality and Type Specimens. Presumed to be in the collection of the University of California,

I not searched. USA, Californa, San Pedro.

Distribution. Eastern North Pacific Ocean, Berg Inlet Alaska to San Benito Islands, Baja California

I Mexico. Depth range, 10-400 m. Biology. Information on the biology of this species not reported in the literature.

Remarks. See above remarks under Abietinaria pacifica.

I

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I

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Abietinaria variabilis (Clark, 1876) I

• Figure 1.20 •

Sertularia variabilis Clark, 1876a: 221-222, pl. 8(figs. 40-48), pl. 9(figs. 49-50).--Kirchenpauer,1884: 35-

36, pl. I4(fig. 6).

Thuiaria variabilis: Nutting, 1901: 203. 1

Abietinaria variabilis: Nutting, 1904: 123.--Fraser, 1911: 63; 1914: 183; 1935: 145, 1936: 25, 1937: 137,

pl. 30 (figs. a-c).--Kudelin, 1914: 406, 408, 428, 443, text figs. 140-141,157-158, pl. 3 (fig. 10, pl. l

4 (fig. 1).--Naumov, 1960 [in 1969]: 405-406, text fig. 265, pl. 3 (fig. 3).

Material Examined. SBMNH specimens from California. •

Description. Trophosome: Hydrorhiza lamellar. Hydrocaulus straight or slightly sinuous. Branching

alternate with branches held at roughly a 45° angle; all in one plane; base of branch constricted but without

a clear branch pedicel; branches attached at proximal end of stem internode; sometimes second order i

branchlets present. Hydrothecae in two rows, alternate, base of one about the midpoint of the one opposite;

hydrothecal margin tilted away from branch, sometimes to point of being turned downwards; adjacent side

50 to 90% immersed. 1

Gonosome: Gonophores typically attached adjacent to base of hydrotheca. Gonotheca shape ovoid,

with small stalk, smooth to rugose; up to 10 denticles inside aperature.

Nematocysts: Unknown. I

---- -- i

'

Figure 1.20. Abietinaria variabilis (Clark, 1876): A. Portion of colony, natural size; B. portion of branch with 1

hydrothecae;C. branchlet with gonotheca (Figures A-B. afterFraser, 1937;C. after Naumov, 1960 []

[in 1969]).

!
!

Type Locality and Type Specimens. Presumed to be in the collections of YPM, not searched. Not

indicated.

I Distribution. Eastern North Pacific Ocean, Alaska to southern California. Depth range 25-4000 m, typically 25-100 m. Also reported from Western North Pacific Ocean, Sea of Okhotsk, Bering Sea.

Biology. Unknown.

I Remarks. This species varies greatly in shape and proportion. Naumov (1969) immersedSertularia

anguina Trask, 1857, and Abietinaria tilesii and A. cartilaginea both Kirchenpauer, 1876 in this taxon. If

i this proves to be a valid decision than A. anguina (Trask) should be the senior synonym.

Genus Dynamena Lamouroux, 1812

I Diagnosis. Colonies erect, branched or unbranched; hydrocaulus monosiphonic. Hydrocautus, and

hydrocladia when present, with hydrothecae in 2 longitudinal rows. Hydrothecae sessile, in opposite to

subopposite pairs, occasionally in groups of 2 or more pairs per internode. Hydrothecal margin tridentate;

I median tooth smaller and less than lateral teeth, with 2 valves adcauline valve

conspicuous Operculum 1

usually smaller than abcauline valve and divided into 2 parts by median line. Hydranth without an adcauline

diverticulum. Gonophores fixed sporosacs.

i Remarks. Sertularia differs from Dynamena in that the hydrothecae, which are also bicuspidate,

are alternate rather than opposite. Sertularella, also with alternate hydrothecae, is then distinguishable from

Sertularia because the hydrothecae are tri-or quadri-cuspidate. There are no true Sertularia species on the

Pacific coast.

I Dynamena pumila (Linnaeus, 1758)

Figure 1.21

I Sertularia pumila Linnaeus, 1758: 807.--Pallas, 1766: 130.--Agassiz, 1862: 326, pl. 32; 1865: 141.-

Nutting, 1904: 51-53, pl. l(figs. 1-3).----Broch, 1910: 173,219.--Fraser, 1911: 74; 1913: 175; 1921:

i Dynamena40.--Kudelin, pumila (Linnaeus).--Naumov, 1914: 247.--Vervoort,19601946: [in 1969]: 252. 53, text fig. 38; 356-357, text figs. 219A-C.--

Cornelius, 1979:271-273.

I [for additional synonyms and references see Nutting, 1904]

Material Examined. California: BLM-OCS Baseline Study 1975-1976 specimens.

I Description. Trophosome: Colonies erect; height typically less than 5 cm; arising from creeping hydrorhiza. Base of stem twisted; internodes on stem with 1-3 pairs of opposite hydrothecae; hydrocladia

originate singly or in opposite pairs on intemode below hydrotheca, branches of the second order may arise.

i Hydr adcauline othecae elongate, slender; sessile, base and about 75% of adcauline side adnate; remaining portion of side curved outward; margin bicuspidate, teeth sharp, subequal. Operculum with 2 valves.

Gonosome: Gonophores arise from internode below each hydrotheca. Gonotheca shape vase-like to

I ovoid; with truncated neck or collar, smooth; stem short.

Nematocysts: Not described.

i Type Locality and Type Specimens. Not searched. Not traced.

!

I 39
A B _11

!

Figure 1.21, Dynamena pumila (Linneus, 1758): A. portion of colony near base showing arrangement of

hydrothecae; B. two pairs of hydrothecae; C. gonotheca and pair of hydrothecae (after Naunov, I

1969).Scales = 1 ram. II

I

Distribution. Eastern North Pacific Ocean, California. Depth range m 250 in. Also reported from

Western South Pacific Ocean, New Zealand. Eastern North Atlantic Ocean, Great Britain; Belgium; Italy; m

Denmark; Norway; Iceland; and Greenland. Western North Atlantic Ocean, Canada (Nova Scotia) and |

USA (New England).

Common Name. Sea Oak. II

Biology. The life cycle is illustrated in Naumov (1960 [in 1969]: fig. 38). Free swimming medusae II

are absent and planulae are released directly from the gonophores.

Although typically an intertidal form which grows under stones, it has been found at depths to 250 m. I

lira

Remarks. Naumov (1969) doubts the reality of Clark's report of this species on the Pacific coast

and he restricts its distribution to the north Atlantic and northern Europe. Fraser bases his inclusion of this

species in the Californian fauna on Clark's citation. Both Dynamena desmoides and D. cornicina also occur il

m

in California where they have been collected at depths ranging from 2-1860 m (BLM-OCS Baseline study,

1975-76). i

° I
!

I Family Thyroscyphidae Stechow, 1920

I monopodial Diagnwith osis. Colonies stolonal or erect, arising from a creeping hydrorhiza; growth commonly terminal growing points. Hydrothecae radially to bilaterally symmetrical; pedicellate, or

both sessile and pedicellate in the same colony, or sessile but adnate only at diaphragm. Hydrothecal margin

I presen entire,t.orHydr witha3-4 nts teeth; with operculum an annular offold1, 3basally, or 4 valves, or with and either an abcauline persistentdiverticulum; or shed early;hypostome annular diaphragm conical,

surrounded by a whorl of f'fliform tentacles. Gonophores fixed sporosacs.

i Remarks. Members of this family were originally included in the family Sertulariidae. They were

subsequently removed to the subfamily Thyroscyphinae by Stechow (1920) and the subfamily Parascyphinae

by Splettst6sser (1929). Most authorities currently place them in a separate family (see Calder, 1986, 1991).

I

Genus Symplectoscyphus Marktanner-Turneretsher, 1890

i Diagnosis. Colonies and hydrotheca similar to Sertularella but hydrotheca margin with 3 teeth and operculum with 3 valves. Gonophores fixed sporosacs; gonotheca walls strongly ridged.

Remarks. This genus has been recognized by most recent authors as similar to but distinct from

I Sertularella the basis of the number of teeth the margin of the and the number of

on on hydrotheca operculum

valves.

I Key to Species of Thyroscyphidae

I Hydrothecal annulated; gonotheca entirely with numerous upcurved spines ...................

1A. wall covered

.............................................................................................................. Symplectoscyphus pedrensis

I lB. Hydrothecal wall smooth ....................................................................................................................... 2

2A. Teeth unequal; stem erect, stout; gonotheca with few low relief spines distally aroundopening ..........

I ................................................................................................................ Symplectoscyphus turgidus

2B. Teeth equal; gonotheca annulated with distinct ridges ........................... SympIectoscyphus tricuspidata

I Symplectoscyphus pedrensis (Torrey, 1904)

Figure 1.22

I Sertularella conica non Allman, 1877.--Torrey, 1902a: 60-61, text figs. 19-21.

i Sert Symplectoscyphus ularellapedrensispedrensis Torrey, 1904b: 27.--Fraser, 1911: 70; 1914: 191; 1937: 155; I938: 110; 1948: 243-244. : Cairns et al., 1991: 27.

Material Examined. California, San Diego.-- I colony; OCSD 8942, Sta. 12 (rep. 2), 60 m; 20

I January 1989.-- several colonies; OCSD 052, Sta. T-6(haul 4), 36 m.

Description. Trophosome: Colony erect; small, usually less than 2 cm; with one or more branches,

I arising directly from creeping stolon. Hydrocladia with 1 hydrotheca per intemode, hydrothecae alternate.

Intemodes fairly short with hydrothecae at distal end; nodes weak but obvious. Hydrotheca with tricuspidate

margin; walls smooth or with faint annulations, stronger distally.

!

i 41
i

I

'

_'" B I

F |

Figure 1.22, Symplectoscyphuspedrensis (Torrey, 1904): A. entire colony, natural size; B. portion of stem and i

smooth hydrothecae; C-D. details of sculptured hydrothecae; E. portion of stem with gonotheea; F.

detail of spinouse texture of gonotheca (Figures A, B, E. after Fraser, 1937; C, D, F. after Torrey,

1905). I

II

Gonosome: Gonophores large; shape spherical to elongate-ovoid structures; walls covered by short, •

upcurved spines.

Nematocysts: Not described. H

lira

Type Locality and Type Specimens. Types presumed to be in the collections of the University of

California, not searched. USA, California, San Pedro. me

Distribution. Eastern North Pacific Ocean, British Columbia, Canada to Costa Rica. Depth range, D

16-96 m.

Biology. Colonies typically form tufts on algae and other living substrates. Naumov 1960 [in 1969] i

considered the species to have a type HI life cycle in which a free medusa stage is suppressed and planulae !1

are liberated from the gonangium. This type of reproductive behavior limits dispersal of larvae to the

immediate vicinity of the parent colony. II

Remarks [] . Fraser's illusla'ations (1937) of this species show the gonophores as more elongate than

typically seen in specimens from southern California.

!

i

42 !
!

I SymplecWscyphus tricuspidata (Alder, 1856)

I Figure 1.23

Sertularia tricuspidata Alder, 1856: 356, pl. 13(figs. I-2).

I Comlina tricuspidata: Agassiz, 1865: 146. Sertularella tricuspidata: Clarke, 1876a: 224, pl. 6(figs. 26-27).--Nutting, 1899: 741; 1901: 201; 1904:

100-102, pl. 25(figs. 3-7).--Calkins, 1899: 360, pl. 4(figs. 2A-C).--Torrey, 1904b: 28.--Fraser,

I 1911: 1948: 71; 246.1913: 154; 1914: 193; 1933: 260; 1935: 145; 1936: 126' 1937: 159, pl. 36 (figs. 191a-c);

Symplectoscyphus tricuspidatus (Alder).---Stechow, 1923: 173.---Vervoort, 1972: 166-168, text figs.

i 54a-b.---Cairns etal., 1991: 27.

Sertularella hesperia Torrey, 1902: 63-64, pl. 7(figs. 57-58).

I CType alifornia, Localitynot and Type Specimens: Types presumed to be in collections of the University of searched. CaLifornia, mouth of San Diego Harbor, 1-9 fm [2-16 m].

For additional synonymy information see Nutting 1904.

I Material Examined. California: 1 colony; OCSD 8721, Sta. C (rep. 1), 59 m; 25 April 1987.

Description. Trophosome: Colony erect, small; arising from creeping stolon; branched in alternate

i onodes r dichmarked otomousbypattern pronoun . Stemced slender, constriction twistedofastem tintervals; but without internodes strongregularly septum;preshydrothecae ent above eac at hdistal hydrends otheca; of

internodes set on slight shoulder. Base and about 20-30% of adcauline side of hydrothecae adnate with

'/ stem. Hydrothecae large, maximum length 0.4 mm, maximum diameter 0.2 mm; cylindrical shape, tubular

II with straight or curved sides, often slightly expanded; walls smooth, without annulations; margin with 3

teeth; operculum with 3 valves. Hydrothecae sometimes prolonged with reduplications of margin.

I Gonosome: Gonophores large, height to 1.6 ram; numerous on stem and branches. Gonotheea shape ovoid; distinct raised ridges or ribs present as continuous rings around gonotheca; with small tubular or

funnel-shaped aperture.

I Nematocysts: Unknown.

Type Locality and Type Specimens. Reported to be in England at MNHNT (Nutting, 1904); not

searched. Not traced.

Distribution.: Eastern NorthPacific Ocean, Alaska to Todos Santos Island, Baja California, Mexico.

Depth range, 1-500 m.

I Britain, Circumpolar, Denmark, boreo-arctic species. Also reported from the Eastern North Atlantic Ocean, Great Iceland, Greenland. Western North Atlantic Ocean, New England to Canada

(Newfoundland).

I Biology. Information on the biology of this species is not reported in the literature.

!

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I

I 43
I

i

I

I

i

i

I

m

Figure 1.23. Symplectoscyphus tricuspidata (Alder, 1856): A. entire colony,natural size; B-C. portion of stem i

showingbranching and hydrothecae;C. branch withattached gonotheca; D. gontheca(Figures A,B, i

D. after Fraser, 1937;C. after Naumov, 1960 [in 1969]).

!

Symplectoscyphus turgidus (Trask, 1857)

Figure 1.24 i

Sertularia turgida Trask, 1857:113, pl. 4(fig. 1). i

Sertularella turgida: Clark, 1876a: pl. 38(figs. 4-5).--Torrey, 1902a: 64, pl. 7(figs. 59-62), pl. 8(figs. 63- m

69); 1904b: 29, text figs. 22-23.--Nutting 1904: 85.--Fraser 1911: 71; 1913: 154; 1914: 193; 1935:

145; 1936: 126; 1937: 160; 1940: 41; 1948: 246.--McCormick, 1965: 141,---Lees, 1986: 106. i

gl

SympIectoscyphus turgidus (Trask).---Cairns et al., 1991: 27.

Sertularella conica non Allman, 1877.--Calkins, 1899: 359, pl. 4(fig. 22). i

SertulareIla nodulosa Calkins, 1899: 360, pl. 5(figs. 29A-B). R

Material Examined. California: off Pt. Conception, BRA-02 (USNM). II

Description. Trophosome: Colony erect; small, up to 2 cm; irregularly branched, branches nearly I

same size as main stem. Hydrothecae arranged in alternate pattern on stem; shape tubular, with slraight

sides; margin tricuspidate; base and 25-50% of adcauline side of hydrotheca adnate with stem.

Gonosome: Gonophores I in axils of hydrothecae. Gonotheca spindle-shaped; distal end with few,

low-relief spines.

Nematocysts: Not described. 1
!

I Type Locality and Type Specimens. Presumed to be in the collections of CAS; probably not

extant; not searched. San Francisco Bay, California.

I Baja California, Distribution. Mexico.EasDepth tern North Pacific Ocean, Alaska to San Benito Islands and Santa Maria Bay, range, low intertidal, 0-200 m.

Biology. Typically attached on rocky bottoms. McCormick (1965) recorded several epibionts on

t Symplectoscyphus turgidus, namely a foraminiferan (Cibicides lobatulus), demosponge, hydroid (Lafoea

adnata) and mollusk (Skeneopsis sp.).

!

!

!

I

I

|

1

I

i Figure 1.24. show Symplingectoshydrotheca cyphus turgi e anddusgonot (Trhaecask,e;1857): C-D. dAe.taporti ils ofongonoth of coleoncay,sculptur naturales(Figu ize; reB.s poA-Brti

.onaftoe

fr bFranc raser,h

1937; C-D. after Torrey, 190).

i

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I 4_
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Brown: Boston, MA. 380 pp. i

Agassiz, A. 1865. North American Acalephae. Illustrated Catalogue of the Museum of Comparative l

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m_

Broch, H. 1910. Die Hydroiden der Arktischen Meere. Fauna Arctica 5(1): 129-247.

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I Campbell, R.D. 1969. A statocyst lacking cilia in the hydroid polyp Corymorphapalma. American Zoologist

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I Kramp, P.L. 1948. Trachymedusae and Narcomedusae from the 'Michael Sars' North Atlantic Deep-Sea

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I U.K. 40: 1-469.

Kudelin, N. 1914. Zur Systematik der Sermlariidae, Gattung Sertularella Gray, 1848. Ezhegodnik

I Zoologicheskago Muzeya Imperatorskoi Akademii Nauk 19:108-113.

Lees, D.C. 1986. Marine hydroid assemblages in soft-bottom habitats on the Hueneme Shelf off southern

California, and factors influencing hydroid distribution. Bulletin of the Southern California Academy

I of Sciences 85: 102-119.

Lele, S.H. and Gae, P.B. 1935. Some common Hydromedusae of the Bombay harbour. Journal of the

I University of Bombay 3: 90-101.

Levinsen, G.M.R. 1893. Meduser, Ctenophorer og Hydroider fra Gronlands Vestkyst. Videnskabelige

Meddelelser fra Dansk Naturhistorisk Forening i KCbenhavn (ser. 5), 4: 143-212.

I Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum

characteribus, differentiis, synonymis, locis. Editio decima, reformata. Laurentii Salvii: Holmiae.

I 823 pp.

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cum characteribus, differentiis, synonymis, tocis. Tomus I. Pars II. Edition duodecima, reformata.

I Laurentii Salvii: Holmiae. 533-1317.

PP.

Ljubenkov, J. 1980. Cnidaria. In: Sraughn, D. and R.W. Klink (eds.). A taxonomic listing of common

,I marine 3, Los Angeles, invertebrateCalifornia. species from Pp. 44-68. southern California. Allan Hancock Foundation Technical Report

Maas, O. 1905 Die Craspedoten Medusen der Siboga Expedition. Siboga Expedition, Monograph 10: 1-85.

I Mayer, A.G. 1910. Medusae of the World. The hydromedusae. Vols. I and II. Carnegie Institution:

Washington, DC. 498 pp.

i McCormick, J.M. 1965. Some aspects of the ecology of hydroids off the Oregon coast. Northwest Science 39(4): 139-147.

I MacGinitie, G.E. 1938. Notes on the natural history of some marine animals. American Midland Naturalist I9: 207-219.

MacGinitie, G.E. and N. MacGinitie. 1968. Natural History of Marine Animals. (2nd Edition). McGrawI Hill Book Co.: New York. 523 pp.

Mereschowsky, M.C. 1877. On a new genus of hydroids from the White Sea, with a short description of

I other new hydroids. Annals and Magazine of Natural History (ser. 4), 20: 220-229. Naumov, D.V. 1957. Structure and taxonomic position of Monobrachium parasitum Mereschk.). Doklady

Akademiya Nauk SSSR 113:1168-1170. [In Russian]

I

i 49
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Ii

Naumov, D.V. 1969. Hydroids and hydromedusae of the USSR. U.S. Department of Commerce: Springfield, BI

MD. 660 pp. [Translation of 1960. Gidroidy i gidromeduzy morskikh, solonovatovdnykh i

presnovodnykh basseinov S.S.S.R. [In Russian]. Akademiya Nauk SSSR, Opredeliteli po Faune _

SSSR 70: 1-626.]. II

Norenburg, J.L. and M.P. Morse. 1983. Systematic implications of Euphysa ruthae n. sp. (Athecate: m

Corymorphidae), a psammophilic solitary hydroid with unusual morphogenesis. Transactions of |

the American Microscopical Society 102(1): 1-17.

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Nutting, C.C. 1900. American hydroids. Part I. The Plumularidae. Smithsonian Institution, United States t

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Nutting, C.C. 1904. American hydroids. Part II. The Sertularidae. Smithsonian Institution, United States

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III

Nutting, C.C. 1927. Report on the Hydroida collected by the United States Fisheries Steamer "Albatross"

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m

Rees, W.J. 1967. A brief survey of the symbiotic associations of Cnidaria with Mollusca. Proceedings of the i

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1111

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498. m

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Publications of the Seto Marine Biological Laboratory 7:111-163. I

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m 2. CLASSANTHOZOA:SUBCLASSOCTOCORALLIA

II ORDERS STOLONIFERA AND PENNATULACEA

l by

i F.G. Hochberg 1and John Ljubenkov 2

I Introduction

I Octocorals are colonial anthozoans in which, as is typical of the class, the medusoid stage is

completely absent. Unlike hydrozoan polyps the mouth leads into a tubular pharynx that extends more than

half way into the gastrovascular cavity. The gastrovascular cavity in turn is divided by longitudinal septa

I into radiating compartments and the edges of septa nematocysts. The gonads are gastrodermal

the bear in

origin and the mesoglea is cellular.

l The Subclass Octocorallia, also know as the Alcyonaria, is comprised of forms whose polyps have very uniform construction. Although sizes vary, the basic polyp plan consists of a cylindrical body and a

single ring of tentacles that surrounds to flattened oral disk. Unlike other anthozoans, the symmetry of this

i group is invariably octomerous. All members have 8 complete or perfect septa and 8 tentacles. Each tentacle bears a series of fingerlike processes, termed pinnules, in a featherlike arrangement along both sides (Figure

2.1). There is a single siphonoglyph and muscle bands are always on

i the sucal side of the septa. Almost all species have an internal skeleton formed by separate of fused calcareous sclerites in the mesogloea. t

In colonial forms, such as sea pens and sea fans, a horny central axis ,_

i may be pThe resent. flagella in the siphonoglyph move water into the polyp. :j

Strong flagella on the asulcal septa, the two septa opposite the

siphonoglyph, force water out of the pharynx. The elongate mouth, flattened pharynx and single siphonoglyph confer a bilateral

symmetry on otherwise radially symmetrical polyps. The

i siphonoglyp Polyps h surfproject ace oftefrom n is termed stolons,the vfleshy entral surf matsace.or vertical

extensions of coenenchyme. Polymorphism in polyps typically occurs

i inandsofunction me ordersmainly . Gastrozoo in feeding. ids or aut Siphonozooids, ozooids are large although bear tetinyntacand les

lacking tentacles, have powerful siphonoglyphs that help to ventilate

I large mesozoids, colonies. may Aalsothirdbe present. group of intermediate sized zooids, the

Figure 2.1 Generalizedillustrationof

anoctocoral.

.l

1Deparmaent of Invertebrate Zoology, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road,

Santa Barbara, CA 93105-2936 USA

I 2MEC Analytical Corp., 2433 Impala Drive, Carlsbad, CA 92009 USA

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II

The common names given to the different octocoral groups are based on the different growth patterns U

observed. In sea pens the shape is similar to a fern frond when alive but resembles a narrow pen when

contracted or preserved. In sea pansies the colony is leaf-shaped with polyps on the upper side of the rachis, i

In both groups the colony is anchored in soft sediment by a stalk or peduncle. The related sea fans grow in II

a lattice-like or branched pattern.

The systematics of west coast octoeorals is in need of critical review. The confusion is due in part to I

the cosmopolitan distribution of a number of species which when_first collected off California were described l

as new species. A second problem relates to the lack of types for may of the taxa described from the west

coast. The excellent reviews by Williams (1965) and Bayer (1956,1981) have helped to resolve much of the i

generic confusion surrounding many species names that potentially represent synonyms that still need to be Q

resolved (see also Dunn, 1982).

I

Collection and Preservation

The majority of the telestaceans and pennatulaceans examined in this study were collected by bottom I

trawl. Although small species or specimens can be collected intact and in good condition, most large species/

specimens are broken or otherwise damaged when the come up in large trawl hauls. Gorgonians which are I

attached to rocks or other hard substrates typically are collected by dredge or grab. Small specimens come

U

up in good shape but again large specimens are often damaged in the process of collecting. For species that

live in depths accessible to divers it is better to collect animals by hand to prevent damage. Sea pens should i

be individually dug out of soft sediments taking care to remove the entire animal including the peduncle. It

Gorgonians can be removed from the rocks to which they attach by using a know or chisel.

Prior to fixation and preservation specimens should be narcotized in order to allow the zooids to I

expand. A variety of techniques can be used to relax octocorals. Each species has to be individually tested

to determine what method works best. Chilled sea water works well with shallow water forms, and allowing

deep water specimens to warm up to room temperature is also effective. Chemical agents such as menthol, •

magnesium chloride, magnesium sulfate and MS 222 or other standard narcotizing agents also can be used. I

Specimens to be stored in wet collections should be fixed in 5-10% buffered formalin for no more a

than 24 hours. Large trays or pans should be used for fixing specimens in order to keep them as straight as I

possible. Specimens should not be fixed in storage jars unless they are very small. Following fixation

specimens should be rinsed in freshwater and then placed in 70% ethyl alcohol for long term preservation. do

Large specimens of sea pens and sea fans are often so long or bulky that they are difficult to house, i

Several methods are used to store long pens. 1) In older collections sea pens are often stored in long glass

tubes with cork or rubber stoppers at each end. Although the specimens are preserved straight with distortion ill

they are difficult to study because the entire specimen (often several meters in length) has to be removed in |

order for any portion of the colony to be examined. In addition, long, thin glass containers are difficult to

house unless special racks are built to prevent breakage. 2) Prior to placing in regular museum storage jars, all

long pens can be coiled and tied in several places with thin cordage while still soft and flexible. However II

with this method they are difficult to remove and handle for study. Following preservation the axis often

becomes brittle and when such specimens are untied and uncoiled, they often break. 3) Specimens may be

broken in several places prior to fLxation and preservation in standard museum jars. The disadvantage with i

this method is that such specimens many be difficult to reassemble unless each piece is tagged to indicate its

position in the colony. In this method it also is advisable to tie all the pieces of each specimen together i

especially if several specimens are placed in the same jar. |

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I Gorgonians, especially large specimens, typically are sun dried following collection or fixation.

Small specimens of gorgonians or sections of large specimens in which the polyps have been relaxed should

i bedrawers. preserved Tags inforalcoh driedol.specimens Dried goshould rgoniansbe tied are pla to acedbranch in boroxesto theor main plasticstembagsandtonotprovi leftdloo e prseotecti in theonboxin

or bag.

I Whenever possible color photographs of live animals should be taken to document the form and color of living colonies.

I

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Laboratory Study

I The study of all octocorals requires an analysis of the calcareous sclerites that are present in different

part of the colony. Temporary mounts are made by first removing small pieces of tissue from various parts

of the body and placing each piece on a separate glass microslide (1 x 4 inch). Be sure to record from what

i part of the colony each tissue sample has been taken. The tissue on each microslide is then dissolved in several drops of sodium hypochlorite (liquid bleach). When the tissue is gone add several drops of water to

rinse the preparation. With a free pipette remove the excess liquid and add a small cover slip (22 x 22 ram)

1 prior to examination under a compound microscope. Measurement and drawings should be made to determine [ -- the ranges in sizes and variations in shapes and textures of sclerites from different sections of the body. In

older specimens that have been stored in unbuffered formalin or acidic alcohol, the sclerites may have

I dissolved and be completely absent. However, several sea pen taxa lack sclerites or have very small ones that easily are overlooked. It is best to examine fresh or recently collected specimens for sclerites.

In the case of pennatulaceans, measurements should be made of the colony's total length and then

I the relative lengths of the rachis and peduncle. The total number of pinna (polyp bearing leaves per colony) and the number of gasterozoids per pinna also are important characters. The number and location of

siphonozoids associated with the rachis and pinna are other key characters to observe. Details of external

I morphology, such as the number and location of siphonozooids, especially small ones embedded in the rachis, often can be enhanced by applying a small amount of aqueous methylene blue to the region being

examined.

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Glossary I

Anthoeodia. Upper tentacular part of polyp which Mesentary. Soft septum_

in most cases can be retracted within the II

rind or calyx, mesogloea. Jelly-like substance separating the 2

cellular epithelial layers of cnidada. II

Anthostele.

Lower, thickened part of polyp body 1

wall into which the anthocodia may be Operculum. Anthocodial spicular apparatus that

more or less closes calyx or protects withdrawn; commonly stiffened with

spicules (see also calyx), tentacles in contraction. I

Oral disc. Distal integument of polyp surrounding Asuleal. Side of polyp opposite siphonoglyph.

mouth, enclosed by ring of tentacles; forms Ill

Autozooid. Feeding polyp with 8 well-developed roof over gastrovascular cavity. I_

tentacles and septa; only kind of polyp in

I

Peduncle. Sterile stalk found in sea pens, typically monomorphic species and major type in

dimorphic species [= gastrozooid; see buried in soft sediments, i

siphonozooid]. Pharynx. Tubular passageway between mouth

Axis. Central supporting structure of Gorgonacea and gastrovascular cavity. I

and Pennatulacea; in the former, it may be Physa. Inflated tip of peduncle in sea pens. !1

spicular, consolidated or unconsolidated, or

horny, with more or less nonspicular Pinnule. Digitate lateral branch of tentacle of !11

calcareous matter, octocoral polyps. |

Axoblast. Individual scleroblast of the axis Polyp. Individual of octocoral colony (= zooid).

epithelium. Polyp leaf. United proximal part of adjacent i

Bark. Rind of holaxonian Gorgonacea. polyps thatproduces a leaf-like expansion,

from margin of which anthocodiae project.

Calyx. Wart-like projecting anthostele.

Polypary. Colony as a whole, i

Coenenchyme. Colonial spiculiferous mesogloea.

Rhachis. The polyp bearing portion of a /

Collaret. Transverse, subtentacular ring of pennatulacean colony, often appears 1

spicules, featherlike (see peduncle).

i

Cortex. Outer coenenchymal layer of Rind. Outer, spiculiferous coenenchyme of 1

gorgonaceans, especially Scleraxonia; outer, Holaxonia gorgonians. U

horny layer of holoaxonian axis, as opposed

to its medulla. Sderite. Calcareous skeletal element of It

mesogloea, irrespective of form (= spicule). I_

Crown and points. Transverse collaret with ul

superposed opercular rays. Seleroblasts. Ectodermal cells of mesogloea that

produce calcareous sclerites. all

Filament. Thickened, convoluted edge of septum; I

in Octocorallia, filaments of 2 septa Septum. Thin radial noncalcareous partition

opposite siphonogIyph are very long and dividing gastrovascular cavity of polyp (see

heavily flagellated, whereas those of mesentery), i

remaining six are shorter and glandular. Siphonoglyph. Strongly ciliated groove extending

Gastrovascular cavity. Interior space of polyp down one side of pharynx, i

radially partitioned by septa. Siphonozooid. Polyp with reduced tentacles or

It

Medulla. Central zone of scleraxonian stem; none, and commonly reduced septal l

rarely, central chord of holaxonian axis. filame autozooids. nts; usually much smaller than i

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I Solenium. Canal lined with gastrodermis Sulcus. See siphonoglyph.

penetrating coenenchyme and

I interconnecting gastric cavities of polyps. Verruea. Calyx; tubercle of sclerite.

Spicule. Properly, a long sharp sclerite (see Zoaid. Any individual of colony, irrespective of

sclerite), its morphological specializations (= polyp;

in most German works; = siphonozooid).

I Sulcal. Side of polyp nearest siphonoglyph.

I List of Species

i Subclass Octocorallia Haeckel, 1866

Order Stotonifera Hickson, 1883

i Family Telestidae Milne-Edwards and Haime, 1857

Telesto californica Kfikenthal, 1913

I Telesto nuttingi Kiikenthal, 1913 Telestula ambi gua (Nutting, 1909)

i Order Pennatulacea Verrill, 1865

I Suborder Sessiliflorae K0&enthal, 1915

Family Stachyptilidae K611iker, 1880

i Stachyptilum superbum Studer, 1894 Family Ombellulidae Williams, 1995

Ombellula magniflora (K611iker, 1880)

I Suborder Subselliflorae Kiikenthal, 1915

Family Halipteridae Williams, 1995

i Halipteris californica (Moroff, 1902)

Family Pennatulidae Ehrenberg, 1834

I Pen Ptilonamla sarcus califomica gurneyi (Gray, Kiike1860) nthal, 1913

Family Virgulariidae Verrill, 1868

I Virgularia agassizii Studer, 1894

Acanthoptilum album Nutting, 1909

t Acanthoptilum gracile (Gabb, 1863) Stylatula eIongata (Gabb, 1862)

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II

Description of Species II

Subclass Octocorallia Haeckel, 1866

Order Stolonifera Hickson, 1883 gl

Family Telestidae m

Telesto californica Kiikenthal, 1913

Figure 2.2 I

Telesto rigida Nutting, 1909: 685. Not Wright and Studer, 1889. i

II Telesto californica Kfikenthal, 1913:229-231, text figs A, B, pl.7 (figs. 1, 2).--Ljubenkov, 1980:56

Material Examined. BLM-OCS collections, m

I1

Description. Colonial octocoral composed of an axial polyp with lateral, branching polyps whose

size is equal to the axial polyp. Base of colony lamellar. The primary (axial) polyp may give rise to secondary

andtertiary polyps. The polyps have eight longitudinal furrows which indicate mesenterial insertions; proximal l[

part of polyp empty. The stiff walls of the polyps contain mainly warty spindles for spicules. II

I

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A B I

Figure 2.2 Telesto SBMNHcalifornica 51530. , views of colony: A, SBMMH 01-65-32; B, USC t948-50; C, detail of polyp I

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I Type Locality and Type Specimens. Holotype, not designated in original description. California,

bearing 8.6° W 2.5 mi off east point San Nicolas Is1and, 31 fm [57 m]; coll. USB CF Albatross, station 4422.

I Distribution. Southern California and probably northern California.

Biology. While noted from 31 fins by Nutting, this is primarily a species from outer shelf andupper

slope areas. BLM-OCS benthic photos often showed this species on the sill between basins.

l Remarks. Nutting thought this was Wright and Studer's T. rigida, but Kiikenthal recognized it as a

new species.

I Telesto nuttingi Kiikenthal, 1913

i Figure 2.3

Telesto nuttingi Kiikenthal, 1913: 231-233, text figs C, D, pl.7 (fig. 3).

I Material Examined. BLM-OCS collections; SBMNH

Description. Colony composed of an axial polyp with lateral polyps whose size is equal to or less

i than of polyp the axial empty. polyp. TheLongitudinal axial polyp also furrows has deep and narrow. Length of longest polyps 75 mm; proximal part short lateral polyps except where the larger branches originate.

Anthocodial spicules are flattened spindles (0.18 x 0.06 mm) thickly studded with warts. Color orange gold

i to brightType gold. Locality and Type Specimens. Holotype not designated; possibly still in the collections at

SIO. Southern California, China Point, 48 fm [88 m].

I Etymology. Named in honor of Nutting for his work on octocorals. Distribution. Southern California.

i Bio Remlogy. arks.Unknown. Kiikenthal found this species in the collections at the "Biological Station in La Jolla"

while reviewing Nutting's material.

I Genus Telestula Madsen 1944

i while zooids Diagnos areisy. ouOct ng;ocorals lower wit parth smal of coel l slende entericr zooids cavity ininwhich older mese zooidsnteripaertiall s extend y filleddownwithto mesogloeal stolon only

tissue. Proximal part of zooid developed into stem-like structure which may constitute largest part of zooid;

i when secondary zooids may arise from this area. Proximal part of secondary zooids developed in similar manner colony reaches certain size. Secondary zooids of higher order may occur. Sclerites of body wall

typically blunt, coarsely tuberculate oval plates; those of intrusion tissue much-branched irregular forms

t more or Tleype ss fused speciestogether. . Telestula septentrionalis Madsen; by original designation.

Remarks. Colonies like Telesto but smaller and less profusely branched.

I

I

I 59
IIiII

,iIIII! B I

Figure2.3 Telesto nuttingi, views of colony: A, SBMNH;B, SBMNH 1348-4.Iii!I!

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I Telestula ambigua (Nutting, 1909)

Figure 2.4

I Telesto ambigua Nutting, 1909: 686, pl. 84(fig. 1, 2), pl. 90(fig. 1).

Telestula ambigua: Bayer, 1952:131.

I Material Examined. Holotype (see below).

Description. Colony with simple monopodial branching; base membranous. No regularity to

I secondary tertiary branching. Height to ram.

and 14

Spicules in polyp wails not fused; slender, thorny spindles in calyx walls.

I animal appears Color inbanded. alcohol pale yellowish-brown, lighter distally; longitudinal furrows lighter than background,

Type Locality and Type Specimens. Holotype - USNM 25421. Monterey Bay, California, bearing

I S 39°E 10.7 mi off Pt. Pinos lighthouse, 524 fm [958 m]; coll. USBCFAlbatross, station 4514.

Distribution. California, currently known only from the type locality.

i Bio Remlogy. arks.Unkno Lackwn. of fused spicules and growth mode separate this genus from Telesto.

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B

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Figure 2.4. Telestoambigua: A, views of 3 polypsfrom colony, Holotype, USNM 25421; B, scleritesfromthe

I body.

i

I 61
I

i

n

Order Pennatulacea Verrill, 1865 Ill

Key to Families of Pennatulaceans m

(Modified from Hickson, 1916) l

1A. Rachis radially symmetrical; axis present, rudimentary or absent ................................ Veretillidae n

lB. Rachis bilaterally symetrical ........................................................................................................... 2

2A. Axis absent ...................................................................................................................................... 3 I

2B. Axis present ..................................................................................................................................... 4

N

3A. Rachis flattened, leaf-shaped ............................................................................................. Renillidae

3B. Rachis cylindrical or club-shaped .............................................................................. Echinoptitidae i

4A. Autozooids irregularly distributed or arranged in longitudinal rows on the rachis ........................ 5 I

4B. Autozooids arranged in whorls or bilateral pairs ............................................................................ 7 II

4C. Autozooids arranged in transverse or oblique rows or in primitive leaves ..................................... 8

4D. Autozooids arranged in well developed leaves supported by spicules ......................................... 10 I

5A. Autozooids without calices .................................................................................. Kophobelemnidae I

5B. Autozooids with calices .................................................................................................................. 6 l

6A. Autozooids of various stages of growth arranged in closely set oblique rows on the rachis ............ n

..................................................................................................................................... Funiculinidae

6B. Autozooids arranged in longitudinal rows on the rachis ............................................... Protoptilidae Nu

7A. Arranged in single terminal whorl or tassle ............................................................. Ombelullulidae

7B. Arranged in several whorls or pairs ............................................................................... Chunellidae il

8A. With ventral radial canals; gonads developed in fully formed antozooids ................... Halipteridae 1

8B. With dorsal radial canals ................................................................................................................. 9 U

9A. Gonads developed in fully formed autozooids ............................................................ Anthoptilidae 1

9B. Gonads developed in young autozooids ....................................................................... Virgulariidae

10A. Siphonozooids on leaves absent .................................................................................... Pennatulidae

N

10B. Siphonozooids on leaves present ................................................................................... Pteroeididae Ill

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Figure 2.5. Genera of sessiliflorae pennatulaceans, entire colonies: A. Renilla amethystina: 10-25 m, Redondo

Canyon, CA (SBMNH 51257), 60 nun RL;B. KophobeIemnon affine: 2430-2710 m, Monterey Bay,

I CA (SBMNH), 100 ram TL;C. Ombellula lindhali: 1005-1060 m, off Avila, SantaLucia Banks, CA (SBMNH 49154), 350 mm TL; D. Anthoptilum grandiflorum: 457-512 m, off Avila, Santa Lucia

Banks, CA (SBMNH 49152), 380 mm TL; E. Stachyptilum superbum: 181 m, off Avila, Santa Lucia

i BLuanks, CA (SBMN-I-/49156), 100 mm TL; F. Distichoptilum gracile: 1260-1277 m, off Avila, Santa cia Banks, CA (SBMNH 49155), 1100 mm TL.
!

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A D

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Figure 2.6. Genera of subselliflorae pennatulaceans, entire colonies: A, Halipteris californica: 475 m, Santa

Barbara Channel, CA (SBMNH 45568), 950 mm TL; B, Stylatula elongata: (modified from

MacGinitie & MacGinitie, 1968), 280 mm TL; C, Virgularia agassizi: 14 m, off Santa Barbara, CA 1

(SBMNH 49487), 300 mm TL; D, Acanthoptilum gracile: 15-35 m, off Santa Catalina Island, CA II

(SBMNH 49691), 650 mm TL; E, Ptilosarcus gurneyi: 15-35 m, off Santa Catalina Island, CA

(SBMNH 49688), 225 mm TL; F, Pennamla californica: 475 m, Santa Barbara Channel, CA im

(SBMNH 45509), 90 mm TL. I
!

I Suborder Sessiliflorae Kiikenthal, 1915

i Family Stachypfilidae K611iker, 1880

! Genus Stachyptilum K6Uiker, 1880

I Diagnosis. Colonies stout and clavate or slender, firm or spongy in texture. Symmetry of rachis bilateral throughout. Axis present throughout length of colony. Polyp leaves absent. Autozooids arranged in

oblique rows in 2 longitudinal series along rachis. Each row disposed alternately to corresponding row on

I other side of rachis. Anthocodiae 3 to several per row, retractile into densely-spiculated calyces. Calyces indistinctly toothed or with 2-3 long terminal teeth. Siphonozooids mostly in oblique rows between rows of

autozooids, often with calyces formed by fan-shaped arrays of sclerites. Sclerites of polyp calyces and

I rahave chisrods with(modified 3-flangedfromneedles Williams, and spindles; 1995) peduncle may have oval- or rod-shaped plates; tentacles may

Type Species. Stachyptilum macleari K611iker, 1880; by monotypy.

I

Stachyptilum superbum Studer, 1894

I Figure 2.7

Stachyptilum superbum Studer 1894: 56.--Nutting, 1909: 708-709; pl. 37(figs. 5, 6).-- Balss, 1910: 36.--

l Kiikenthal and Brock, 1911: 261-265; text-figs. 84-91.--Kiikenthal, 1913: 259-261; text-figs. C', D1,pl. 8(fig. 9).

I 0-10 em.,(2). Material Examined. California, Santa Mafia Basin, off Pt. Sal, CAMP 1-1, Sta. R7, rep. 3, 0.5,

Description. A short, stubby colony with completely adnate leaves that appear to fall into spirals.

I 4-7 polyps per "whorl". Polyps fully retractile into a heavily spicularized calyx, with at least 2-6 larger spicules protruding at top. On one side calyces absent from strip containing 2 rows of siphonozooids. Patches

of siphonozooids also present below each calyx. Axis not running into physa. Balloonlike physa perforated

I with "cinclides". An empty chamber parallel to axis.

Type Locality and Type Specimens. See Studer, 1894. Panama, Gulf of Panama, 7°16'45"N,

79056'30"W, 210 fm [384 m]; coll. USBCF Albatross, station 3389.

I Distribution. Central and southern California.

Biology. Occurs on Slopes.

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Diagnosis, Colonies with long, slenderstalk and terminal duster of polyps, Symrnetry of rachis

bilateral. Conspicuous axis present throughout colony; round to quadrangular in cross-section. Polyp leaves •

absent. Autozooids restricted to terminal cluster containing 1-40 polyps. Anthocodiae non-retractile, calyces I

absent. Siphonozooids present on rachis at base of autozooids and below terminal cluster on upper part of

stem. Sclerites present (in only 3 species) or totally absent (in all other species). When present sclerites I

spindles, rods, ovoid rods, or needles; three-flanged or round in cross-section; surfaces often rough with I

numerous low knobby tubercles. Sclerites often present in tentacles, walls of autozooids and peduncle

(modified from Williams, 1995). I

Type Spedes. Isis encrinus Linnaeus, 1758; by subsequent designation. I

Ombellula magniflora Kiilliker 1880 i

Figure 2.8

IlI

Umbellula magniflora K611iker,1880: 240.--Nutting, 1912:420.---Pastemak, 1962: 126.--Ljubenkov, 1980:58. I

UmbeUula loma Nutting, 1909: 712-713, pl. 87 (fig. 9). I

Type Locality and Type Specimens. Types presumed to be at University of California, Scripps Institution I

of Oceanography; not extant. USA, California, off San Clemente Island, 330 fm [604 m].

!
!

I Material Examined. California: BLM-OCS,

SCCWRP collections. Pt. Piedras Blancas, 35°29.3'N,

i Ap121ri°l35.8%V, 1974; SIO906B174-12; m; coll. RNNofAgass Catalina iz, 25Island, ft otter33°traw33.5q_1, l, 1

118°39_, 534-585 m; SIO 76-95, 40 ft otter trawl, 9 March

I 33_33.5'N, i976; SIO118CO-422; °38%V,556-585 N end m; of coll. SantaSIOCatalina 76-96, 40Island, ft otter

trawl, 9 March 1976; SIO CO-426.

I rach Description. Distinctive long stem with tuff (or is) of rather large polyps atdistal end. Rosette shaped

l mchis with only one ring of secondary autozooids atouter

I margin. Up to twelve zooids; axis square. Sclerites absent throughout colony.

Type Locality and Type Specimens. West of

i Kerquelen Island, 1600m. Types not extant. Distribution. Southern California.

I "" Biology. Occurs in deep basins usually below about

Remarks. Many more species of Umbellula have

: : ._.,:_ been described since early specimens were taken from

_i many places. However, Pastemak (1962) collapsed all

forms which locked sclerites into 3 species, namely: U.

I . magniflora, U. lindahli, and U. thomsoni.

oO

Figure 2.8. Ombellula magniflora: view of

I single polyp (SBMNH49154).

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m 67
!

Ii

Suborder SubseUiflorae Kiikenthal, 1915 •

Family Halipteridae Williams, 1995 n

u

Genus Halipteris Krltier, 1869

Diagnosis. Colonies long, slender, whip-like. Rahcis bilaterally symmetrical throughout. Axis extends n

throughout colony length; round to quadrangular in cross section. Polyp leaves present in mature colonies I

as ridges. Autozooids disposed in many oblique rows along 2 longitudinal series. Adjacent autozooids often

united attheir bases forming raised ridges, which emerge from opposite sides ofrachis. Anthocodiae retractile

into calyces with 2 terminal teeth. Siphonozooids sparsely scattered on rachis between oblique rows of m

autozooids. Sclerites: calyces with 3-flanged spindles; peduncle with smooth rods; tentacles often with

three-flanged rods. N

Type Species. Virgularia christii Koren and Danielssen, 1847; by subsequent monotypy (see KrUiker,

1869). m

Halipterus eahfornica (Moroff, 1902)

Figure 2.9 N

Pavonaria californica Moroff, 1902: 393-394.--Kfikenthal, 1913: 249-250; table 8; pl. 8(figs. 6, 7), textfigs. V,W. I

Balticina californica: Nutting, 1912: 40.

Balticina pacifica Nutting, 1909: 704-705; pl. 87(figs. 1,2); 1913; pl. 6(fig. 4). B

Type Locality and Type Specimens: Holotype - USNM 25426. USA, California, bearing S 59_E 5.6 m

mi off Pt. La Jolla, Soledad Hill, 243-280 fm [444-512 In]; coll. USBCF Albatross, station 4326

Halipteris contorta Nutting, 1909: 707-708; pl. 86(figs. 3,4). I

Type Locality and Type Specimens. Holotype - USNM 25427. USA, California, Santa Catalina

Island, bearing SW 2.1 mi off SE Point, 52-88 fm [95-161 m]; coll. USBCFAlbatross, station 4409.

Stachyptilum quadridentatum Nutting, 1909: 709; pl. 87(figs. 7,8). •

Type Locality and Type Specimens. Holotype: USNM 25428. USA, California, bearing N 86o30,E

9.4 mi off Point Loma lighthouse, 92-108 fm [168-198 m]; coll. USBCF Albatross, station 4360. I

Material Examined, SBMNH specimens from California.

Description. Colony large. Axis runs into physa. Polyps very large, non-retractile; arranged in

114+ rows, 2-5 per row; polyps surround and obscure stem. Calyces with thorny spines; decreasing in size m

from fast to last to last polyp in each row. Siphonozooids, 15 to 20 between each row of polyps; not surrounded

by spicules. Spicules confined to polyps and calyces; tentacles spiculate. Spicules bar shaped.

I Type Locality and Type Specimens. Types unknown. Presumed to be from the coast of California,

details of locality not provided in original description.

Distribution. Off California, about 200 m and deeper, i

Remarks. Moroffmay not have macerated the polyps and therefore overlooked their spicules which

led Nutting to place this in Balticina. _

Biology. An Outer Shelf to Basin floor species. Benthic photographs of living Balticina show the II

polyps expanded to great size while the colony leans over so that the distal end nearly touches the substrate.

I

68 !
i

I Figure 2.9. Halipterus californica : A, B, C views of colony; D, sclerites from the body. SBMNH 45568.

I

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I

I 69
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i

Family Pennatulidae Ehrenberg, 1834 •

Genus Pennatula Linnaeus, 1758 I

I

Diagnosis. Colonies feather-shaped in appearance. Symmetry of rachis bilateral throughout. Axis

present throughout length of colony. Polyp leaves present, usually large and conspicuous, deltoid, sickle- It

shaped, or fan-shaped. Autozooids arranged in 1 or more rows along margins of polyp leaves. Anthocodiae i

retractile into spiculiferous calyces. Calyces tubular, most often with 8 terminal teeth. Siphonozooids confmed

to rachis, often extending between polyp leaves. Mesozooids also may be present on rachis or on margins of

polyp leaves opposite the autozooids. Sclerites: calyces with 3-flanged needles; peduncle surface with 3- Ii

flanged rods; peduncle interior with tiny ovals.

u

Type Species. Pennatula phosphorea Linnaeus, 1758; by monotypy. I

m

Pennamla cah'fornica Kiikenthal, 1913

Figure 2.10 i

Pennatula acuIeata Nutting, 1909: 688-689. i

Pennatula phosphorea var. californica Kiikenthal 1913: 241-243.

Material Examined. BLM-OCS collections.

Description. A relatively small pennatulid with 20+ pairs of leaves (pinnae) with up to 10 autozooids II

per leaf; leaves stiff and heavily spicularized. Stem swollen below the rachis. Siphonozooids on venla'al

surface of stem in bands on either side of a bare median strip. Spicules are needles and bars of a bright 1

carmine color which give color to the colony. II

Type Locality and Type Specimens. Holotype not designated, but all specimens apparently come

from Pt. Loma or the Channel Islands. i

II

Distribution. Throughout California possibly the west coast.

Biology. Occurs at the base of slopes and basins generally below 400 m.

Remarks. Nutting (1909) insufficiently described his species Pennatula aculeata. Kiikenthal (1913), II

who re-examined some of Nutting' s specimens, states that a note in the jar in Nutting' s handwriting identifies

10 specimens ofP. phosphorea from Albatross "Station 4407". Nutting does not list this station but he does I!

list an Albatross Station 4417 which is perhaps what Kiikenthal meant.

i

Genus Ptilosarcus Verrill, 1865 i

Diagnosis. Colonies stout; feather-shaped. Rachis bilaterally symmetrical throughout. Axis extends

length of colonly. Polyp leaves present; kidney-shaped, margins often sinuous. Autozooids crowded on i

distal margins of polyp leaves. Anthocodiae retractile into calyces. Calyces spiculiferous, usually with 1-2 II

indistinct to very conspicuous terminal teeth. Siphonozooids numerous; in 2 longitudinal tracts along rachis,

clumped or continuous, not present on polyp leaves. Sclerites - calyces and polyp leaves with three-flanged lil

needles and spindles or longitudinally grooved oval-shaped plates and rods; peduncle with smooth, relatively I

large ovals.

Type Species. Sarcoptilus gurneyi Gray, 1860; by subsequent monotypy. •

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Figure 2.10. PennatMa californica:A, lateral view of pinna; B, scleritesfrom the body. SBMNH 45509.

I

I Ptilosarcus gurneyi (Gray, 1860) Figure 2.11

i Material Examined. BLM-OCS, County, and SBMNH collections.

Orange

Description. Colony shaped like a fleshy feather with a very fat stem; with 52+ leaves per side, and

i about 150 polyps per leaf in 3 rows. Two wide rows of siphonozooids present on ventral side of rachis with bare zone between them. Spicules include spindles and needles colored violet, red, orange, and purple. Stem

square in cross section. Nutting (1909) contains a detailed anatomical description of an individual.

i TDype Locality and Type Specimens. Not searched. Not traced. istribution. Pacific coast from Alaska through Baja California.

Biology. A shelf species which occurs intertidally in colder waters. Often forms dense beds where

I they are preyed on by several sea stars.

Remarks. See Batie, 1972 for the extensive synonyms of this species including Sarcoptilus,

i Leioptilum and their anagrams.

I 71
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Figure 2.11. Ptilosarcus gurneyi: sclerites from the body. SBMNH 49688. I

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I Family Virgulariidae Verrill, 1868

t Genus Virgularia Lamarck, 1816

Diagnosis. Colonies long, slender and vermiform or more stout, robust and rigid. Rachis symmetry

bilateral throughout. Axis extends throughout most of colony length; sometimes extending beyond apex of

I rachis; mostly round in cross section. Polyp leaves present; relatively short, often congested or sometimes

with intervals of bare rachis between adjacent leaves. Autozooids tubular, fused for most of length to form

relatively thin poly leaves; 3-100+ autozooids per polyp leaf. Anthocodiae retractile into bulbous flesh;

I portions polyps, calyces. Siphonozooids sparsely on polyp proximal of forming fleshy distributed leaves

below free parts of autozooids or more commonly on rachis between polyp leaves. Sclerites absent except

for minute oval bodies in peduncle interior.

I Type Species. Pennatula mirabilis Linnaeus, 1758; by subsequent monotypy.

I Virgularia agassizii Studer, 1894

Figure 2.12

I Virgularia bromleyi Ktlliker, 1880: ??--Kiikenthal, 1913: 243.

Halisceptrum cystiferum Nutting, 1912: 698.

I Material Examined. BLM-OCS and OCSD collections.

Description. Typical elongate sea pen with 32 or more pairs of short (in contraction) leaves, 4-5

I ppolyp. olyps Entire per leaf. colony Virguwith larifoonly rm paofewlypssmall with 8spicules bumps orornone small(especially projectionsinrevealing long presthe ervedoctospecimens). -symmetry Physa of the

swollen with axis visible within. Stem with distinct groove.

I Type Locality and Type Specimens. see K611iker,1880. Challenger Reports, Zoology, pt. 2.

Distribution. Throughout California.

i Bio Remar logy.ks.APolyps very common of genusseaVirgularia pen alongarethedistin Pacific ct incoast. their poor spiculation and beautiful feathery

form when expanded. Colonies can retract into the substrate.

I Genus Acanthoptilum Kolliker, 1870

i throughout. DiagAxis nosis.presen Colonies t throulong, ghoutslender, length ofto colony. somewhat Polypfeather-shaped. leaves present,Rachis often bilaterally sickle-shaped, symmetrical biseriate

along rachis; numerous and mostly crowded with extent of naked rachis between leaves usually not exceeding

i length of single leaf. Autozooids 4-9 per polyp leaves, grading larger distally. Anthocodiae retractile into sparsely spiculated calyces which may have several indistinct rounded terminal teeth. Siphonozooids minute,

in 1 or 2 longitudinal rows on rachis between polyp leaves. Sclerites: at base of each polyp leaf an irregular

cluster of 3-flanged spindles; calyces, rachis and peduncle with 3-flanges spindles or small to minute ovals

I or rods (modified from Williams, 1995).

Type Species. Acanthoptilum pourtalesii Kolliker, 1870; by monotypy.

!

m 73
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I

I

Figure 2.12. A, B. Virgularia agassizii: views of portion of racNo, SBMNH 51250. C. Virgularia catifornica:

lateral view of portion of rachio showing single pinna. SBMNH49487. I

I

Acanthoptilum album Nutting, 1909 Ig

Figures 2.13 _l

It

Acanthoptilum album Nutting, 1909: 701-702.--Ktikenthal, 1913: 246-247.--Ljubenkov, 1980:58.

Material Examined. Orange County collections. I

Description. Typical member of genus with characteristic spicularized leaves and calyces. 4-5

polyps per leaf, 3 siphonozooids between adjacent leaves. All spicules are clear which gives the colony a 1

white appearance. The rachis occupies about 2/3 of the total length. I

Type Locality and Type Specimens. Holotype - USNM 25424. California, bearing S 15° E 2.8 mi

off Point Pifios lighthouse, 59-65 fm [108-119 m]; coll. USBCF Albatross station 4473. II

Distribution. Central and southern California. |

Biology. Unknown. m

Remarks. See remarks under Acanthoptilum gracile. 1

74 I
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I Figure 2.13. Acanthoptilum album: A, B, C views of colony; D, sclerites from the body. SBMNH 49268.

i

I 75
!

Acanthoptilum gracile (Gabb, 1863) I

Figure 2.14

Virgularia gracilis Gabb, 1863: 120.--Verrill, 1868: 383-384. i

?Acanthoptilum1980: 58.gracile:Nutting, 1909: 700-701, pl. 86(figs. 3, 4).--Ktikenthal, 1913: 245-246.--Ljubenkov, I

Acanthoptilum gracile: Kukenthal, 1915: 65.--Ricketts, Calvin and Hedgepeth, 1968:317, text-fig. 234.--

Morin, 1976: 630, text-fig, lb.--Williams, 1995: 123, text-figs. 5E, F, 9H. I

Material Examined. BLM-OCS, Orange County collections.

alcohol light Descripti tan obrown. n. LongLeaves slender withwhiplike 8-9 autozooids; colony with calyces basalandphysa leavesandheavily withoutspicularized a purple band. with Color needles;in I

spray at base of leaves with large spicules; 6-12 prominent siphonozooids between adjacent leaves. Needle

spicules with slight yellow tinge, not carmine. I

Type Locality and Type Specimens. Not designated; presumed not extant (see Coan and Bogan,

1988). Monterey Bay, California, 20 fm [36 m].

Distribution. Monterey and southern California. I

Biology. Unknown.

bases, calyRemcesarks. usually Acancontain thoptilumlongspp. needles. have whiplike The leaves colonarey shapes, attachedleaves by their withbase a spratoy the of spstem; iculestheyat their are I

heavily laden with spicules. The stem usually contains spicules lightly scattered and the purple band is thick

with spicules. The genus needs a thorough revision. I

I

I

I

I

I

I

I

I

I
I Figure 2.14. Acanthoptilum gracile: A, B, C, D views of colony; E, scledtes fi'om the body (A-C, SBMNH 49693. D, SBMNH 49691).

I 77
!

n

Genus StyIatula Verrill, 1864 •

Diagnosis. Colonies long, slender, often vermiform. Rachis bilaterally symmetrical throughout.

Axis present through entire colony; mostly round in cross section. Polyp leaves present, short, densely II

arranged on rachis. Each polyp leaf subtended by strong fan-like armature of long sclerites. Autozooids 5- g

30 per polyp leaf. Anthocodiae retractile into bulbous basal portions of polyps. Siphonozooids absent from mR

polyp leaves, arranged on rachis between leaves. Sclerites: polyp leaf armature with elongate, partly or ll

entirely 3-flanged, flattened needles, often broad and rounded at one end, tapering and pointed at opposite

g

end; calyx-like proximal portions of polyps may contain spindles; peduncle interior often with minute oval

bodies; absent from other parts of body. 1

IO

Type Species. Virgularia elongata Gabb, 1862; by subsequent designation.

Stylatula elongata Gabb, 1863 I

Figure 2.15 n

Virgularia eIongata Gabb, 1863: 167.--Gray, 1870: 19.

Stylatula elongata: Verrill, I864: 30;1868: 384; 1870: 560.--Nutting, 1909: 699-700, pl. 86(fig. 2).-- •

Kukenthal and Broth, 1911: 317-319, text-figs. 114-116.--Kiikenthal, 1913: 244-245; 1915: 68- l

69, text-fig. 70.--Morin, 1976--Ljubenkov, 1980: 59.--Williams, 1995: 122-123, figs. 5C, D, 9G.

Stylatula Ringei Pfeiffer, 1886: 59. H

IN Type Locality and Type Specimens. Types not searched. "Unter-Califomien, yon Diego" [California,

San Diego]. i

Stylatula columbiana Verrill, 1922:11-12, pl. 3 (figs. 1-4a). i

Type Locality and Type Specimens. Types not searched. [Canada, British Columbia], W coast of

Vancouver Island, Ucluelet, 13 fm [24 m]. !

i

Material Examined. BLM-OCS and Orange County Collections.

Description. Colony shaped like test-tube brush; 20-24 polyps per leaf; leaves attached by inner •

margin to stem. Each leaf supported by spray of 12-16 large needle spicules, with leaves slightly overlapping Ii

on one side of stem; with about 10 pairs of leaves per inch, with numerous papilliform siphonozooids

scattered about free portions of rachis. No spicules in polyps. •

Color in life: colony grayish-brown; physa salmon. Color in alcohol: colony brownish-white.

Type Locality and Type Specimens. Types not designated; presumed not extant (see Coan and

Bogan, 1988). California [further locality details not provided in orignial description]. 1

am

Distribution. California.

Biology. Numerous benthic photographs show that many colonies have had their tips eaten by I

fishes. These nipped tips provide a substrate for some hydroids and Bryozoa. 1

!

!

!

78 I
I Figure 2.15. StyIatula eIongata: A, B, C, views of colony; D, sclerites from the body (SBMNH 49704).

I 79
!

Acknowledgments I

The illustrations were prepared by Joann T. Rounds (pen and ink) and Jamie Calhoun (graphite). I

Specimens loans were facilitated by Ted Bayer (USNM); Spencer Luke (SIO benthic collection); James I!

Nybakken (MLML); and the late Dusty Chivers (CASIZ). 1

II

Literature Cited

I1

Balss, H. 1910. Japanische Pennatuliden. In: F, Doflein (ed.), Beitr_ige zur Naturgeschichte Ostasiens. 1

Abhandlungen der Mathematisch-Physischen Classe der Krniglich S_ichsischen Gesselschaft der m

Wissenschaften l(suppl. 10): 1-106. !I

Batie, R.E. 1972. Investigations concerning the taxonomic status of the sea pen Ptilosarcus gurneyi (Cnidaria,

Pennatulacea). Northwest Science 46(4): 290-300. n

Bayer, F.M. 1956. Octocorallia. Pp. F166-231. In: R.C. Moore (ed.). Treatise on Invertebrate Paleontology,

Part F. Coelenterata. Geological Society of America and University of Kansas Press: Lawrence KS.

498 pp. I

Bayer, F.M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa),

with diagnoses of new taxa. Proceedings of the Biological Society of Washington 94(3): 902-947. n

Coan, E.V. and A.E. Bogan. 1988. The Recent invertebrate taxa described by William More Gabb 1839-

1878. Proceedings of the Academy of Natural Sciences of Philadelphia 140(1): 273-284. 1

Deichmann, E. 1936a.'Notes on Pennatulacea and Holothurioidea collected by the first and second Bingham I

Oceanographic Expeditions 1925-1926. Bulletin of the Bingham Oceanographic Collection 5(3):

1-11. I

Deichmann, E. 1936b. The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of the Museum

of Comparative Zoology 53: 1-317.

m

Deichmann, E. 1941. Coelenterates collected on the Presidential Cruise of 1938. Smithsonian Miscellaneous I

Collections 99(10): 1-17.

Dunn, D.F. 1982. Cnidaria. Pp. 669-706. In. S.A. Parker (ed.), Synopsis and Classification of Living I

Organisms. Vol. I. McGraw-Hill Book Co.: New York, NY. II

Gabb, W.M. 1863. Description of two new Pennatulidae from the Pacific Coast of the United States. I1

Proceedings of the California Academy of Natural Sciences 2: 166-167. I

Gabb, W.M. 1864. Description of a new species of VirguIaria from the coast of California. Proceedings of llll

the California Academy of Natural Sciences 3: 120. 1

Gray, J.E. 1860. Revision of the family Pennatulidae, with some descriptions of some new species in the

British Museum. Annals and Magazine of Natural History, (ser. 3) 5: 20-25. n

Gray, J.E. 1870. Catalogue of sea-pens or Pennatulariidae (sic) in the collection of the British Museum.

London. 40 p. 1

Hickson, S.V. 1915. Some Alcyonaria and aStylaster from the west coast of North America. Proceedings of U

the Zoological Society, London, 1915: 541-557.

I
!

I Hickson, S.J. 1916. The Pennatulacea of the Siboga Expedition, with a general survey of the order. Siboga

Expedition Monogr. 14(Livre 77): 1-265.

I S.J. 1921. On in the Museum. of the

Hick,son, some Alcyonaria Cambridge Proceedings Cambridge

Philosophical Society 20(3): 366-373.

i Hickson,Society S.J. 1930. of London, Some aleyonarians 1930(1): 209-227. from the Eastern Pacific Ocean. Proceedings of the Zoological

Hickson, S.J. 1937. The Pennatulacea. Scientific Reports, John Murray Expedition, 1933-34, 4(5): 109-130.

I Hyman, L.H. 1940. The Invertebrates: Protozoa through Ctenophora. McGraw-Hill: New York. 726 p.

Krlliker, R.A. von. 1869. Anatomisch-Systematische Beschreibung der Alcyonarien. I. Die Pennatuliden.

I Abhandlungen vonder Senckenbergischen naturforschenden Gesellschaft 7:11

1_ 5 _

K61liker, R.A. yon. 1872a. Anatomisch-Systematische Beschreibung der Alcyonarien. I. Die Pennatuliden.

I Abhandlungen vonder Senckenbergischen naturforschenden Gesellschaft 8: 85-275.

Krlliker, R.A. yon. 1872b. Morphotogie und Entwickelungsgeschichte des Pennatuliden-stammes nebst

allgemeinen Betrachtungen zur Deseendenzlehre. Christian Winter: Frankfurt a. Main.

I Krlliker, R.A. yon. 1875. Die Pennatulide UmbeUuIa und zwei neue Typen der Alcyonarien. In: Festschrift

zu Feier des fiinfundzwanzigjiihrigen Bestehens der Physikalisch-medizinischen Gesellschaft in

I Wiirzburg. 23 pp.

K611iker, R.A. yon. 1880. Report on the Pennatulida dredged by H.M.S. Challenger during the years 1873-

1876. Report on the Scientific Results of the Voyage of the H.M.S. Challenger during the years

I 1873-1876. Zoology 1(2): 1-41.

Kiikenthal, W. 1913. Ueber die Alcyonarienfauna Californiens und ihre tiergeographischen Beziehungen.

I Zoologische Jahrbucher Abteilung fur Systematik 35(2): 219-270.

Kiikenthal, W. 1915. Pennatularia. Das Tierreich, 43: 1-132. Verlag von R. Friedlander: Berlin.

I Kiikenthal, Expedition W. and auf H. Bro demthDampfer . 1911. Pennatulacea. "Valdivia" 1898-1899, Wissenschaftlie 13(1)hLieferu e Ergebni ng 2:113-576. sse der deutschen TiefseeLjubenkov, J. 1980. Cnidaria. In: Sraughn, D. and R.W. Klink (eds.). A taxonomic listing of common

I marine invertebrate from southern California. Allan Hancock Foundation Technical Report species

3, Los Angeles, California. Pp. 44-68.

I Morin, J.G. Coelenterate 1974. CoeBiology: lenterateReviews bioluminescence. and New Perspectives. Pp. 397-438. Academic In: L. Muscatine Press: Newand HYork, .M. Lenh NY. o501 ff (eds. pp.)

Morin, J.G. 1976. Probable functions of biolurninescence in the Pennatulacea (Cnidaria, Anthozoa). Pp.

I 629-638. In: G.O. Mackie (ed.). Coelenterate Ecology and Behavior. Plenum Publishing Corp: New York, NY.

I Moroff, Th. und 1902 Biologie . Studien der Theire, ueber Octocorallien. 18(I 8): 363-410. Zoologische Jahrbucher, Abteiling fur Systematik, Geographie

Nutting, C.C. 1909. Alcyonaria of the California coast. Proceedings of the U.S. National Museum, 35: 681-

I 727.

Nutting, C.C. 1912. Descriptions of the Alcyonaria collected by the U.S. Fisheries Steamer "Albatross"

i primarily in Japanese waters during 1906. Proceedings of the U.S. National Museum, 43(1923): 1- 104.

II 81
!

!

Pasternak, F.A. 1962. Pennatulids of the genus UmbeUuIa Cuvier from the Antarctic and Subantarctic. [In m

Russian]. Issledovaniya Fanny Morei 1: 105-128. I

Pasternak, F.A. 1970. Sea pens (Octocorallia, Pennatutaria) of the hadal zone of the Kurile-Kamtschatka

Trench. Trudy Instituta okeanologii. Akademiya nauk SSSR 86: 236-248. [In Russian]. I

I

Pfeffer, G. 1886. Neue Pennatuliden des Hamburger Naturhistorischen Museum. Mitteilungen Zoologisches

Museum und Institut, Hamburg 3: 53-61.

Ricketts, E.F., J. Calvin and J.Hedgpeth. 1968. Between Pacific Tides. 4th Ed. Stanford University Press:

Palo Alto. 614 pp.

Stearns, R.E.C. 1873a. Remarks on a new alcyonoid polyp from Burrard' s Inlet. Proceedings of the California !

Academy of Sciences 5(1): 7-12.

Steams, R.E.C. 1873b. Description of a new genus and species of alcyonoid polyp. Proceedings of the •

California Academy of Sciences 5:147-150

Studer, T. 1889. Supplementary report on the Alcyonaria collected by the H.M.S. Challenger during the

years 1873- 1876. Challenger Reports: Zoology 32(81): 1-31. I

Studer, T. 1894. X. Note preliminaire sur les alcyonaires. Reports on the dredging operations off the west

coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California,

in chage of Alexander Agassiz, carried on by the U.S. Fish Commission steamer 'Albatross', during

1891, Lieut. Z. L. Tanner, U.S.N., commanding. Bulletin of the Museum of Comparative Zoology

25(5): 53-69. I

Verrill, A.E. 1864. List of the polyps and corals sent by the Museum of Comparative Zoology to other

institutions in exchange, with annotations. Bulletin of the Museum of Comparative Zoology at mm

Harvard

College 1(3): 29-60. I

Verrill, A.E. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expedition, under

Commodore C. Ringgold and Captain John Rogers, U.S.N., from 1853 to 1856. Collected by Dr. •

Wm. Stimpson, naturalist of the Expedition. With description of some additional species from the

west coast of North America. Proceedings of the Essex Institute, Salem, Mass 4: 181-196.

Verrill, A.E. 1868. Notes on Radiata in the Museum of Yale College. No. 6. Review of the corals and I

polyps of the west coast of America. No. 7. Geographical distribution of the polyps and corals on

u

the west coast of America. Transactions of the Connecticut Academy of Arts and Sciences 1(2):

377-558. I

Verrill, A.E. 1870. Notes on Radiata in the Museum of Yale College. No. 7. Geographical distribution of

the polyps and corals on the west coast of America. Transactions of the Connecticut Academy of •

Arts and Sciences 1(2): 558-570. I

Verrill, A.E. 1922. Revision of additional Canadian Alcyonaria, with descriptions of two new genera and

some new species. Report of the Canadian Arctic Expedition, 1913-18. Vol. 8. Mollusks,

Echinoderms, Coelenterates, etc. Part G. Alcyonaria and Actinaria. pp. 9-87. I

Williams, G.C. 1995. Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key •

and synopses. Zoological Journal of the Linnean Society 113:93-140. I

Wright, E.P. and T. Studer. 1889. Report of the Alcyonaria collected by H.M.S. Challenger during the years

1873-1876. Challenger Reports: Zoology, 31(4): 1-314. D

II

82 I
!

I 3. CLASSANTHOZOA

I ORDERS ACTINIARIA, CEmWrHAaIA,AND ZOANTHINARIA

by

I Daphne GaffFaufin _

!

Introduction

I Members of the three orders covered inthis section, which belong to anthozoan subclass Hexacorallia,

resemble one another inlacking a skeleton. Corallimorpharia, a fourth order sharing those attributes, occurs

I in the northeast Pacific, but there were no specimens of it in this collection. The fifth non-skeleton-forming hexacoral order,Ptychodactiaria, consists of three monotypic genera confined to high latitudes. All of these

animals may be referred to as sea anemones, but I prefer to reserve that term for actinians. The common

I name group.often used for cerianthids is "tube anemones," and Irecognize none other than "zoanthids" for the third

Few hexacoral faunas are sufficiently well known thatspecies identifications can be made exclusively,

I or even primarily, from living specimens. Descriptions of many species do not specify color, expanded posture, behavior, or other attributes absent in preserved specimens. This is because many species of

especially deep-sea and tropical anthozoans were described by expeIts who did not see them alive, not

I having participated in the expeditions on which they were collected (Dunn, 1981). These anatomically simple animals possess few distinguishing features. Further, the size and form of askeletal anthozoans can change

while the animals are alive and certainly during the processes of collection and preservation. Thus, variety,

I size, and distribution of cnidae, and histological features of the musculature may be critical in diagnosing species. Nature of the marginal sphincter muscle, if any,can be especially important. Although not included

in this report, I used these charactersin verifying identifications of the specimens inthis collection from the

I Santa Mafia Basin and Western Santa Barbara Channel. Because identification of these animals relies largely on features that persist in preserved specimens, it is likely that cryptic/sibling species are unrecognized

(Knowlton, 1993).

I No single external feature can unambiguously be used to assign these non-skeletalized anthozoan polyps to order. A key to the orders precedes the narrative concerning each order. That narrative includes

information about external anatomy, which suffices for assignment to order, and internal anatomy, which is

I essential for most generic and species identifications. The narrative of Order Actiniafia is followed by a dichotomous key to the 12 actinian species (three identified only to genus). There is one species of Zoanthinafia

in the collection and no specimen of Ceriantharia could be identified to species. A glossary contains terms

I necessary to understand the keys and taxonomic discussions.

The mid-water fauna of the vicinity where these specimens were collected has northern, southern,

western, and endemic components (Ebeling et al., 1970). The distribution of most deep water actinians is

I poorly documented. Members of Edwardsia, Epizoanthus, Pachycerianthus, and Peachia are

genera

widely distributed, if not cosmopolitan; those of Actinostola are typical of the deep seas at high latitudes; and

those of Hormathia are worldwide in deep seas. Sea anemones of the genera Edwardsia, Halcampa, and

!

I _ DepartmeUSntAof Systematics and Ecology, and KU Natural History Museum, University of Kansas, Lawrence, KS 66045
!

|

Anemonactis occur off Plymouth, England (Walton and Rees, 1913), and those of Metridium are common at •

least in shallow British waters (Fautin et al., 1990). Thus, at the generic level, this fauna appears unremarkable

for northern latitudes/deep seas. Halcampoides purpurea apparently occurs in all boreal waters, Peachia i

quinquecapitata occurs in both the eastern and western North Pacific, Hormathia digitata had previously l

been reported from boreal/North Atlantic waters, and Actinostola carlgreni was previously known from

Sagami Bay. Thus, these species are widely distributed. In contrast, Edwardsia californica, Epizoanthus •

induratum, Halcampa decemtentaculata, Metridium giganteum, and Pentactinia californica may be

endemic to the northeast Pacific. A great deal more research on both taxonomy and distribution is necessary

to understand the biogeography of actinians, cerianthids, and zoanthids, i

Biology I

The animals in this collection are infaunal except for the zoanthid (members of this class probably

are never infaunal) and three species of actinians. This habitat is correlated with inconspicuousness, and, in i

actinians, with small body size, which contribute to a dearth of information. Indeed, as mentioned in the l

remarks of some species, other than descriptions, there is no primary literature on them, and little or nothing

is known of their biology, i

Glossary of Morphological Terms ii

Aeontia. Nematocyst-studded threads arising from acontia may be carried through cinclides by., 

mesenteries of some actinians. May be the fluid, i 

extruded through mouth or body wall. Their

i

cnidom (qv.) may define actinian families. Clone. Collective term for genetically identical

Some cerianthids possess comparable individuals formed by asexual reproduction I

that are not physically connected to one i

structures called acontioids, another.

Aclinopharynx. Tube of tissue extending from i

mouth into coelenteron, continuous with Cnidae. Collective term for nematocysts,

oral disc. Unique to and present in all ptychocysts, and spirocysts. The sine qua

non of cnidarians, these intracellular

allthozoans.

"stinging capsules" secreted by the Golgi i

Actinostola rule. Members of pair of mesenteries apparatus are used mainly in offense and

unequal in size such that retractor muscle of defense. i

the better-developed member faces a

Cnidom. Types of cnidae possessed by a i

mesentery of the preceding cycle;

characterizes some genera in actinian family cnidarian or a part thereof. i

Actinostolidae. Coelenteron. Central body space of a cnidarian 1

Capitulum. Division of column in zoanthids and (also called gastrovascular cavity).

i

some actinians distal to and thinner than Coenenehyme. Tissue connecting polyps of •

scapus, colonial anthozoans. II

Cinclides. Plural of cinclis. Pores in the body Colony. Collective term for genetically identical

wall, or thinnings that can be ruptured to individuals formed by asexual reproduction 1

form pores, through which coelenteric fluid that are physically connected. II

can be expelled; in animals with acontia,

!
!

I Column. Tubular "body" of an anthozoan. Perfect. See mesentery.

Complete. See mesentery. Physa. Basalmost portion of actinian in some of

I those acfinians that have a divided colunm; Conehula. See siphonoglyph, used to dig or anchor in sediment.

i Couplme e. Tsenteries) wo mesenonteriopesposite (or 2 spides airs ofof column Protocnem attached e. One to cerian of 3 ctohiduplsiphonoglyph ed mesenteriesand

that arise simultaneously, immediately flanking the radius bisecting the

I Distal. Toward oral disc, away from base. siph symmetry onoglyphof the(thupolyp). s forming plane of

Lhnbus. Where pedal disc meets column.

Proximal. Toward base, away from oral disc.

I Maeromcnuemsclee. andMesentery filaments;with oftenstrong gametoretract genic,or Ptyehocyst. A type of cnida occurring only in

commonly complete, cerianthids; used in tube formation.

I Where oral disc meets column. Scapulus. Histologically differentiated division of Margin.

column in some actinians that is distal to

Mesentery. Radial sheet of tissue extending from

scapus and typically thick.

I bod are y"complete" wall. Thoseor rea"perfect"; ching actinhoseopharearynxching Scapus. Division of column in zoanthids and

only part way are "incomplete" or some actinians distal to base; commonly

I "imperfect." May be added singly (in constitutes most of a polyp's length. cerianthids) or in pairs and couples (qv.) (in Siphonoglyph. Longitudinal channel in

actinians and zoanthids), actinopharynx down which water can be

I Microcneme. Incomplete mesentery with weak driven due to longer cilia lining it. Oral end

retractor muscle; lacking filaments and may be drawn out into lobed projection

gametes, called conchula.

I Nematocyst. A type of cnida; there are about 30 Solitary. Polyps not physically attached to others.

varieties. Solitary polyps may not be genetically

unique: see clone.

I Pair. Two adjacent mesenteries that arise

simultaneously; equal in size (typical of Spirocyst. A type of cnida made only by

acfinians) or not (typical of zoanthids), anthozoans; its tubule is sticky.

I

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!

I

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I

Key to the Orders of Non-Skeletalized Hexacorallia •

1A. Colonial ......................................................................................................................... Zoanthinana m

1B. Solitary ............................................................................................................................................ 2

2A. Polyp encased in felt-like tube ...................................................................................... Ceriantharia m

2B. Polyp naked or with adherent detritus ............................................................................................. 3

l

3A. Tentacles in two distinct cycles ..................................................................................... Ceriantharia

3B. Tentacles around the margin only, or not arrayed in cycles ........................................................... 4 mw

4A. Tentacles arising from oral disc (and perhaps from margin as well) ................................ Actiniaria m

4B. Tentacles arising only at the margin ............................................................................................... 5 m

5A. Aboral end rounded or pointed, unattached ...................................................................... Actiniaria m

5B. Aboral end fiat, adherent or not ...................................................................................................... 6

6A. Column encrusted with sand or other debris ................................................................. Zoanthinaria m

6B. Column smooth ................................................................................................................. Actiniaria

I

Order Acfim_a n

Sea anemones are exclusively solitary, although some reproduce asexually to form clones. They are m

far more diverse ecologically and morphologically than are either cerianthids or zoanthids. Most oftbe taxa l

in this collection are infaunal, and therefore the animals are small, vermiform, and not attached to a hard

substratum -- attributes atypical of members of the order as a whole.

An external feature important to identification is number, arrangement, length, and form of tentacles, m

They may be cyclically or radially arrayed on the oral disc, be scattered, or exhibit a combination of those

arrangements; commonly they are simple, tapered cylinders, but they may be branched, capitate, or swollen m

along their length. The other major external feature is nature of the column. It may be divided into regions. I

The form of the basalmost portion determines whether the animal can attach to firm substrata or burrow in

sediment (abasilarian anemones, lacking basilar muscles, cannot adhere to substrata f'Lrmly so generally •

burrow; basilarian anemones have basilar muscles so can adhere firmly to substrata but may burrow). Some |

or part of the column may be encrusted with debris, have a cuticle, or be naked; it may have tubercles,

adhesive warts, or other specializations. •

Internal anatomy is also critical to accurate identification. For specimens in this collection, number I

and array of mesenteries, and features of the actinopharynx suffice.

Nearly 1,000 species of sea anemones are known (Carlgren, 1949). About 50 have been documented 1

from the northeast Pacific, half primarily from the intertidal zone and the others primarily subtidal. However, m

the bathymetric and geographic distributions of most are poorly resolved. In this collection I identified 50

specimens in 12 species (but see the caveat concerning identification in the Introduction) belonging to seven m

m
!i

i

I families; I was able to identify three species only to genus. Halcampoides purpurea was not previously

recorded from the Pacific; Anemonactis mazeli, Hormathia digitata, and Actinostola carlgreni were not

I previously recorded from the eastern Pacific, although, based on museum collections, the last two appear to be common in these waters.

The remaining specimens, which were too small and/or poorly preserved to be identified, seem to

I bel specimens ong to thefromspecPhase ies other I (Sta. than21th[oxsel] Iand identified Sta. 27to[xeither 3]) belong genustoorone species. speciesFourof very familyelonEdwardsiidae, gate, vermiformbut

cannot be identified further because their internal anatomy is indistinct. Their body form, absence of a strong

I cPhase uticle,I (Sta. and lack 1 [x4], of lSta. ongitudinal 14 [xl], grooves and Sta. 21distinguish [xl]) belong themtofroma singleEdward species, sia but californica the bases. Six are tomothersfromfroall m

of them and some are otherwise damaged also; they cannot be identified. Some of the remaining unidentified

I toI 1verify specimens with may the species belong dtoiscussions these species. identifica Because tions these madeanimals by use ofarethenotkey. included in the key, it is important

I Key to the Species of Actiniaria

I 1A. Column lacking adherent base; commonly vermiform (long, narrow; basal end may be pointed) or globular ............................................................................................................................................ 4

lB. Column with adherent base; not vermiform (may be as wide as long) ........................................... 2

i!

i 2A. Column smooth ................................................................................................................................ 3

I 2B. Column with cuticle and grooves (large specimens nodular) ............................. Hormathia digitata

i 3A. Column not divided into regions (commonly white, short relative to width; tentacles short but most margmat) ................................................................................................... Actinostola cf. carlgreni

3B. Column divided into long scapus, short capitulum (commonly yellowish, often very long relative to

I width; entire oral disc tentaculate) ................................................................. Metridium giganteum

4A. Tentacles capitate (usually 20, may be fewer) ................................................. Anemonactis mazeli

i 4B. Tentacles not capitate ...................................................................................................................... 5

I 5A. Five pairs of complete mesenteries (other, smaller ones may be present) ...................................... 6

5B. Other than five pairs of complete mesenteries ................................................................................ 8

I 6A. One distinct siphonoglyph ................................................................................................................ 7

6B. No distinct siphonoglyph; tentacles ten, stubby; five pairs each of macrocnemes and mierocnemes

I ........................................................................................................... Halcampa decemtentaculata

7A. Siphonoglyph drawn into conchula (column indistinctly divided; smooth but may be wrinkled,

I sausage-like constrictions) ................................................................ quinquecapitata

have Peachia

7B. No conchula; mesenteries bilateral about siphonoglyph (column often sand-encmsted; 20 tentacles)

i .................................................................................................................... Pentactinia californica
!

m

8A. Eight complete mesenteries ............................................................................................................. 9 []

8B. Six pairs of complete mesenteries ................................................................................................. 10

!

9A. Column cross-section resembles 8-pointed star ............................................ Edwardsia califomica

9B. Column globular; densenly covered with nemathybomes ......................................... Scolanthus sp. m

10A. All mesenteries complete ........................................................................... Halcampoides purpurea m

10B. Six pairs of macrocnemes; at least six pairs of microcnemes ....................................................... 10 m

l

11A. Column with adherent debris ................................................................................. Calamactinia sp. m

II

1lB. Column with dark cuticle; physa may be flattened ................................................. HalcampeUa sp.

m

Description of Species m

Genus Hormathia Gosse, 1859 I

Diagnosis. Hormathiidae (Acontiaria) with column divided into scapus and scapulus; scapus with m

protrusions except sometimes in young individuals, and with a cuticle. Mesogleal sphincter strong. Maximum l

number of tentacles about 96. Mesenteries of first cycle complete, sterile.

im

Remarks. This genus (indeed, the entire family Hormathiidae) is well represented in the deep sea. •

Carlgren (1949) listed 15 species in the genus, two with uncertainty, inexplicably omitting one species he I

himself had described. The considerable taxonomic confusion is probably partly attributable to the variable

development of promasions on the scapus, mm

Hormathia digi_ta (Miiller, 1776)

Figure 3.1 1

Actinia digitata Miiller, 1776:231. m

Hormathia digitata: Carlgren, 1942:44 (includes synonymy); 1949:93 (synonymy).- Grebel'nyi, 1980:12 m

(includes synonymy).

Material Examined. California: Santa Maria Basin, Phase I, Sta. 76 (x2); Phase II, Sta. R-6 (xl). mI

Description. Column faintly or strongly divided into longer scapus, with small, pointed to domed,

scattered protuberances that may be arrayed in vague longitudinal columns; and short scapulus, with a I

coronal ring of protuberances just below the tentacles. Animals examined columnar, with pedal disc expanded 1

or not, to about 10 mm column diameter and height; members of the species grow much larger. Commonly

opaque white with golden-brown cuticle adherent in cracks and on tips of protuberances.

Pedal disc may adhere to gastropod shell (as that from station R-6). Oral disc usually retracted; I

with 48-72 short, often nipple-shaped tentacles in specimens examined; large individuals with up to 96.

Mesenteries regularly hexamerously arranged; those of 6 primary pairs complete and sterile, others 1

fertile. Acontia prominent in specimens examined: coiled spring-like, protruding into actinopharynx and even

out mouth.

Cnidom: Spirocysts, basitrichs, microbasicp-mastigophores (acontia contain only basitrichs). I

88 m
I

I

I

I

I

I

I

I

I A B

I

i Figure 3.1. Hormathiadigitata:A,specimenwithpronouncedprotuberances(fromGrebel'nyi, 1980);B,specimen with lessprominent protuberances attached to crustacean Hyas (fromCarlgren, 1942).

I

Biology. Actinians of this species axe abundant at depth in the northeast Pacific. They apparently

i require a firm substratum but do not seem particular about it -- they often occur on gastropod shells or other living animals (see Fig. 3.1B) and have even been collected attached to rubber gloves.

Remarks. The identity of this species has been uncertain. Grebel'nyi (1980) convincingly argued

I for (typethespesyno ciesnymiz of theatigenus) on of inHormath the Northia Atla nodosa ntic,, Hasser . jostiephi ng ,thatH.variability incubans, between and H. individuals marioni withis responsible H. digitata

for the plethora of names. He suggested that a fifth species, H. coronata, belongs in hormathiid genus

I Acti America, nauge.despite Therelarge are vnumbers irtually noof rec specimens ords of any in museums, species ofprobably this genubescause alongoftheuncertainty Pacific coainsttaoxonomy f North

resulting from variability among individuals.

I Type Locality and Specimens. Species described from the waters of Norway or Denmark; no type specimen designated.

Distribution. Circumarctic; southern limits of distribution in North Pacific unknown. Depth range

I at least 10-1,000 m, based on Arctic and North Atlantic records.

I

I 89
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Genus Actinostola Verrill, 1883 •

Diagnosis. Actinostolidae (Mesomyaria) with thick, firm column and comparatively weak sphincter. •

Inner tentacles longer than outer, bases sometimes thickened on the aboral side; outer ones with microbasic I

b-mastigophores distally. Longitudinal muscles of tentacles and radial muscles of oral disc mesogleal.

Mesenteries of the first two cycles sterile; retractor muscles of those from the third or fourth cycle onward •

arrayed so that the one nearer the mesentery of the preceding cycle is better developed than its parmer (the I

Actinostola rule).

Remarks. This genus (indeed, the entire family Actinostolidae) is well represented in the deep sea. •

Carlgren (1949) listed 12 species in the genus, one of which Riemann-Ziirneck (1978) placed in its own I

genus. None is unequivocally known from the northeast Pacific.

n

Aetinostola cf. carlgreni Wassilieff, 1908 1

Figure 3.2 1

Actinostola carlgreni Wassilieff, 1908:28. -- Carlgren, 1949:78.

Catadiomene carlgreni: Stephenson, 1920:558. 1

Material Examined. California: Santa Maria Basin, Phase I, Sta. 17 (xl).

Description. Column nearly as long (42 mm) as wide (50 ram). Oral disc only slightly retracted so •

tentacles visible. Limbus extended beyond level of pedal disc but same diameter as column. Tan ectoderm 1

remaining only in shallow circumferential wrinkles. Tentacles short, outer mostly shorter than inner, all

lacking ectoderm. So entire animal white in color (from exposed mesoglea). •

Short sphincter muscle mesogleal, weak; narrow, adjacent to endoderm of margin, less than 20% 1

width of margin.

All mesenteries except those of highest order complete, sterile. Those of highest order fertile 1

(questionable in individual examined), extend from base to oral disc; members of highest-order pairs slightly

unequally developed (that is, they follow the Actinostola rule).

Cnidom: The genus is characterized by spirocysts, basitrichs, microbasic p-mastigophores, and 1

microbasic b-mastigophores, but because of the erosion of ectoderm, this could not be ascertained in the

u

specimen examined.

Biology. Unknown. n

Remarks. Clearly a member of family Actinostolidae, this animal appears tobelong toActinostola.

The only nominal species of the genus that has been documented in the eastern North Pacific, A. •

spetsbergeneis, was transferred by Riemann-Ziirneck (1978) to the new genus Glandulactis based partly 1

on the absence of mesenteries arrayed according to the Actinostola rule. This specimen agrees well with

Wassilieff's brief description of A. carlgreni. •

Type Locality and Specimens. Sagami Bay, 730 m. Holotype in the Zoologische Staatssammlung

des Bayerischen Staates, Munich, Germany.

Distribution. Actinostola carlgreni is definitely known only from Sagami Bay. If this specimen 1

really belongs toA. carlgreni, the species is at least circum- or trans-North Pacific. This specimen was from

654 m.

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i Figure 3.2. Actinostolagroenlandica (fromCarlgren,1921;slightlylargerthanlife):the numerous,shorttentacles and wrinkled column are typical of preserved specimens of all speciesof the genus.

I

I Genus Metridium

i Diagnosis. Metridiidae (Acontiaria) with mesenteries not divisible into macrocnemes and microcnemes; acontia with basitrichs, microbasic b-masfigophores, and microbasic amastigophores.

Remarks. Authorship of this genus is in doubt, as summarized by Manuel (1981 a). Oken is credited

made with ittobyhave Carlgre de Blainville n (1949), rebuctognized his nameassthe haveauthor. been declared nomenclaturally invalid. An appeal has been

I Metridium giganteum Fautin, Bueklin, and Hand, 1990

Figure 3.3

i Metridium senile: see below under Remarks

Material Examined. California: Santa Maria Basin, Phase I, Sta. 4 (x2); Phase I, Sta. 27 (x2);

i Phase I, Sta. 28 (xl), in addition to all type specimens and many others.

Description. Column long (subtidal individuals may attain 1 m height), relatively narrow (column

i height/pedal disc diameter >2:1); tapered distally; prominent collar between long scapus and shorter capitulum

(Fig. 3.3). Commonly opaque white, occasionally salmon orange or brown. Lacking cinclides.

Pedal disc adherent to firm substratum; diameter typically 125 mm, maximum 200 mm. Oral disc

i much broader than column; stiffly lobed (Fig. 3.3B); marginal portions densely covered with hundreds of

m 91
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A B 1

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Figure 3.3. Metridium giganteumin str_ngcurrent (fr_mK_eh___976): A_side view;B_c__umnand underside _f I

oral disc. Note raised collarnear oral end of column, and bracing of oral disc lobes.

!

short, pointed tentacles giving disc frilled or fluffy appearance (Fig. 3.3A); tentacles of central oral disc

longer, sparser. Tentacles and oral disc same color as column. No catch tentacles.

I

Mesenteries regularly hexamerously arranged due to the virtual absence of asexual reproduction. 1

Three specimens examined are very small (3-10 mm long), resembling Diadumene. However,

cnidae agree very closely with those of Metridium and differ from those of species of Diadumene and allied •

genera. Moreover, of actinians with acontia, thin walls, and columnar shape, only Metridium is known from

depth in the northeast Pacific.

Cnidom: Spirocysts, basitrichs, microbasic b-mastigophores, microbasicp-mastigophores, microbasic

amastigophores.

Biology. Actinians of this nearly exclusively subtidal species can achieve the greatest height and

mass (but not breadth) of any sea anemone. They are common on pilings in shallow water, and occur deeper 1

on firm objects such as shells. Populations consist of solitary individuals or groups of genetically distinct ones.

1

Reproductive potential is almost exclusively sexual: individuals rarely fragment, and excised fragments

i

regenerate infrequently and very slowly, in contrast to those of M. senile.

Remarks. Three species of Metridium occur along the west coast of North America: M. exilis, M.

senile, and M. giganteum (see Fautin et al., 1990). Metridium exilis is morphologicallyl biologically, and

biochemically distinctive. The other two were previously considered to belong to one species, M. senile, a

|

highly variable species first described from the northeastern Atlantic. Compared withM, senile, M. giganteum

is less translucent, its column mesoglea is thicker and stiffer, it feels more slippery, and its height:width ratio !

92 I
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I is greater; its retractor muscles are larger, and its sphincter muscle is relatively much larger; permanent oral

disc lobes are reinforced by mesogleal thickenings in larger individuals of M.giganteum, whereas the oral

I Mdisc. senile of M. aresenildistinct, e is unlowhile bed when the oral small discanofd lMacks . giganteum mesoglealappears thickenings fluffy.when For biochemical large; individdifferences, ual tentaclesseeof

Bucklin and Hedgecock (1982). Nematocysts do not differ consistently between the two, their size seemingly

I correlated more with size than species of the individual. This interesting, ecologically important genus is the subject of a variety of research, including

biomechanics (e.g. Koehl, 1976, 1977) and theoretical ecology (e.g. Sebens, 1981). From the literature, it is

l not always obvious which species was studied. Type Locality and Specimens. Monterey County, California, 2.4 km SW of Point Pifios Lighthouse,

80 m. Holotype CAS #003349 fromAlbatross station 4439; paratypes CAS #004370 fromAlbatross station

I 5787: San Mateo County, California, 13.3 km W of Point Montera Light, 75 m (x2); CAS #003508: north of

Famllon Islands, off Cordell Bank, 182 m (x2); USNM #85789 from Albatross station 5789: San Francisco

County, California, 6.3 km W of Farallon Light, 84 m (x l); SBMNH #45511: off Gaviota, Santa Barbara

I County, California (ca. 34°23'N 120°10_¢), 256 m (xl); SBMNH #45512: off Gaviota, Santa Barbara County, California (ca. 34°23q_ 120°10'W), 201 m (xl); RBCM #986-79-11: Hecate Strait, British Columbia (54° 12'N

13I°3.4"W, 99 m) (x3).

I Archipelago, (56-58°N) (or north) along coast

Distribution. From Alexander Alaska further Pacific

of North America through California, in bays or open water. Typically subtidal; previously known range

extended by specimens in this collection to 611 rm

Genus Anemonactis Andres, 1881

i Diagnosis. Haloclavidae (Athenaria) with elongate, cylindrical column not divided into distinct regions, and lacking sphincter. Column with cinclides in uppermost part, and scattered solid papillae; rounded

aboral end perforated by numerous apertures. Capitate tentacles 20, inner longer than outer.

I Remarks. Anemonactis mazeli, the species, is the best known of the three nominal species;

type

the other two are from Australia, and may be synonymous (Carlgren, 1949).

I Anemonactis mazeli (Jourdan, 1880)

Figure 3.4

llyanthus mazeli Jourdan, 1880:41.

Anemonactis magnifica Andres, 1881:329.

I Eloactis mazeli: Andres, 1883:257. 1892:380. Faurot, 1895:152. Rees, 1913:70.

Garstang, l

Walton and Rees, 1913:68. -- Stephenson, 1935:91.

i Halcampella Eloactis mazeliminuta i: Carlgren, Wassilieff, 1921:111. 1908:7. -- Uchida, 1938:288.

Haloclava minuta: Carlgren, 1949:30 (synonymy).

I Anemonactis mazelii: Carlgren, 1949:31 (synonymy).

Anemonactis mazeli: Manuel, 1981a:184 (includes synonymy).

i Material Examined. California: Santa Basin, Phase II, (xl).

Maria Sta. PJ-7

Description. Cylindrical scapus 7 mm long, 4 mm broad; yellowish, with minute solid papillae;

connected with physa by constriction. Physa flattened, diameter 4 nun, with adherent debris.

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B

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Figure 3.4. Anemonactis mazeli (fromUchida, 1938):A, whole specimen;B, cross-section. /

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Tentacles 10, relatively short; each with distinct, lighter-colored capitate end (Fig. 3.4) comprising i

dense, long (more than 100 ram), thin (3-4 ram) basitrichs, as well as sparse spirocysts. |

Cnidom: Spirocysts (few seen in this specimen but reported by Carlgren [1921] to be absent),

basitrichs, microbasic p-mastigophores. I

Biology. Reported by Manuel (1981a: 185) to burrow "in mud, sand or gravel, always offshore." iN

Remarks. Rees (1913) provided the most thorough description of the species, Stephenson (1935)

summarized biology and external morphology, and Carlgren (1921) detailed internal anatomy and histology, i

Jottrdan (1880:41) remarked "Comme chez tousles Actiniens, sa forme est tl'_Svariable et d6pend beaucoup

u

de l'6tat d'extension ou de contraction darts lequel se trouve l'animal." This specimen conforms to those

accounts except in having only 10 tentacles. However, it is also smaller than those studied, and Carlgren l

(1921) examined what he reported as a "young" specimen in which he found only 11 tentacles.

It

Wassilieff's (1908) description and figures of Halcampella minuta are clearly of this species. A

Carlgren's (1949) uncertainty about the generic placement of the species may be because of its possession of |

only 15 tentacles (which presumably was related to its small size -- 10 mm long) and its Japanese locality.

These attributes are now known to be consistent with Anemonactis mazeli.

Iml

Type Locality and Specimens.No type specimens recorded. Originally described from 60-80 m in I

the northeastern Gulf of Marseilles.

Distribution. Previously known from the Mediterranean, North Sea, Japan, from 20-650 m. This i

specimen is from 123 m. I

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Genus Halcampa Gosse, 1858

i Diagnosis. Halcampidae (Athenaria) with elongate, cylindrical column usually divided into physa, scapus, and capitulum, and with a weak sphincter. Mesenteries divided into macrocnemes and microcnemes.

Remarks. Carlgren (1949) listed six species in the genus, including one with a query. Since then,

t three additional species, including Halcampa decemtentaculata, have been described.

i Haleampa decemtentaculata Figure 3.5 Hand, 1955

l Halca HaIcampa mpa decemtentaculata Hand, 1955: 360 (includes synonymy); 1957:413. duodecimcirrata: Carlgren, 1936:19; 1949:34 pro parte. -- Cutress, 1949:12.

I Phase I, Mater Sta. 52ial(xl)Examin ; Phaseed.I, StCal a.i6for5 ni(xl)a:; Sant Phasea Mari I, Staa.Basin, 74 (xl);PhaPsehaseI, SI,taS.ta.9 79(xl);(xl)Phase ; PhasI,e SII,ta.Sta25. P(J-10 xl);

(xl); Phase II,Sta. PJ-14 (xl).-- Weatem Santa Barbara Channel, Phase I, Sta. 88 (xl).

I tan; 5 x D2emm, scripti6xon.1.5Col ram, umn9 xvermiform, 2 ram, 17xdivided 3mm, in18toxd2.5istinmm, ct regions; 28x 8 specimens ram, 30 x 4examined mm, 45 xyellowish 8 ram, 45tox

10mm. Physa often bulbous, occasionally pulled in; scapus may have adherent debris; in expanded specimens

I capitu per endocoet lum slightly flared, thin so that at least macrocnemic insertions (10) visible through it. Several cinclides reported (Hand, 1955); only that at tip of physa discerned in some specimens examined.

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I Figure 3.5. Halcampa mesenteries. decemtentaculata:cross-sectionfrom Hand(1955)illustratingdistinctive arrangementof

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Ten tentacles stubby, cylindrical in expansion (to 2 mm long), most nearly as wide as long (about

0.5 mm) in contraction. Darkly pigmented endoderm in specimens examined. Tentacles and mouth area

darkly pigmented in two specimens; Cutress (1949) described variations in markings of oral disc. Mouth Ill

protuberant in one expanded specimen.

Ten macrocnemes and 10 microcnemes mn entire length of column; arrayed in 4 pairs each plus 1

pair composed of macrocneme and microcneme. Two pairs of macrocnemes directives: 1 pair flanked by i

pair of microcnemes then pair of macrocnemes; other pair flanked by mixed pair then pair of microcnemes 1

(Fig. 3.5). No or few microcnemes evident in smallest specimens examined. No acontia.

Crtidom: Spirocysts, basitrichs, microbasicp-mastigophores. 1

Biology. Anemones burrow in sand and gravel; Cutress (1949) reported they may expand and be

visible intertidally on overcast days. He also stated that the larvae are parasitic on medusae. I

Remarks. Hand (1955) provided detailed information about distinguishing Halcampa i

decemtentaculata from the similar species H. duodecimcirrata, with which it has been confused, and H.

arctica. Type Locality and Spec I imens. Havens Neck, Mendocino County, California. l_e,ctotype USNM

50637 (Hand, 1957).

Distribution. At least from Washington to southern California. It was previously known only from 1

the lower intertidal zone; specimens in this collection came from 98-398 m.

m

Genus Peachia Gosse, 1855 I

Diagnosis. Haloclavidae (Athenaria) with elongate, cylindrical column, and lacking sphincter.

Tentacles normally 12, inner shorter than outer. Oral end of single very deep siphonoglyph drawn out into If

lobate conchula, occasionally siphonoglyph entirely separated from rest of actinopharynx so it forms a tube.

m

Remarks. Larvae of all 11nominal species of this genus (and the one unnamed one) live parasitically /!

on medusae.

l

I

Peachia quinquecapitata McMurrich, 1913b ,_

Figure 3.6

df_

Peachia quinquecapitata McMurrich, 1913b:963.- Carlgren, 1949:32 (synonymy). l

Bicidium aequoreae McMurrich, 1913b:967.

I

Material Examined. California: Santa Mafia Basin, Phase II, Sta. PJ-6 (xl); Phase II, Sta. R-1 I

(x2); Phase II, Sta. R-2 (xl).

Description. Column of larger individuals vermiform, not divided into distinct regions; smooth It

aside from circumferential wrinkles that are presumably due to contraction; may also be constricted at one

or more points so animal appears like link sausages. Specimens examined retracted. Extended ones 30 mm

long, one with uniform diameter of 13 mm, other 22 mm at broadest, 12 mm at constriction. Two smaller •

specimens globular, also with circumferential wrinkles but longitudinally grooved at mesenterial insertions !

as well; one 6 mm broad by 4 mm long (it resembled a pumpkin), the other 10 mm long by 8 mm diameter.

Aboral end may be abruptly narrowed, nipple-like; commonly indented to depth of several ram. i

Twelve tentacles either nipple-like, mostly 2 mm long by 3 mm greatest diameter at base (some slightly

elongated radially), or digitiform and up to 4 nun long by 1.5 mm at base. Ectoderm of column and tentacles

pinkish-brown or yellowish, column may have indistinct white reticulation, particularly in larger individuals, i
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Figure 3.6. Peachiaquinquecapitata(fromMcMurrich, 1913b):A, whole animal;B, oral disc and tentacles.

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I prCionper clidillumination es not visible (Spaulding, in specimens1972). examined; reportedly discernible only in well-expanded specimens under

Mesenteries continuous entire length of column, distinctively arrayed in larger individuals: 6 pairs

i complete, musculatureincluding lack filaments. 2 pairs ofExocoels directives,flanking with diffuse 1pair retractor of directives muscles; lack secondaries; 4pairs secondaries other pair withof very directives weak

associated with deep siphonoglyph that is separated from actinopharynx, forming distinctive conchula at

I oral specimens end. Smaller examinedspecimens sterile. had No acontia. only 6 pairs of mesenteries, which extended entire length of animal. All

Cnidom: Spirocysts, basitfichs, microbasic p-mastigophores.

i Larvae of this of all members of the is medusae Biology. animal, as genus, parasitic on (Spaulding,

1972).

i Taxonomic Remarks. Some of the specimens examined conform precisely to the detailed original description (McMurrich, 1913b), including the depression in the center of the base, and the end of one being

narrower than the rest of the column. Although its vermiform or globular shape is not unusual, its internal

i anatomy is very distinctive. Contrary to my experience, McMurrich (1913b) wrote that its column is "intense brownish red," a coloration that may be retained in preservation. Bicidium aequoreae was described from

the parasitic larva. Although he thought it likely thatB, aequoreae represented the larva ofP. quinquecapitata,

McMurrich (1913b) considered it prudent to describe them separately because an association between parasitic

I larvae and the adult form had not been def'mitely established.

Type Locality and Slmeimens. Nanoose Bay, Vancouver Island, 15-20 fathoms. Type specimens

i could noDistribution. t be located: they FromareBritish not at CAS, Columbia RBCM, or USNM. south at least to the Santa Maria Basin. Known from low

intertidal; the previously known range was extended to 161 m by these specimens.

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Genus Pentactinia Carlgren, 1900

Diagnosis. Halcampoididae (Athenaria) with elongate cylindrical column and no sphincter. Four i1

pairs and two single perfect mesenteries, the latter paired with microcnemes. One siphonoglyph, 20 tentacles. I!

Remarks. Pentactinia is a monospecific genus.

i

Pentactinia cah'fornica Carlgren, 1900

Figure 3.7 i

Pentactinia califomica Carlgren, 1900:1166; 1931:33; 1949:29.

Material Examined. California: Santa Mafia Basin, Phase I, Sta. 4 (xl); Phase I, Sta. 16 (xl); i

Phase II, Sta. PJ-7 (xl); Phase 11,Sta. PJ-8 (xl); Phase II, Sta. PJ-22 (xl).

1

Description. Column vermiform, reportedly divided into regions but not obvious in specimens II

examined; ectoderm encrusted with fine sand. Aboral end (physa) of specimens examined indented; said to II

be distinct in extension (Fig. 1). Column of specimens examined 2-10 mm long x less than 1 to 3 mm broad.

20 tentacles small, short, outer nearly as broad as long, inner longer than outer; yellowish in extension or dark ll

pink when retracted. 1

Mesenteries continuous entire length of column, but strong retractor muscles, filaments, and gonads

(if any) of 10 macrocnemes confined to distal half. Single strong siphonoglyph connected with directive pair;

five other macrocnemes arrayed on either side of each directive, imparting very pronounced bilaterality to II

internal anatomy (Fig. 3.7A). Retractors sla'ong,reniform (Carlgren, 1931).

Data based mainly on 2 larger specimens; it was difficult to discem detail in the three smaller ones. i

g

Cnidom: Spirocysts, basitrichs, microbasic p-mastigophores.

Biology. Nothing is known about this animal. Aside from Cafigren' s original description and his i

redescription, this is the only published record of its occurrence. II

Type Locality and Specimens. San Pedro, Califomia; beach. Six specimens were said by Carlgren

(1900) to be in SMNH. Two syntypes are SMNH 1162. The others were thought by Lennart Sandberg iI

(personal communication), Curator at SMNH, to be in the Zoologiska Museet in Lund, Sweden, but they are II

not (personal communication from Lennart Cederholm, Curator, Zoologiska Museet).

Distribution. Southern California. Known from low intertidal to at least 591 m (specimens in this i

collection).

1

Genus Edwardsia de Quatrefages, 1842 I

Diagnosis. Edwardsiidae (Athenafia) with elongate, cylindrical column divided into physa, scapus,

scapulus, and capitulum; lacking sphincter. Short physa without nemathybomes (spheroidal invaginations of i

the ectoderm into the mesoglea, laden with nematocysts), long scapus with nemathybomes. Eight macrocnemes ll

and at least 4 microcnemes (very weak and confined to the distal portion of the body). Tentacles at least 12,

the inner shorter than the outer, i

I Remarks. Edwardsia is the largest genus of athenarian actinians and one of the largest of all sea

anemones, Carlgren (1949) having listed 56 species, one questionably. Williams (1981) recognized 40 species, i

and summarized the status of other nominal species. I

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i Figure 3.7. Pentactinia californica: A, cross-section, illustrating distinctive arrangement of mesenteries(from

Carlgren,1900);B, wholeanimal(fromCarlgren,1931).

l' Edwardsia cah'fornica (McMurrich, 1913a)

i Figure 3.8

Edwardsiella californica McMurrich, 1913a:551.

I Edwardsia californica: Carlgren, 1936:18; Carlgren, 1949:24. -- Williams, 1981:346 (synonymy).

Material Examined. California: Santa Maria Basin, Phase I, Sta. 56 (x2); Phase I, Sta. 78 (xl);

i Phase II,Description Sta. R-7 (xl); . Vermiform; Phase II, Sta. specimens PJ-10 (xl). examined to 30 mm long, 7 mm diameter. Seapus long, with

brown cuticle, prominent nemathybomes; some circumferentially wrinkled; most specimens examined

lostrngi iped, tudinall withyrgirdgooesvedligalong ht comacrocnem lored, groovesessodar thaktco(tlheumnse rsetripe sembs les ma8-point y be singedlestarorindcro oubsle). s-sectPiohnys;longi a short, tudinall wely

separated from scapus by constriction in most specimens, generally encmsted with t-me sand. 12 yellowish

I tentaclesMaincsrocn pecimemenes examined. s eight; microcnemes presumably present in distal portion of column butnot prominent.

Cnidom: Spirocysts, basitrichs, microbasic p-mastigophores. i
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Pigure 3.8. Diagrammatic sketch of Edwardsia (fromWilliams, 1981): A, oral disc and tentacles; B, expanded

animal, illustrating basal physa, long scapuswith nemathybomes,and distal scapus, i

Biology. Nothing is known about this species. /I

Remarks. McMurrich' s (1913a) description of whole animals and their histology conforms precisely |

to the specimens examined. Carlgren's (1936) brief treatment of the species provides data on cnidae, which

differ slightly from what I found. However, the cnidae of one specimen differed slightly from those of the 1_

other three examined, and so cnidae may be variable in this species. II

Carlgren (1936) transferred the species to the genus Edwardsia, a placement with which Williams

(1981) agreed. I Type Locality and Specimens. "Anaheim Bay (Creek), California: Muddy shore, about one mile

inland, at or below low water, one example; mudflats andshore at, or below, low tide, 'Stingaree Hole,' six

examples" (McMurrich, 1913a:551). Type specimens USNM 30716. I

11 Distribution. Southern California. Known intertidally to 900 m (this collection).

Genus Scolanthus Gosse, 1853 I

Diagnosis. Edwardsiidae (Athenaria) with column divided into scapus and scapulus; physa externally :Ul

identical to scapus, with cuticle and tenaculi. No sphincter. Eight macrocnemes. |

Remarks. Manuel (1981 b) discussed his resuscitation of Gosse's (1853) genus Scolanthus and his

opinion that Carlgren's (1921) Isoedwardsia is synonymous with it. However, he believed that not all six Ill

species Carlgren (1949) listed in that genus belong in Scolanthus. II

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I Scolanthus sp.

Material Examined. California: Santa Maria Basin, Phase I, Sta. 21 (xl); Phase II, Sta. R-1 (x7).

i Description. All specimens examined contracted, pear- to American football-shaped; maximum

length 4 ram, maximum diameter about 1 mm. Column orangish brown due to rough cuticle, with numerous,

i easily visible Ten tentacles nemathybomes. counted Eight in onemesenterial individual. insertions Blunt conical, visibleslightly entirelonger lengththan of columrt. broad.

Character, number, and arrangement of mesenteries obscure due to small size of animals.

t Cnidom: Spirocysts, basitrichs, microbasicp-mastigophores found.

Biology. Unknown.

i animals Rinemawhich rks. theyThesewereanimals countedagreehadwith 10 tentacles, Manuel's as(198contrasted 1b) diagnosis with of16 Scolanthus supposedly except in Scolanthus that two.

However, these specimens axe very small and so might be expected to have fewer tentacles. No species of

! • this genus or other similar edwardsiid is known from California. The specimens are so small as to preclude

B a more detailed diagnosis.

Genus Halcampoides Gosse, 1858

Diagnosis. Halcampoididae (Athenaria) with elongate, smooth, cylindrical column not divided into

I regions no microcnemes. (although aborai Two indis endtinmayct siphonoglyphs, resemble a physa), 12 cylindrical and no sphitentacles. ncter. Six pairs of perfect, fertile mesenteries;

Remarks. Several species of the genus have been described, but Carlgren (1921) synonymized all

I as Halcampoides

purpurea.

I Halcampoides purpurea (Studer, 1879) Figure 3.9

l Halcampa purpurea Studer, 1879:545. Halcampoides purpurea: Carlgren, 1921:82 (includes synonymy); 1949:27 (synonymy).- Manuel,

1981a: 182 (synonymy).

i Material Examined. California: Santa MariaBasin, Phase I, Sta. 13 (xl); Phase II,Sta. PJ-14 (xl).

Description. Column vermiform, not divided into distinct regions, although may be circumferentially

I cmesenterial ontracted along its length so it somewhat resembles a chain of sausages; longitudinally furrowed along insertions (12); smooth aside from circumferential wrinkles that arepresumably due to contraction;

aboral end expanded, reportedly with cinclides but not detectable in specimens examined. Ectoderm brown,

i scr column aped diameter, off particu15larlymmaroaboral und base end .diameter; Retracted30 specimen mm longs xexamined 10 mm maximum 48 mm loncolumn g x 13diameter, mm maxim13 ummm

aboral end diameter.

I Tentacles long, cylindrical, abmptly terminating in threadlike extension; retracted in specimens examined. Pale brown color.

Mesenteries distinctive: 6 pairs complete and fertile, including 2 pairs of directives, with very

i strong retractor muscles.

Cnidom: Spirocysts, basilrichs, microbasicp-mastigophores.

i

. 101
I

1

i

[2Sr I

I

I

I

Figure 3.9. Halcampoidespurpurea (fromManuel, 1981a): distal end of expanded animalin situ. I

III

Biology. This animal is said to be capable of great extension when alive-- Manuel (198 la) reported II

it is up to 100 mm long when not buried in sediment and can greatly exceed that when buried, with tentacles l

reaching 100 mm in length. It appears to be nocturnal in Galway Bay, to which Manuel (1981a) attributed the

paucity of records. _l

Remarks. Carlgren (1921, 1949) and Manuel (1981a) considered there to be only one species of n

this genus in the world. It is very distinctive. This species was given several names in various parts of the

world before its extraordinary range was recognized. Additional synonyms were given by Carlgren (1921); n

none was used for animals on this coast.

Carlgren (1921) summarized reports of some nematocysts and color in life: column is often suffused

with pink or red, and oral disc may have reddish radial lines on oral disc, vivid green lips, and white spots near I

mouth. He also illustrated muscle variability. U

Type Locality and Specimens. Kerguelen, South Central Indian Ocean, 6-100 fathoms; no type

specimens located, i

Distribution. Previously reported from both poles, around Britain, and the Mediterranean; thought

by Carlgren (1949) to be cosmopolitan. This is the first record of the species along the west coast of North II

America. Known from 1-1,134 m. I

Genus Calamactinia Carlgren, 1949 i

I!

Diagnosis. Halcampoididae (Athenaria) with elongate column divided into scapus and scapulus.

Sphincter absent. Six pairs of perfect, fertile macrocnemes, including 2 pairs of directives; microcnemes _i

also present. II

Remarks. This genus and its only species (C. goughiensis) are both listed as new in Carlgren's

(1949) catalog. There is no other description of either, Carlgren apparently not having found time to elaborate d

on them before his death within a few years of his catalog's being published. The genus Calamactinia is

listed in volume VI of Nomenclator Zoologicus (F_Awards and Hopwood, 1966) with the 1949 Carlgren

citation. Six lines of text in Carlgren's catalog are devoted to diagnosing the genus. In addition, there is a i

102 _i
!

I specific entry consisting of "C. goughiensis n.sp. SW of Gough Isl., 141-162 m." I interpret this presentation

as equivalent to a combined description of new genus and new species, which is deemed in Article 13(c) of

i timplies he International to me that,Code there of beingZoological only oneNospecies menclature in the genus "to confer as Carlgren availabiknew lity oit,n the eachgenename." ric description This further can

be interpreted as applying to the species.

I Calamactinia sp.

t Material Examined. California: Santa Maria Basin, Phase I, Sta. 2 (x4).

Description. Column bulbous, varying from club-shaped to American football-shaped. All animals

I examined scapus. Noretracted; physa, shor totalt length scapulus. 5-I5 mm, maximum width 2-5 mm. Sand, small bivalve shells, etc. adhere to

24 tentacles: inner longer than outer, but each about 1 mm long; narrow, tapering to pointed end.

I sphincter or siphonoglyphs discernible; pairs macrocnemes (including pairs

No muscle 6 of fertile 2

of directives) and 6 pairs of microcnemes extend entire length of column.

i Cnidom: Remarks.Spirocys Carlgrents, (1949) basitrichs, mademicrobasic no mention p-maofstigophores. the adherent debris, which is a conspicuous feature

of this animal. His generic diagnosis included the phrase "scapus ... with a rather thick, easely [sic] deciduous

I cuoftiCcle." . goughiensis I omitted. that Thefeature specimens from mfromy diagnosis California of thehave, genus, instead, choosing adherent to interpret debris,thatandfeature so, I ainfer, s characteristic belong to

a second species of the genus. Before it can be described, type material of C. goughiensis must be

I examined. Additional material of the Califomia species should also be studied.

Genus Halcampella Andres, 1883

i Diagnosis. Halcampoididae (Athenaria) with elongate column divided into physa, scapus (which

has tenaculi), and scapulus; no sphincter. Six pairs of perfect, fertile macrocnemes, including 2 pairs of

I directives; microcnemes also present. Two indistinct siphonoglyphs; tentacles short, stubby. Remarks. Three species were listed in this genus by Carlgren (1949), including H. robusta.

Presumably in Carlgren's (1931) description of H. robusta and discussions of H. mamma by Wassilieff

I (1908) and Carlgren (1931), references to a "capitulum" were actually to a scapus.

i HalcampeUa sp.

Material Examined. California:Santa Maria Basin, Phase I, Sta. 52 (×2); Phase II, Sta. R-8 (×1).

I Description. Column vermiform. Physa discoidal, mostly smooth but with remnants of cuticle, pulled in along mesenterial insertions, broader than most proximal part of scapus; crusty cuticle thicker on

scapulus than scapus, perhaps due to scapulus being introverted in all specimens examined; scapus, which

I represents most of animal's length, with easily eroded cuticle, may be greatly narrowed proximally. Ectoderm light brown, visible where cuticle eroded, particularly proximally; no adherent material. Retracted specimens

examined 23-25 mm long x 6-8 mm maximum column and basal diameter.

l Tentacles short, stubby; 12 in specimens examined.

Six pairs of mesenteries complete and fertile, including 2pairs of directives, with very strong retractor

I muscles. At least 1 cycle of microcnemes.

i 103
!

IN

Cnidom: Spirocysts, basitrichs, microbasic p-mastigophores. I!

Remarks. The gestalt of these animals is virtually identical to that of Halcampoides purpurea.

However, specimens of H. purpurea are smooth, have long tentacles, and lack microcnemes. I

lit Itis difficult to determine whether these three specimens belong to a known species. The description

of the type species, Halcampa endromitata Andres, 1881, is so brief as not to be diagnostic. The figure in m

the more informative redescription, in which Andres (1883) transferred the species to his new genus l_

HalcampelIa, resembles these specimens except that the physa of these is discoidal rather than rounded and

u

H. endromitata has 24 tentacles rather than 12, but that may simply be a function of size. The Mediterranean i

and presumably shallow habitat of H. endromitata also makes it unlikely to be the same as those from i

California. V

The specimens on which the other two species in the genus were based were large (to 150 mm long), ml

which could account for some differences between them and the three California specimens (if both of the

others actually do possess a capitulum and not a scapulus, they belong not only to a different species but to

another genus). The physa was torn from the sole specimen of H. robusta Carlgren, 1931, which could /

easily with the specimens from California. I

occur

Order Ceriantharia i

Tube anemones are exclusively infaunal, each animal forming im

a tube that is a feltwork composed mostly of ptychocysts (a type of I

cnida), mucus, and mud (Mariscal et aI., 1977). When disturbed, the

sensitive cerianthid rapidly retracts into itstube, whichtypically extends ill

above the surface of the sediment. The tube is unique to this group, I

but because it is often not collected, it cannot be relied upon to assign

a specimen to order. Related to the possession of tubes, cerianthids in

are exclusively solitary. A cerianthid has a long column that tapers to I

an end that is either narrow and rounded, or is bulbous (Fig. 3.10).

The most distinctive external attribute of a cerianthid, aside from its m

is its simple tentacles arrayed in two cycles: a whorl of short _mi

tube,

ones immediately surrounds the mouth, and longer ones are arrayed

at the margin (Fig. 3.10). Internally, a cerianthid has a single lira

siphonoglyph and its mesenteries, which are all complete, arise singly.

There is no marginal sphincter muscle.

Because of their subtidal, infaunal habitat, little is known about

the natural history or live appearance of animals of most species, and I

the polyps may be difficult to collect intact. This dearth of knowledge

contributes to taxonomic uncertainty: so few specimens of most l

species have been studied that range of intraspecific ontogenetic and •

geographic variability are unknown. Of about 50 species of cerianthids

n

recognized worldwide, perhaps four or five occur in the waters of

southern California. I

The number of specimens in this collection is uncertain

because some of the seven lots contain pieces (commonly the distal ill

end). Cerianthids may grow to more than 300 mm in length (Arai, I

1965), but the longest intact specimens in this collection were only Figure 3.10. Ceriantharian(after

25 ram. Character and arrangement of mesenteries are essential to Hyman,1940). I

104 '1
I

I family and genus placement; mesenteries of most of these specimens were insufficiently developed or wellpreserved for such determinations. None of the specimens in this collection were identified to species, but,

i • based a member on itsofshort familyproCerianthidae. tocnemes, oneAll frombutSone ta. R-of1the(Phase otherlI;specimens 91 m) wasinplthe acedcollection in the genappear us Pachycerianthus similar to that,

of Pachycerianthus; thus, all but that one may be at least congeneric, if not conspecific. Animals of the

I species as elsewhere; P.fimbr/atus the species are common was originally and variable described(Arai, from1971) Indonesia along th(Me cwest Murrich, coast o1910). f North America, as well

I Order Zoanthinaria i

m_ Zoanthids may be solitary, clonal, or colonial. Polyps are always(?) attached to f'krrnsubstrata,

I although, of course, in species with a thick coenenchyme, they arise from the common tissue that is fixed to

the substratum. Colonies of many deep-water taxa form symbioses with hermit crabs. Zoanthids typically

J inandcorporate coenenchyme, sand, sponge in somespicules, species tests sohicofklyforaminiferans, as to form essenortiother ally ansmall, exoskeleton. hard objects Although into their somebodyactinians wll

attach debris to their columns, in zoanthids such material is incorporated into the living tissue, the difference

i being shorterobvious capitulum; to andebris experienced is usuallyzoologist. thicker in theA zoanthid's former. Thecolumn short, issimple dividedtentacles into aarelonger arrayedscapus in a single and a

marginal cycle (Fig. 3.11).

i

!

I.

I

I

t

i

I

Figure 3.11. A colony ofzoanthids (afterI-IN, 1940). I

i

-I 105
!

!1

The mesenteries of zoanthids are paired and coupled; the typical pair is composed of one complete I!

and one incomplete mesentery. However, a pair of compete directive mesenteries is attached to the single

siphonoglyph; the other pair of directives is incomplete. The marginal sphincter muscle is endodermal or in

mesogleal, and single or double. tl

Perhaps 250 species are valid, although more have been described. Because of the debris in the

body wall, these animals are difficult to study. Nine species have been described from the northeast Pacific; I

only Epizoanthus induratum is in this collection. Its scapus is thickly encrusted by uniformly fine sand, and II

its coenenchyme is thin so that most of the length of each polyp is free of the common tissue.

l

Description of Species of Zo thinaria

I

Genus Epizoanthus Gray, 1867 I

Diagnosis. Epizoanthidae with single mesogleal sphincter. Scapus and coenenchyme are encrusted Ill

with foreign material. The arrangement of mesenteries is macrocnemic. This means that, of the mesenteries _!

flanking the incomplete directives, the member of the pair nearer the directives is complete whereas the other

member of the pair is incomplete, and both members of the pair of mesenteries beside that flanking pair are in

complete. In the rest of the pairs, except for the directives attached to the siphonoglyph, the member of the I

pair on the side toward the siphonoglyph is complete.

,Ul

Remarks. Nine species of this large genus are known from the eastern Pacific (Cutress and /

Pequegnat, 1960). II

Epizoanthus induratum Cutress and Pequegnat, 1960 i

Epizoanthus induratum Cutress and Pequegnat, 1960:95. I

I1

Material Examined. California: Santa Maria Basin, Phase I, Sta. 4 (x28); Phase I, Sta. 20 (x40).

Description. Column pillar-like, from 1- 9 mm length, 1.5 - 3.5 mm diameter; completely encrusted 1

with f'mesand grains of uniform size, including prominent ridges of scapulus. Aboral end of most specimens

examined tom -- presumably they had been embedded in a common coenenchyme. About 20 tentacles in

specimens examined. I

Macrocnemes about 20; microcnemes only in distal portion of column.

Cnidom: Spirocysts, microbasic p-mastigophores, microbasic b-mastigophores, holotrichs.

Biology. The only publication about this animal is the original description, which noted that large I

colonies occur "principally upon shale but may occasionally be found on the gorgonianMuricea californica"

and that it "is bioluminescent and pale salmon in color" (page 96). These observations could not be verified

in the material examined. II

Remarks. Holotype: colony of about 60 polyps from Corona del Mar, California, 1 mi (1.6 km) SE

of Newport Harbor entrance and 0.5 mi (0.8 km)off shore at 75 ft (23 m), encmsted on base and lower 1

portion of Muricea californica that was attached to shale, USNM 51054. Paratypes: colony of about 20 II

polyps from type locality, enerusting a piece of shale, USNM 51055; about 100 polyps constituting fragments

of a colony from 1.3 mi (2.1 km) SE of Corona del Mar, California, at 75 ft (23 m), encrusting a piece of I

shale, USNM 51058. II

Distribution. Southern California. Known from 23-396 m (this collection). 1

106 I
!

!

i Literature Cited

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i Carlgren, O. 1921. Actiniaria. I. The Danish Ingolf-Expedition 5(9):1-241.

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I Carlgren, O. 1942. Actiniaria. II. The Danish lrtgolf-Expedition 5(12):1-92.

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Cutress, C.E. 1949. The Oregon shore anemones (Anthozoa). Unpublished M. S. thesis, Oregon State

i Cutress, CoC.llege, and W.E. Corvallis Pequegnat. . 1960. Three new species of Zoantharia from California. Pacific Science

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I Dunn, D.E 1981. The clownfish sea anemones: Stichodactylidae (Coelenterata: Actiniaria) and other sea

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i Ebeling, 71(1):1-115. A.W., R.M. Ibara, R.J. Lavenberg, and F.J. Rohlf. 1970. Ecological groups of deep-sea animals off

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I 43.

Edwards, M.A. and A.T. Hopwood, eds. 1966. Nomenclator Zoologicus, vol. VI, 1946-1955. Zoological

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I Faurot, L. 1895. l_tudes sur l'anatomie, l'histologie ette drveloppement des actinies. Archives de Zoologic

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i Fautin, D.G., A. Bncklin, and C. Hand. 1990. Systematics of to Metridium

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(Coelenterata: Actiniaria), with a description of M. giganteum new species. Wasmann Journal of

Biology (for 1989) 47(1-2):77-85. !

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Garstang, W. 1892. On some new or rare marine animals recently discovered on the coast of Devonshire. HI

Reports and Transactions of the Devonshire Association for the Advancement of Science, Literature

and Art 24:377-386.

n

Gosse, P.H. 1853. Notes on some new or little-known marine animals XVI (No. 2). Annals and Magazine

of Natural History, series 2, 12:153-159. tll

Gosse, P.H. 1855. Description of Peachia hastata, a new genus and species of the class Zoophyta; with

observations on the familyActiniadae. Transactions of the Linnaean Society of London 21:267-276.

Gosse, P.H. 1858. Synopsis of the families, genera, and species of the British actiniae. Annals and Magazine U

of Natural History, series 3, 1:414-419. nq

Gosse, P.H. 1859. Characters and descriptions of some new British sea-anemones. Annals and Magazine I

of Natural History, series 3, 3(13):46-50. II

Gray, J.E. 1867. Notes on Zoanthinae, with the descriptions of some new genera. Proceedings of the

Zoological Society of London 1867:233-240. I

Grebel'nyi, S.D. 1980. The northern representatives of the genus Hormathia (Hormathiidae, Actiniaria).

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invertebrates, pp. 12-28. Zoological Institute, Academy of Sciences of the USSR (in Russian with II

English summary).

Hand, C. 1955. The sea anemones of central California part I. The corallimorpharian and athenarian I

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Hand, C. 1957. Another sea anemone from California and the types of certain Californian anemones, in[

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Hyman, L.H. 1940. The Invertebrates: Protozoa through Ctenophora. McGraw-Hill Book Co., New York lira

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Jourdan, E. 1880. Les zoanthaires du Golfe de Marseille. Armales des Sciences NatureUes, ser. 6, 10(1):1-

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Knowlton, N. 1993. Sibling species in the sea. Annual Review of Ecology and Systematics 24: 189-216.

Koehl, M.A.R. 1976. Mechanical design in sea anemones. Pp. 23-31 in:G.O. Mackie, ed., Coelenterate I

Ecology and Behavior. Plenum Press, New York and London. 41

Koehl, M.A.R. 1977. Mechanical diversity of connective tissue of the body wall of sea anemones. Journal

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McMurrich, J.P. 1910. Actiniaria of the Siboga Expedition, Part I. Ceriantharia. Siboga Expedition 10

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II McMurrich, J.P. 1913a. Description of a new species of actinian of the genus Edwardsiella from southern

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Manuel, R.L. 1981a. British Anthozoa (Synopses of the British Fauna, no. 18). Academic Press, London I

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Manuel, R.L. 1981b. On the identity of the sea anemone Scolanthus callimorphus Gosse, 1853 (Actiniafia:

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l Mariscal, R.N., E.J. Conklin, and C.H. 1977. The ptychocyst, a major new of cnida used Bigger. category

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I Walton, C.L. and O.M. Rees. 1913_ Some rare and interesting sea anemones from Plymouth. Journal of Marine Biological Association of the United Kingdom 10:60-69.

l Wassilieff, AbhAandlungen . 1908. Japanische der mathematisch-physikalischen Actinien. In Beitrfige Klasse zur Naturgeschichte der krniglich bayerisc Ostasiehns,en EAkademie Doflein, der ed.

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I Williams, R.B. 1981. A sea anemone, Edwardsia meridionalis sp. nov., from Antarctica and a preliminary revision of the genus Edwardsia de Quatrefages, 1841 (Coelenterata: Actiniaria). Records of the

Australian Museum 33(6):325-360.

I

I
IIiIIIIiiIII

,ilIIIi

I10I
Appendix

Lists and Maps of Stations

Table A.1. Position of soft-substrate stations taken during the Phase I Reconnaissance.

Station Latitude Longitude Depth

(m)

1 35°27.86"N 121°05.33"W 98

2 35°27.70'N 121°06.52'W 200

3 35°27.07'N 121° 10.20"W 291

4 35°26.56'N 121° 14.93'W 393

5 35°25.77"N 121°21.69'W 585

6 35°20.88"N 120°59.62"W 109

7 35°20.65"N 121°02.57"W 197

8 35°20.00"N 121°06.58"9/ 308

9 35°19.48'N 121°10.06"W 398

10 35°18.28'N I21°18.65"W 591

11 35°17.80'N 121°22.13"W 690

12 35°15.03"I'4 120°57.3 I'W 98

13 35°14.54'N 120°59.77"W 197

14 35°t4.15"N 121°02.04"W 299

15 35°13.98'N 121°04.54"W 393

16 35°12.23"N 121°16.29"W 591

17 35°11.61"1',,1 121°22.55"W 654

18 35°09.08'N 120°56.55"W 197

19 35°08.93'N 120°59.66"W 296

20 35°15.72'N 121°04.68'W 396

21 35°06.11 "N 120°44.82"W 49

22 35°05.85"N 120°50.23"W 99

23 35°05.60"N 120°55.18"W 195

25 35°05.07'N 121°00.75"W 390

26 35°04.38"N 121°15.99"W 590

27 35°04.30"N 121°19.27'W 611

28 35°04.22'N 121°19.65'W 603

30 34°54.19'N 120°47.07"W 98

31 34°53.76'N 120°52.96"W 200

32 34°53.56"N 120°56.8 I'W 297

33 34°53.43'N 120°59.66"W 396

34 34°53.15"N 121°04.40"W 492

35 34°52.96"N 121°10.30'W 548

36 34°52.77"N 121°15.37'W 492

38 34°49.8 I'N 120°52.66'W 197

39 34°49.53'N 120°56.85"W 294

40 34°49.24"N 121°00.8 I'W 392

41 34°48.35'N 121°19.14'W 495

42 34°48.04"N 120°47.50'W 100

43 34°46.59'N 120°5 2.92'W 197

45 34°44.9 I'N 120°59.59'W 395

46 34°41.22'N 121°13.56'W 597

47 34°41.99'N 121°10.81'W 378

48 34°45.1 I'N 120°52.85"W 196

49 34°45.03'N 120°56.31"W 290

50 34°37.80'N 121°01.66"W 591

52 34°39.56"N 120°47.64'W 98

53 34°37.69'N 120°50.38"W 196

54 34°36.57"N 120°52.02"W 396

55 34°33.66"N 120°56.3 I'W 590

56 34°30.32"N 121°01.02'W 900

111
!

Table A.1 (Continued)

dlB

Station Latitude Longitude Depth n

(m)

58 34°34.35"N 120°45.18'W 99 _l

59 34033.65"3I 120°47.18"W 216 II

60 34°33.25"5I 120°48.34'W 275

61 34°33.01"N i 20°48.89"W 345

62 34°30.46'N 120°52.13'W 582

63 34°26.29"N 120°58.08'W 930 U

64 34°33.15'N 120°40.90"W 59

65 34°31.27"N 120°43.27'W 107 n

66 34°30.46'N 120°¢4.55'W 201

67 34°30.29'N 120°45.50'W 282 I

68 34°29.24'N 120°45.99'W 390

69 34°22.88'N 120°54.20"W 927 m

70 34o29.67'N 120°43.70'W 200 !!

71 34°29.04'N 120°44.01'W 306

72 34°28.4 I'N 120°44.76"W 401

73 34°28.2 I'N 120°36.8(YW 98 R

74 34°26.84'N 120°38.6 I'W 201 II

75 34°26.08'N 120°39.65"W 293

76 34°25.59'N 120°40.98"W 387

77 34°22.62'N 120°44.02'W 578 !

78 34°18.78'N 120°49.3(YW 762

79 34°24.12'N 120°28.32'W 98

80 34°22.86'N 120°28.34'W 196

81 34°21.26'N 120°28.83'W 294

82 34°18.71'N 120°29.55'W 394 l

83 34°17.20'N 120°30.20"W 444

84 34°13.54'N 120°31.19"W 394 •

85 34°25.88"N 120°16.3 I'W 113 Ii

86 34°24.45'N 120°17.02"W 197

87 34°21.60'N 120°17.11"W 299

88 34017.89"N 120°16.86'W 393 B

89 34013.79'N 120°16.56'W 471

90 34°09.44'N 120°16.30'W 375

91 34°11.73'N 120°07.43'W 540

92 34°08.70"51 120°07.50'W 444 •

93 34°07.63"N 120°07.5 I'W 357 U

96 34°22.9 I'N 120°05.42'W 296

94 34°24.54'N 120°05.47'W 96 I

95 34°23.70"3I 120°05.47'W 198

97 34°22.28'N 120°05.49"W 393 a

98 34°12.8TN 120°05.59'W 561

99 34°11.22"N 120°05.86'W 540 Lm,

100 34°08.67"N 120°05.50'W 443

|

101 34°07.5 I'N 120°05.65'W 357

102 34°59.71'N 120°48.22'W 99

103 34°59.63"N 120°53.56'W 197

104 34°59.45'N 120°56.49"W 294 ti

105 34°59.23'N 120°59.60'W 392

106 34°58.95"N 121°04.42'W 492 m

107 34°58.65"N 121ot5.08"W 573 •

108 34°58.21"N 121°17.88'W 492

Note: Sample labels from the Soft-substrate stations have several identification codes which include a station number, sample /

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); 0t-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.

II

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Figure A.1. Map showing location of soft-substrate stations from the Phase I Reconnaissance and Phase H

i Monitoring Programs.

I 113
!

Table A.2. Location of soft-substrate stations taken during the Phase II Monitoring Program. I

Eli

Sration Latitude Longitude Depth •

(m)

R-1 35°05.83'N 120°49.16_ 91 I

R-2 35°05.5_N 120°53.40_ 161 g

R-3 35°05.3_N 121°_.90_ _9

R4 _°43.0 I'N 120°47.39_ _ I

l

R-5 _°42.69'N 120°50.83_ 154

R-6 M°41.4_N 120°57.90_ 410

R-7 M°52.9_N 121°10.30_ 565

R-8 M°55.3_N 120°45.87_ _ l

R-9 34°53.68'N 120°59.12_ 410 I

PJ-1 M°55.7_N 120°49.91_ 145

PJ-2 M°55.32'N 120°49.59'W t42

PJ-3 M°56.26'N 120°49.58'W 138

PJ-4 M°56.26'N 120°50._'W 150 g

P J-5 _°55.32'N 120°50._'W I52

PJ-6 _°54.71"N 1_°49.91_ 148 m

I

P J-7 M°55.7_N 120°48.60_ 123

P J-8 _°56.87'N 120°49.9t_ 142

PJ-9 M°55.7_N 120°51.23_ ! 69

PJ- 10 _°53.63'N 120°49.91 _ 147 m

|

PJ- 11 M°57.95'N 120°49.91_ 136

PJ- 12 _°55.58'N 120°49.91_ 145

PJ-13 _°56.01'N 120°49.91_ 1_

PJ- 14 _°55.7_N 120°49.26_ 134

|

P J-15 M°55.7_N 120°50.57_ 155

PJ- 16 _°55.03'N 120°48.99_ 130

PJ- 17 M°56.56_ 120°48.98_ 126 /

PJ- 18 M°56.56'N 120°50._ 158 •

PJ- 19 M°55.03'N 120°50.M_ 167

PJ-20 M°50.38'N 120°49.91_ 148

PJ-21 35°01.23_ 120°51.15_ 143 I

|

PJ-22 M°55.25'N 120°49.93_ 143

PJ-23 _°56.33'N 120°49.90_ 143

I

|

Table A.3. Sampling dates of MMS Phase II Monitoring Program. I

Cruise Date .l'

1-1 October 1986

1-2 Jan_ 1987

1-3 May 1987

2-1 J_y 1987 [

2-3 Oc_r 1987

24 Jan_ 1988

2-5 May 1988

3-1 Oc_r 1988

34 May 1989 I

..|

i 14 1
Table A.4. MMS Phase I - Locations of hard-substrate transects.

Station Beginning End Depth

Latitude Longitude Latitude Longitude (m)

1 A/B 34°24.454'N 120°01.876'W 34024.464'N 120°00.878'W 69-73.5

1 C/D 34°24.076'N 120°00.443'W 34°24.184'N 120°01.480"W 73.5-78

2A/B 34°i 1.377'N 120°29.318'W 34°11.289"N 120°28.774'W 110-126

2 C/D 34°10.984"3/ 120°28.094'W 34° 10.78fin 120027.554"W 120-123

4 A/B 34°27.539'N 120°40.364"W 34°28.162'N 120°40.189'W 168-237

6 A/B 34°30.246'N 120%5.555'W 54-63

6 C/D 34°30'.42 I'N 120°3'4.315'W 54-63

13 A/B 34°42".570"N 120°47.899"W 34°42.107'N 120°48.253'W 92-100

13 C/D 34°42.556'N 120°48.147"W 34°42.974'N 120°47.424'W 88.5-100.5

14 A/B 34°43.589'N 120°49.093"W 34°42.826'N 120°48.370'W 96-105

14 C/D 34°43.244"N 120°49.406"W 34°42.893"N 120°48.822'W 105-117

16 A/B 34°46.544"N 120°50.197'W 34°45.912"N 120°49.726'W 91.5-123

17 A/B 34°49.382'N 120°50.768"W 34°49.600'N 120°50.688'W 160.5-168

19 A/B 34°47.833"N 120°51.425"W 34°47.097"N 120°50.793'W 148.5-177

20 A/B 34°46.470'N 120_50.289"W 34°46.140'N 120°49.885'W 90-130.5

21 A/B 34°47.335"N 120°45.903"W 34°47.548"N 120°46.123'W 75-90

22 A/B 34°50.365"N 120°48.221"W 34°50.990"N 120°48.365'W 114-115.5

23 A/B 34°49.868"31 120°47.393'W 34°50.003'N 120°47.480'W 93-102

25 A/B 35°05.662'N 120°47.562'W 35°06.036'N 120°47.652"W 64.5-72

26 C/D 35° 11.586"N 120°55.556'W 35°11.555"N 120°55.233'W 108-111

27 A/B 35°20.906'N 120°59.657'W 35°21.035'N 120°59.603"W 96-126

28 A/B 35°21.539'N 120°59.641'W 35°21.867'N 120°59.299"W 96-105

29 A/B 35°27.864"N 121°05.331"W 35°27.805"N 121°05.277"W 102-106.5

'l_le A,$. MMS Phase II - I.,oeafions of hard-substrate photosm'vey stations.

Stufion Latitude Longitude Depth

(m)

PH-E 34°30.26"N 120°42.76"W 119

PH-F 34°30.81'N 120°42.36'W 105

PH-I 34029.96'N 120°41.68'W 107

PH-J 34°29.82'N 120°41.82"W 117

PH-K 34029.37"N 120°42.26"W 160

PH-N 34029.21 "N 120°42.05"W 166

PH-R 34o29.11 'N 120°42,67"W 213

PH-U 34°31.4 8'N 120°43.51 "W 113

PH-W 34°31.52"N 120°45.86'W 195

115
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Figure A.2. Map showing location of hard-substrate stations from the Phase I Reconnaissance and Phase II

Monitoring Programs. Phase II stations are indicated in the inseL I

i

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I

The Department of the Interior Mission

I As the Nation's principal conservation agency, the Department of the Interior has responsibility

for most of our nationally owned public lands and natural resources. This includes fostering

sound use of our land and water resources; protecting our fish, wildlife, and biological diversity;

preserving the environmental and cultural values of our national parks and historical places;

P and providing for the enjoyment of life through outdoor recreation. The Department assesses

our energy and mineral resources and works to ensure that their development is in the best

interests of all our people by encouraging stewardship and citizen participation in their care.

i The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.

The Minerals Management Service Mission

t _____% As a bureau of the Department of the Interior, the Minerals Management Service's (MMS)

primary responsibilitiesare to manage the mineral resources located on the Nation's Outer

Continental Shelf (OCS), collect revenue from the Federal OCS and onshore Federal and Indian

I lands, and distribute those revenues. Moreover, in working to meet its responsibilities, the Offshore Minerals Management Program

administers the OCS competitive leasing program and oversees the safe and environmentally

i sound exploration and production of our Nation's offshore natural gas, oil and other mineral resources. The MMS Royalty Management Program meets its responsibilities by ensuring the

efficient, timely and accurate collection and disbursement of revenue from mineral leasing and

production due to Indian tribes and allottees, States and the U.S. Treasury.

i The responsive MMS sttorivthe es topublic's fulfill itconcerns s responsibiliti and inter es through ests by the maintaining general guiding a dialogueprinciples with allof:potentially (1) being

affected parties and (2) carrying out its programs with an emphasis on working to enhance the

quality of life for all Americans by lending MMS assistance and expertise to economic

i development and environmental protection.