DOE/EVIOO703-T1 -Vol . 2

(DE870061 1 1 )

The Natu re H sto ry of

Enewetok Ato

Vo lume II B iogeog raphy and Sys temat ics

Edited by : Denn i s M . DevaneyBer n ice P . Bi s hop Mu s eum

Hono lu lu , Hawai i

Un ited States

Depar tment Of Energy Er n s t S. Rees e

Un iver s ity of Hawai iOf f ice of Energy Res earchHono lu lu , Hawai i

Of f ice of Health and

Envi ronmen tal Res earch Beat r ice L . Bu rch

Ber n ice P . B i s hop Mu s eum

Eco log ical Research D ivi s ion Hono lu lu ,Hawai i

Prepared by Ph i l ip H el f r ich

Off ice of Sc ient if ic and Techn ical Information Un iver s i ty of Hawai i

US Department of Energy Hono lu lu , Hawai i

[Photograph by Annabelle Lyman .)

a w l"

ln temat ional Copyr ight , ! U . S. Depar tment of Energy , 1 987, under the provi s ions of the Un iver salCopyr ight Convention . Un ited States copyr ight i s not asser ted under the Uni ted States Copyr ightLaw , T i tle 1 7, Uni ted States Code.

L i b r ar y o f Con g r es s Catal og i n g-l n -P n b l i ca t l on D ata

The natur al history of Enewetak Atol l .

DOE/EVOO703-T I -Vol . l (DE87006 1 1 0) .

DOE/EVOO703-T I -Vol . l l (DE87006 1 l

Includes indexes .

1 . Natural history— Mar shall i s lands — EnewetakAtol l . l . Devaney , Denn is M . II . Un ited States .

Dept . of Energy . Office of Scientific and Techn icalInformation .

QH 1 98 .E53N38 1987 87-24863

ISBN 0—87079—579- 1 (set)

lSBN 0-87079-580-5 lpbk. set )

ISBN 0-87079-581 -3 (m icr ofiche set )

Work performed under contr act No . DE-ACO8-76EV00703

The set of two vo lumes is avai lable as DE870061 1 0 (DOE/EV/0073-T I -Vol . 1 ) and DE87006 1 1 1

(DOE/EV/0073-T l -Vol .2) fr omNTlS Energy D istr ibution CenterP . 0 . Box 1 300

Oak R idge, Tennessee 3783 1

Pr ice Code: Paper Copy A99Micr ofiche AO1

Pr i nted in the Un ited States of Amer ica

F o r eword

As act iv i ty and fund ing at the M id-Pac i fic Resear ch Laborator y began to d im in ish in the ear ly 1 9805 ,i t seemed fi tt ing that a s ynthes is be pr epared of the thr ee decades of r esear ch that had been conducted atth is Laborator y on Enewetak Ato l l . For 30 year s the Atol l ser ved as a conven ien t , access ible locat ion fors tud ies of M id-Paci fic i s land ecosys tems , and several hundr ed sc ien tis ts ut i l i zed the faci l i ty . Pr imar y funding was pr ov ided by the Off ice of H ealth and Env i r onmental Resear ch , Eco logical Resear ch D iv is ion ,

U . S. Depar tment of Energy (former ly the Atom ic Ener gy Commi ss ion and the Energy Resear ch and

Development Adm in i strat ion) .Th is i s an attempt to syn thes i ze in two vo lumes the results of the M id-Paci fic Resear ch Laborator y

s tu dies that have been publ ished in hundreds of w idely di sper sed publications . It is hoped that presen t andfuture scien tis ts invo lved in s tudies of M id-Paci fic i s lands w i l l find thi s syn thes is a conven ien t resour ce forthei r resear ch .

Cons iderable time and effor t were expended by many con tr ibutor s to make th is syn thes is poss ible.

Thanks are extended to all these author s for thei r manuscr ipts . Special appreciation i s expressed forDr . Denn is Devaney ’s dedi cation in fi ll ing gaps in the taxonom ic descr ipt ions of several inver tebrategroups . Thi s publ ication would not have been poss ible, however , wi thout the determ ination and per s istence of Dr . Er ns t Reese in organ i z ing and col lect ing the mater ial . Deepes t grati tude is acknowledged forhis conscientious effor ts .

H elen M . M cCammon, D i rectorEco logi cal Resear ch D iv is ionOffice of H ealth and Env i r onmental Resear chUni ted States Depar tment of Energy

Acknowledgmen ts

Many people have contr ibuted in many ways to the production of these two volumes . Regar dless of

the nature of the contr ibut ion, everyone listed below has given thought and time, that mos t pr ecious comr

‘

nodi ty of thinking individuals, to br ing The Natural H istory of Enewetak Atoll into publication . Author s ofthe chapter s are not l is ted separately , even though , in most cases , they cr i ti cally read other chapter s . No

doubt we have over looked many who have contr ibuted In Impor tant ways , and for these over s ights weapolog i ze. To all of you we w ish to extend our deepest thanks .

Donald P . Abbottlsabella A. AbbottHazel K . AsherGeorge H . BalazsJer r y L . Bar nard

Freder ick M . BayerHenry R . BennettRichard A . Booloot ian

Thomas E. Bowman

Har ry U . Brown

Fenner A . Chase, Jr .

G . Ar thur Cooper

The two volumes of The Natur al H is tory of EnewetakAtoll summar i ze resear ch done at the M id-Paci fi c Resear chLaborator y from 1 954 to 1984 under the auspices of the

Depar tment of Energy . The h is tor y of the labor ator y andthe reasons for i ts suppor t by the Un i ted States Depar tment of Energy ar e descr i bed in Chapter 1 of Volume 1 .

Over a thousand per sons —es tablished scientis ts , thei r

ass is tants , and gr aduate s tudents — conducted resear ch atthe labor ator y dur i ng the 30-year per iod. Thei r effor tsresulted in 223 publ i cations . These have been col lected infour volumes of repr ints ent i tled M id-Pac ific Mar ine Labor atory Contr ibutions, 1 955— 1 979, U . S. Depar tment of

Energy, Publication NVO 628-1 . The laboratory has con

tinued oper ation on a limi ted scale to the pr esen t . A col

lection of paper s recent ly appear ed in the Bulletin ofMar ine Science, Vo lume 38 , 1986 .

Much of the r esear ch conducted at the labor ator y wason the mar ine env i ronmen t . The reason was that themajor i ty of Scient is ts applying to wor k at Enewetak weremar ine bio logis ts . For many , th is was the fi r st oppor tun i tyto study the biota of a coral atol l . Fewer s tudies were con

ducted in the ter r es tr ial env i r onment and i ts biota.

Never theless , as these volumes attes t, the coverage is

amaz ingly complete and thorough , and there ar e few , i f

any, studies of an equ ivalent ecosystem that equal thetotal resear ch effor t r epor ted in these volumes .

Volume i prov ides a syn thes is of the resear ch car r iedcut under the subject headings of the respective chapter s .

Cer tain of the chapter s , e.g . , those on geo logy , subtidaland inter t idal env i r onmen ts and ecology , and those on reefprocesses and trophic r elationsh ips , summar i ze a greatdiver s i ty of resear ch car r ied ou t by many sc ien ti s ts formany year s . in contras t , the chapter s on meteoro logy andoceanog r aphy summar ize r esear ch car r ied out under one

integrated program invo lving fewer scien tis ts wor k ing over

a shor ter per iod .

Volume 11 of The Natural H is tory of Enewetak A tollpr ov ides in format ion on the taxonomy of an imals and

plan ts known to occur at Enewetak Atoll . Th is taxonomyr epr esents a fulfi l lment of one of the fi r s t ass ignments to

the laborator y— to determine the scien ti fi c names of the

biota of the atol l . The col lections on wh ich the check lis tsin each chapter are based are housed at the Ber n ice P .

B ishop Museum in H onolulu and the U . 5 . NationalMuseum of Natural H is tor y , Smi thson ian ins ti tution , Washington , D . C.

in addi tion to the species check lis ts , each chapter inVo lume i i prov ides a succinct summar y of the biota wi thr espect to endemism , range extens ions , and other featuresthat set the Enewetak biota apar t from those one mightexpect to fi nd on equivalent lndo-Paci fi c is lands . Th is compendium of taxonomic information for an ato l l shouldprove of immense value to s cientis ts interes ted in biogeography and evolut ionar y bio logy of is land ecosys tems foryear s to come.

One of the problems of edi ting these volumes has beenthe cor rect use of place names . in some cases author sused the m i l i tar y code names for is lands wh i le other s usedthe native names . Even the native names have changedfr om ear ly phonetic spell ings to the spell ings cur r ently inuse and pr efer r ed by the Enewetak people. For example,

the name of the atol l has changed fr om En iwetok toEnewetak, and, although the cor r ect cur r ent spell ing is

used throughout , the old spel ling occur s in o lder r efer encesand maps wh ich appear in these volumes . Maps giv ing themi l i tary code names and the native names prefer r ed by theEnewetak people are located in Chapter 1 of Vo lume 1.Sur pr i s ing ly , i t is di fficult to determine the exact number ofi s lands . Due to the effects of s torms , small is lands are

ephemer al , and two i s lands and par t of a th i r d were ob

li terated by nuclear explos ions . Cur r ently there are 39 rec

ogn izable is lands , and these are shown on the map usedthroughout the book .These volumes do not r eport on the extens ive radiolog

ical sur veys and s tud ies wh ich have been conducted by theLawr ence L ivermore Labo ratory , Un iver s i ty of Cal i for n ia,

and the Rad iat ion Laborator y , Un iver s i ty of Wash ington ,

also under the auspices of the U . S. Depar tmen t of

Ener gy .Denn is M . Devaney , sen ior edi tor of th is vo lume, disap

peared wh i le col lect ing specimens off the i s land of H awai ion Augus t 1 3 , 1 983 . Denn is was do ing what he lovedbes t , col lect ing mar ine inver tebrates , at the t ime of his

death . H e col lected extens ively at Enewetak , and be undertook the task of organ i z ing the sys tematic chapter s of

Volume i l . Beatr ice L . Bur ch , Devaney ’s ass is tant at the

B i shop Museum , completed the task , and she has wr i ttenthe introduction to Vo lume i i .

i t i s fi tt ing that the two volumes of th is book are dedicated to the people of Enewetak Atoll . They , like so manyother human beings , were caught up by for ces beyondthei r con tr ol and under s tanding in an immense cataclysm

of human h is tor y . in a smal l way, this book s tands as

someth ing good that has res ul ted fr om those year s .

E r ns t 8 . Reese

Pr ofessor of Zoo logyUn iver s i ty of H awai i , H onolu lu

Con t r ibu tors

A . Char les Ar nesonScr ipps ins t i tut ion of Oceanogr aphy , La Jo l la, Cal i for n iaJul ie H . Bai ley-B rockU n iver s i ty of H awai i , H ono lulu , H awai i

Andr ew J . Ber gerU n iver s i ty of H awai i , H ono lu lu , H awai i

A . J . B r uceNor ther n Ter r i tor y Museum of Ar ts and SciencesDarw in , N . T . , Aus tr al ia

Beatr ice L . Bur chB . P . B ishop Museum ,

H ono lulu , H awai iEl i zabeth H . ChaveU n iver s i ty of H awai i , H ono lulu , H awai i

C . A l lan Ch i ldSm i thson ian ins t i tut ion , Wash ington , D . C .

Rober t 8 . Cox

U n iver s i ty of M ich igan ,Ann Ar bo r , M ich igan

Roger J . CuffeyPenns ylvan ia State U n iver s i ty , Un iver s i ty Par k , Pennsylvan ia

Ber tha M . Cu t res s

U n iver s i ty of Puer to R ico ,Mayag ii ez , Puer to R ico

Char les E . Cu t r ess

U n iver s i ty of Puer to R ico ,Mayaguez , Puer to R ico

Denn i s M . Devaney (Deceased)B .

‘

P . B i s hop Museum , H ono lulu , H awai iPaul H . DunnR iver s ide F i r e Labor ator y , R iver s ide, Cal i for n ia

L ucius G . Eldr edgeU n iver s i ty of Guam , Mang i lao ,

GuamJohn S. Gar thUn iver s i ty of Souther n Cal i for n ia, Los Angeles , Cal i for n ia

Ray P . Ger berSt . Joseph ’

s Co l lege, Nor th W indham ,Maine

R ichar d E . Gr antSm i thson ian i ns t i tut ion , Wash ington , D . C .

Janet H aigUn iver s i ty of Souther n Cal i for n ia,

Los Angeles , Cal i for n ia

Er ns t S . Reese

U n iver s i ty of H awai i at Manoa, H ono lu lu , H awai iDon Reyno ldsNatural H i s tor y Museum, Los Angeles , Cal i for n iaFr anci s W . E . RoweThe Aus tr al ian Museum, Sydney , Aus tral iaG . A llan Sam uelsonB . P . B ishop Museum , H ono lulu , H awai i

R ichar d H . T itgen

B . P . B i s hop Museum , H ono lulu , H awai iRoy T . TsudaU n iver s i ty of Guam , Mang i lao ,

Guam

Scott JohnsonU n iver s i ty of H awai i at Manoa, H ono lulu , H awai i

E . A l i son KayU n iver s i ty of H awai i at Manoa, H ono lulu , H awai i

Lou is S . Kor n icker

Sm i thson ian ins t i tut ion , Wash ington , D . C .

Jens W . KnudsenPaci fic L utheran U n iver s i ty , Tacoma,

Wash ingtonJanet O . Lamber sonU . S. Env i r onmen tal Protect ion Agency ,Newpor t , O r egonJud i th C . Lang

U n iver s i ty of Texas at Aus t in , Aus tin , TexasRaymond B . Mann ingSm i thson ian ins t i tut ion , Wash ington ,

D . C .

Go r don M . N ish idaB . P . B i shop Museum

,H ono lu lu , H awai i

H elen A . Randal lB . P . B i shop Museum , H ono lulu , H awai iJohn E . Randall

B . P . B ishop Museum , H onolulu , H awai iMar jor ie L . BeakaU n iver s i ty of Mar yland , Co l lege Par k , Mar yland

Con ten ts

Chapterin troduct ion

Beatr ice L . Bur chMar ine Benth ic A lgae of Enewetak Ato l l

Roy T . TsudaFungi of Enewetak Atol l

Pau l H . Dunn and Don Reyno ldsNatur al H is tor y of Ter r es tr ial Vascular Plan ts of Enewetak Ato ll

Janet O . Lamber son

Recent Foram in i fera and Nonp lankton ic Pr otozoansE . H . Chave and Denn is M . Devaney

Po r i fera of Enewetak Ato l lDenn is M . Devaney

Sea Anemones of Enewetak Ato llC . E . Cut ress and A . C . A r neson

Octocorall ia of Enewetak Atol lDenn is M . Devaney

Scler act in ia (Ston y Corals) of Enewetak Atol lDen n is M . Devaney and Judith C . Lang

Br ach iopods of Enewetak Ato l lR ichard E . Gr an t

Reef-Dwel l ing B r yozoans of Enewetak Ato l lRoger J . Cuffey and Rober t 8 . Cox

Sipuncu lans and Echiu r ans of Enewetak Atol lDenn is M . Devaney

Platyhelm in thes , Nemer tea, and Nematoda of Enewetak Atol lDenn is M . Devaney

Po lychaetes of Enewetak Atol lDenn is M . Devaney and Ju lie H . Bai ley -B rock

Mo l lusca of Enewetak Atol lE . A lis on Kay and Scott John son

insects and A l l ies (Ar thr opoda) of Enewetak Ato l lG . A l len Samue lson and Gordon M . N is h ida

Pycnogon ida of Enewetak Atol lC . A l lan Ch i ld

Stomatopod Cr us tacea of Enewetak Ato l lMarjor ie L . Reaka and Raymond B . Man n ing

Ci r r iped ia of Enewetak Ato l lR ichar d H .Titgen

ChapterNonplankton ic Copepoda of Enewetak Atol l

Denn is M . DevaneyLagoon Plankton of Enewetak Atol l

Ray P . Ger berOs tr acoda (Myodocop ina) of Enewetak Ato ll

Louis 8 . Kor n icker

Cr us tacea Decapoda (Penaeidea, Stenopodidea, Car idea, and

Palinu ra) of Enewetak Atol lDenn is M . Devaney and A . J. B ruce

Cr us tacea Decapoda (B r achyur a and Anomura) of Enewetak Atol lJohn S. Gar th, Janet H aig, and Jens W. Knudsen

H olothur ians of Enewetak Atol lBer tha M . Cu t ress and Fr ancis W. E . Rowe

Ech inodermata Other than H o lothur ians of Enewetak Atol lDenn is M . Devaney

Protochor dates of Enewetak Ato l lLucius G . Eldredge

Annotated Check l ist of the Fishes of Enewetak Ato ll andOther Mar shal l i s lands

John E . Randall and H elen A . Randa llRept i les of Enewetak Atol l

Janet O . Lamber sonB i r ds of Enewetak Atol l

Andr ew J. Berger

Mammals of Enewetak Ato l lEr ns t 8 . Reese

M iscel laneous Spec ies Recor ds of Enewetak Atol lR ichard H . Ti t

“

en and Beatr ice L . Bur ch

In troduction

Beatr ice L . Bur ch

B . P . B ishop Museum ,H ono lu lu, H awa i i 968 1 7

The organ i zat ion and coor dination of the taxonom icsect ion of th is vo lume was in i t iated by Dr . Denn is M .

Devaney of the Bernice P . B ishop Museum and was com

pleted by Beatr ice L . Bur ch after Dr . Devaney died in a

tr agic scuba-diving acc ident on Augus t 13 , 1 983 , as he

was inves tigating shr imp offshore from the B ig i s land of

H awai i . H is great interes t in all inver tebrates in the tropicswas increased when the oppor tun i ty ar ose for him to wor kat the Mid-Paci fic Resear ch Laborator y at Enewetak Atol l .Devaney made hi s fi r s t col lect ing tr ip to the atol l in the

ear ly 1 9605 wh i le he was s t i l l in graduate s choo l . As thecol lection grew and taxonom y of the organ isms becamebetter known , i t was soon apparent that the refer ence col

lection at Enewetak was becom ing increas ingly valuable.

Devaney was p leased to par t ic ipate in the Cor al ReefWor kshop held at the atol l in 1 976 , because he bel ievedthat the scler act in ian coral co l lect ion was the key for the

s tudy of other organ isms . The wor kshop br ought togetherinternat ional coral special is ts to es tabl i sh species l imi ts on

th is impor tant and var iable g roup . After the wor kshop washeld , reference mater ial fr om Enewetak was depos i ted inEuropean and Amer ican museums for r eady r efer ence by aw ider audience of scient is ts . Each year after the Cor alReef Wor kshop , Devaney wen t to Enewetak to cur ate thereference col lect ion and to conduct his own r esear ch on

ech inoderms . At the same t ime , he encour aged the wor kof special i s ts to comp i le taxonom ic and other resear chfrom Enewetak for this publ icat ion .

The d iver s i ty of the or gan i sms at Enewetak made i t

d i fficult to find spec ial is ts to s tudy all groups , so Devaneypr epared several chapter s h imself . Un for tunately , mos tgroups wer e co l lected in the cour se of other wor k such asphys iology , tox ici ty , etc . , and wer e not extens ively col

lected by special is ts for a par t icular taxonom ic g roup .

The number of fam i l ies , gener a, and spec ies r epor tedin th is vo lume ei ther fr om the l i ter atur e o r fr om new

r ecords determ ined by the author s of th is vo lume ar e

pr esented in Table 1 .

Refer ences in th is vo lume show that some or muchwor k was done on a par t icular taxon . Man y gr oups r emain

Taxonom i c Gr oups at Enewetak A to l l

Taxon

Totals

TABLE 1

N o . of

species

4671

N o. of

genera

2284

N o. of

fam i l ies

AlgaeFungiVascular plantsForams and nonplankton ic

protozoaPor i feraAct in iar ia

.

Octocoral l iaScleract in ia

B rach iopodaB r yozoaSipunculaEchiur a

P latyhelm inthesNemer teaNematodaPo lychaetaMo l lusca (fos s i l , r ecent)insects and r elated

ar thr opodsPycnogon idaStomatopodaCi r r ipediaLagoon planktonOs tracodaNatantiaReptant iaAnomuraB rachyuraH o lothuro ideaEchinodermata other thanH o lothuro ideaFishes of the Mar shal li s lands

Rept i l iaAves

Mammal iaM i scel laneous

to be wor ked on more thoroughly by special is ts in par t icular fields , such as Por i fer a or Tun i cata, wh ich , at the

presen t t ime, seem to be r epr esented so l ightly at

Enewetak Ato l l . By hav ing a named r efer ence co l lect ion ,

the r esear cher s ther e wer e able to iden t i fy or gan i sms usedin thei r s tud ies on biochem is tr y , eco log y , product iv i ty ,an imal o r plant assoc iat ions , phys io logy , immuno logy ,rad iobio logy , growth r ates , and r epr oduct ion . They wer ealso able to make broad in ter pretat ions of r eef chrono logy ,

geochem is tr y , s tr at igr aph y , and biogeog raph ic di s tr ibut ion .

The checkl i s t contained in each chapter has a codedentr y s ymbo l placed befor e the gener ic des ignat ion to indicate ( 1 ) i f the organ i sm r epresen ts a newly r ecor dedspecies fo r Enewetak or for the Mar shal l i s lands , (2) i f i t isa foss i l recor d , or (3 ) i f i t has some other r eason to be so

mar ked . The explanat ions for these codes fo l low eachspec ies checkli s t .

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TSUDA

wer e heter-ocys tous blue-

gr eens s tud ied by W iebe et ai . Enewetak Ato l l : Cyanophyta ( 1 6 species ), Chlor ophyta (89Webb and W iebe Webb et ai . and species ) , Phaeoph yta (24 spec ies ) , and Rhodophyta (1 09

W iebe (1 976) in thei r s tud ies on the n i trogen cycle. spec ies ) . N ine of the species (pr eceded by a dagger inOn the bas is of the paper s c i ted above, 238 spec ies Table 1 ) wer e descr ibed as new w i th the Enewetak speci

( 1 06 genera) of mar ine benth ic algae are known fr om mens ser v ing as ho lotypes . i t is in ter es t ing to note that 40

TABLE 1

Check l i s t of Mar i ne Ben th i c A lgae of Enewetak

D iv i s ion CY'ANOPH YTA (blue-

g r een algae)Class CYANOPH YCEAEOrder CH ROOCOCCALESFam i ly CH AMAESiPH ONACEAEEn tophys al is confer ta (K i i tz ) D r . and Dai ly : Dawson , 1 957 .

En tophysal is deus to (Menegh ) D r . and Dai ly : D r ouet and Dai ly , 1 956 ; Dawson , 1957 .

Fam i ly CH ROOCOCCACEAEAnacys t is dim idiata (Kii tz ) D r . and Dai ly : Dawson , 1957 .

Coccochlor is s tagn ina Sprengel : Dawson , 1 957 .

G omphosphaer ia apon ina Kii tz . : Taylo r , 1 950b; D r ouet and Dai ly , 1 956 ; Dawson , 1 957 .

Order OSCILLATORIALESFam i ly NOSTOCACEAECalothr ix cr us tacea Schousboe and Thuret : Dawson , 1 957 ; Bakus , 1 967; Dr ouet , 1 973 ; Webb et ai . , 1 975 ;Webb and W iebe , 1 975 ; W iebe et ai . , 1 975 ; W iebe , 1 976 .

R ivu lar ia at r a Roth : Dawson , 1 957 .

R ivu lar ia po lyo t is (J . Ag . ) B . and FL: Dawson , 1 957 .

H ormotham n ion en ter omorphoides B . and Fl . : Tay lor , 1 950b; Dawson , 1957; Gi lmar t in , 1960 ; Webb et ai . , 1 975 ;W iebe et ai . , 1975 .

Scytonema hofman i i Ag .

Scytonema myocyr ous (D i llw . ) C. Ag . : Dawson , 1 957 .

Scytonema polycys tum B . and F i . z Dawson , 1 957 ; Gi lmar t in , 1 960 .

Fam i ly OSCILLATORlACEAEM icr ocoleus lyngbyaceus (Kli tz ) Cr ouan .

H ydr ocoleum can thar idosmum (Mont ) Gomon t : Tay lor , 1 950b .

H ydr ocoleum coccineum Gomon t : Dawson , 1 957 .

H ydr ocoleum como ides (H ar v ) Gomon t : Dawson , 1957; Gi lmar t in , 1 960 .

H ydrocoleum g lut inosum (C. Ag . ) Gomont : Dawson , 1 957 ; G i lmar t in , 1 960 .

H ydr ocoleum lyngbyaceum Kii t z Dawson ,1 957 .

Lyngbya aes tuar i i (Mer t . ) Lyngb. : Dawson , 1 957 .

Lyngbya confer vo ides C. Ag . z Tay lor , 1 950b; Dawson , 1 957

Lyngbya majuscu la (D i llw . ) H ar v : Dawson , 1 957 ; Gi lmar t in , 1 960 .

Lyngbya meneghin iana (Kli tz ) Gomont : Taylor , 1 950b; Dawson , 1 957

Lyngbya sem ip lena (Ag . ) J . Ag . z Dawson , 1 957 .

Osci l lator ia submembranacea A rd. and Straff : Dr ouet , 1 968 .

Phorm idium pen ici llatum Gomont : Dawson , 1 957 .

Porphyros iphon no tar is i i (Menegh ) Ku'tz .

Osci llator ia n igr o-vi r idis Thw . : Dawson , 1 957 .

Sch iz othr ix ar enar ia (Berkeley) Gomont .Symp loca laete-vi r idis Gomont : Dawson , 1 957 ; Gi lmar t in , 1 960 .

Schiz othr ix calcicola (C . Ag . ) Gomont : Bakus , 1 967 ; D rouet , 1 968 ; Johannes et ai . , 1972 ; Webb and W iebe , 1 975 .

Phorm idium crosbyanum T i lden : Tay lor , 1 950b; Dawson , 1 957 .

Plectonema nos tocor um Bo r net : Tay lor , 1 950b ; Dawson, 1 957Plectonema ter ebr an s B . and F i . z Dawson , 1 957 .

Sch iz othr ix lacus t r is A . B r . : Dawson , 1 957 .

Schiz othr ix mex icana Gomont : Drouet , 1 968 .

Lyngbya gr aci lis (Menegh ) Rabenh . : Tay lo r , 1 950b ; Dawson , 1 957

Lyngbya sordida (Zanard ) Gomont : Dawson , 1 957 ; Gi lmar t in , 1 960 .

Symp loca hydno ides (H ar v ) K i i tz : Dawson , 1 957 ; G i lmar t in , 1 960 .

Schiz o thr ix tener r ima (Gomont) D r . : D rouet , 1 968 .

M icrocoleus tener r im u s Gomont : Dawson , 1957 .

Sp i ru l ina s ubsalsa Oer sted : Dawson , 1 957 .

Sp i ru l ina majo r K i l tz . : Dawson , 1 957 .

Sp i ru l ina tener r im a K ii tz . : Dawson , 1 957 . (T hi s table con ti nued on next page. )

MARINE BENTH IC ALGAE

TABLE 1 (con t'd)

D iv is ion CH LOROPH YTA (green algae)Class CH LOROPH YCEAEOrder CH AETOPHORALESFam i ly CH AETOPH ORACEAE

‘En tocladia vi r idis Reinke: Dawson , 1 957 .

Order ULVALESFam i ly ULVACEAE

‘En ter omo rpha acan thophora Kutz . : Dawson , 1 957 .

En teromorpha clathrata (Roth) J . Ag . : Dawson , 1 957 .

En ter omorpha in tes t inal is (L . ) L ink : Dawson , 1 957 .

En teromorpha kyl in i i B l iding : Dawson , 1 957 .

En teromorpha l ingu lata J . Ag . z Tay lo r , 1950b .

En ter omorpha ralfs i i H ar vey : Dawson , 1 957 .

‘En ter om orpha tubu losa (K ijtz ) Kil tz . : Dawson , 1 957 .

Order CAULERPALESFam i ly BRYOPSlDACEAEB ryop s is hypno ides Lamx Dawson , 1 957 .

B ryop s is indica A . and E . S. Gepp : Tay lor , 1 950b ; Dawson , 1 957 .

B ryop s is pennata Lamx . : Dawson , 1 957 ; Gi lmar t in , 1 960 .

Fam i ly CAULERPACEAECau lerpa ambigua OkamuraCaulerpa vicker s iae Boerg . : Dawson , 1 957; Gi lmar t in , 1 960 .

Cau lerpa an toens is Yamada : Dawson , 1 957 .

Cau lerpa bikinens is Tay lor : Dawson , 1 957 .

Cau lerpa br achypus H ar v . : Taylor , 1950b; Dawson , 1 957 ; Gi lmar tin , 1 960 .

Cau lerpa elongata W . v . Bosse: Odum and Odum , 1 955; Dawson , 1 957 ; Ca lver t et al . , 1 976 .

Cau lerpa f i l ico ides Yamada: Dawson , 1957 .

Caulerpa acuta (Yamada) Yamada: Gi lmar tin , 1 960 .

Caulerpa peltata Lamx . : Gi lmar t in , 1 960 .

Caulerpa r acemosa (For sk ) J . Ag . z Tay lor , 1 950b; Dawson , 1 957; Gi lmar tin , 1 960 .

Cau lerpa ser ru lata (Por sk ) J . Ag . z Tay lor , 1 950b; Dawson , 1 957; G i lmar t in , 1 960 ;

Calvert et ai . , 1 976 .

Caulerpa ser tu lar io ides (Gmel ) H owe: Gi lmart in , 1 960 .

Cau lerpa tax ifo lia (Vahl) C. Ag . z Dawson , 1 957 .

Caulerpa u r vi ll iana Montagne: Tay lor , 1 950b; Dawson , 1 957; Ca lver t et ai . , 1 976 .

Cau lerpa ver t ici l lata J . Ag . : Gi lmar tin , 1960 .

Caulerpa webbiana Montagne: Dawson , 1 957; Gi lmar t in , 1960 ; Ca lver t et ai . , 1 976 .

Fam i ly CODIACEAEAvr ainvi llea lacer ata H ar v . : Tay lor , 1 950b ; Dawson , 1 957 .

Aur ainvi llea n ig r icans Deca isne: Gi lmar t in , 1 960 .

Cod ium ar abicum Dawson , 1 957 .

Cod ium edu le Si lva: Dawson , 1 957 .

Codium gepp i i O . C. Schm idt : Dawson , 1 957; Gi lmar t in , 1 960 .

‘Cod ium saccatum Okamur a: Dawson , 1 957 .

Cod ium ten ue K ii tz . : Tay lor , 1 950b; Dawson , 1957 .

H a l imeda copiosa Goreau and Graham : H i l l is-Co linvaux , 1 977 .

H al imeda opun t ia f . hederacea Bar ton : Tay lor , 1 950b ; Dawson , 1957; Gi lmart in , 1960 .

H alimeda cyl indr acea Decai sne: H i ll isC olinvaux , 1 977 .

H alimeda mon i le (So lander ) Lamx . : Dawson , 1 957 ; Gi lmar tin , 1 960 , 1 966 .

H al imeda discoidea Deca isne: Gi lmar t in , 1 960 .

H al imeda dis tor ta (Yamada) Colinvaux : H i l l is-Coi invaux , 1 977 .

H al imeda fr ag i l is Tay lor : Tay lor , 1 950b; H i l l i s , 1959; Gi lmar t in , 1960 , 1966 ;

H i l l is -Colv inaux ,

TH al imeda gigas Taylor : Tay lor , l 950b ; Dawson , 1 957 ; H i l l i s , 1 959 ; H i l l is -Col invaux , 1977 .

H a limeda graci lis H ar v . : Gi lmar tin , 1 960 ; H i l l i s—Col invaux , 1 977

H a limeda incras sata (El l is ) Lamx . : H i l l is -Colinvaux , 1977 .

H al imeda t r idens (E . and S . ) Lam x . : G i lmar t in , 1 960 .

H a limeda t r idens f . lamou roux i i (J . Ag . ) W .v . Bos se: Tay lor , 1 950b ; Dawson , 1 957 .

H a limeda t r idens f . t r ipar t i ta (Bar ton ) Co l l ins : Taylor , 1 950b .

H alimeda t r idens f . typ ica (Bar ton ) Co l l ins : Taylor , 1950b .

‘On ly specimen r ecorded fr om M icrones ia .

TNew species ; ho lotype based on Enewetak specimens . (Th i s table continued on next page. )

TSUDA

TABLE 1 (con t’d)

Fam i ly CoDiACEAE (con t’d)TH al imeda

‘

lacunal is Tay lor : Tay lo r , 1 950b ; Dawson , 1 957; H i l l is , 1 959 ; Gi lmar t in , 1 960 ; H i ll is—Col invaux , 1 977 .

H alimeda macr op hysa Askenasy: Dawson , 1 957; H i l l is —Colinvaux , 1 977 .

H alimeda m icrones ica Yamada: H i l l is -Colinvaux , 1 977 .

H alimeda m in ima (Tay lor ) Col invaux : H i llisC olinvaux , 1977.

H a l imeda opun t ia f . m in ima Tay lor : Dawson , 1 957; Gi lmar t in , 1960 .

H a limeda opun t ia (L . ) Lamx . : Tay lor , 1 950b; Dawson , 1 957; Gi lmar t in , 1 960 ; H i l l is -Colinvaux , 1 977 .

H al imeda s tuposa Taylor : Tay lor , 1 950b; Dawson , 1 957; H i l l is -Col invaux , 1 977 .

H al imeda taen icola Tay lor : Tay lor , 1 950b; Dawson , 1 957; Gi lmar t in , 1960 ; H i l l i s-Col invaux , 1977 .

Pseudochlorodesm is fu r cellata (Zanard ) Boer g . : Dawson , 1957 ; G i lmar t in , 1 960 .

Rh ip i l ia diaphana Tay lo r : Tay lor , 1 950b; Dawson , 1 957 .

Rh ip i l ia gepp i i Tay lor : Taylor , 1 950b; Dawson , 1 957 .

Rh ip i l ia or ien tal is A . and E . S. Gepp : Dawson , 1 957 .

Tydeman ia expedit ion is W . v .

‘Bosse: Tay lor , 1 950b; Dawson , 1 957 ; Gi lmar t in , 1 960 , 1 966; H i l l is -Col invaux , 1 977.

Udo tea indica A . and E . S. Gepp: Tay lor , 1 950b; Gi lmar t in , 1 960 .

Udo tea javens is (Mon t ) A . and E . S. Gepp : Tay lor , 1 950b ; Dawson , 1 957 ; Gi lmar t in , 1 960 .

Udo tea or ien ta lis A . and E . S. Gepp : G i lmar t in , 1 960 .

Udo tea palmetto Deca i sne : Dawson , 1 957 .

Fam i ly DERBESlACEAEDer bes ia attenuate Dawson : Dawson , 1957 .

Der bes ia mar ina (Lyngb) So l ier : Dawson , 1 957 .

‘Der bes ia neg lecta Reinbo ld : Gi lmar t in , 1 960 .

Der bes ia ryukiuens is Yamada and Tanaka : Dawson ,1957 .

Fam i ly PH YLLOSIPH ONACEAEOs t reobium r einecki i Bo r net : Dawson , 1 957 .

Order SiPH ONOCLADALESFam i ly BOODLEACEAEBoodlea compos ita (H ar v ) B r and : Tay lor , 1950b ; Dawson , 1 957; Gi lmar t in , 1 960 .

Boodlea s iamens is Reinbo ld : Tay lor , 1 950b ; Dawson , 1957 .

St r uvea anas tomosons (H ar v ) Piccone: Dawson , 1 957 .

Fam i ly SiPHONOCLADACEAECladophorop s is gr aci ll ima Dawson : Dawson , 1 957 ; Gi lmar tin ,

1 960 .

Cladophorops is s undanens is Reinbo ld : Dawson , 1 957 ; G i lmar t in , 1 960 .

Siphonocladus r ig idus H owe: Dawson , 1 957 .

Fam i ly VALON lACEAEBoergesen ia for bes i i (H ar v ) Feldmann : Dawson , 1 957 .

D ictyosphaer ia cauer nosa (Por sk ) Boerg . : Taylor , 1 950b; Dawson , 1 957; Webb et ai . , 1975 .

D ictyosphaer ia in termedia W . v . Bo s se: Tay lor , 1 950b; Odum and Odum ,1955 ; Dawson , 1 957 .

D ictyosphaer ia ver s luys i i W . v . Bos se: Dawson ,1957 .

Valon ia aegagr op i la C. Ag . z Tay lor , 1 950b ; Dawson , 1 957; Gi lmar t in , 1 960 .

Valon ia fas t ig iata H ar v . : Dawson , 1 957 .

Valon ia ut r icu lar is (Roth) C. Ag . z Dawson , 1 957; Gi lmar tin ,1 960 .

Va lon ia ven t r icosa J . Ag . : Dawson , 1 957; G i lmar tin , 1960 .

Valon iop s is pachynema (Mar tens ) Boer g . : Tay lor , 1 950b; Dawson , 1 957 .

Order DASYCLADALESFam i ly DASYCLADACEAEAcetabu lar ia moebi i So lm s

-Laubach : Dawson , 1 957 ; G i lmar t in ,1 960 .

Neomer is bi l imbata Kos ter : Dawson , 1 957 .

Order CLADOPH ORALESFam i ly ANADYOMENACEAEAnadyomene wr ight i i Gr ay : Dawson , 1 957 .

M icr odictyon japon icum Setch Dawson , 1 957 .

M icrod ictyon okam u rai Setch : Taylor , 1 950b; Dawson , 1957 .

Rhip idiphyllon r et iculatum Askenasy: Tay lor , 1 950b; Dawson , 1 957 .

Fam i ly CLADOPH ORACEAEChaetomo rpha indica Kijtz . : Dawson , 1 957 .

‘C ladophora albida (H udson ) K ii tz . : G i lmar tin , 1 960 .

Cladop hor a cr ys ta l l ina (Roth) Ku'tz Dawson , 1 957 .

‘Cladophora g r aci l is (Gr i ffi ths ) K ij t z . : G i lmar tin , 1 960 .

‘On ly specimen recorded fr om Micrones ia .

TNew species ; ho lotype based on Enewetak specimens . (Th is table con tinued on next page )

MARINE BENTH IC ALGAE

TABLE 1 (con t’d)

Fam i ly CLADOPH ORACEAE (cont ’d)‘C ladophor a inser ta D ick ie: Dawson , 1 957 .

C ladopho ra soc ial is K i i tz . : Dawson , 1 957 ; G i lmar tin , 1 960 .

‘Rhizoclon ium imp lexum (D i llwyn ) Kli tz . : Dawson , 1 957 .

D iv i s ion PH AEOPH YTA (brown algae)Class PH AEOPH YCEAEOrder ECTOCARPALESFam i ly ECTOCARPACEAEEctocarpus br eviar t icu latus J . Ag . z Dawson , 1 957 .

Ectocarpus m itchellae H ar v . : Tay lor , 1 950b; Dawson , 1 957 .

Feldmann ia indica (Sonder ) Womer s ley and Bai ley .Ectocarpus indicus Sonder : Tay lor , 1 950b; Dawson , 1957 .

Feldmann ia i r regu lar is (K ijtz ) H amel .Ectocarpus i r regu lar is Kii tz Dawson , 1 957

Order SPH ACELAR IALESFam i ly SPH ACELAR IACEAESphacelar ia fu r cigera K ii tz . : Dawson , 1 957 ; Gi lmar tin , 1 960 .

Sphace lar ia novaehollandiae Sonder : Tay lor , 1 950b; Dawson , 1957 .

Sphacelar ia t r ibulo ides Menegh. : Dawson , 1 957 .

Order D icTYOTALESFam i ly DiCTYOTACEAED ictyopter is repen s (Okam ) Boerg . : Dawson , 1 957 ; G i lmar t in , 1 960 , 1 966 .

D ictyota cer vicor n is K i l tz . : G i lmar tin , 1 960 .

D ictyo ta dichotoma Lamx Gi lmar t in , 1 960 .

D ictyota divar icata Lamx . : Dawson , 1 957 ; G i lmar tin , 1960 ;

Lowman and Palumbo , 1 962 ; Gi lmar t in , 1 966 .

D ictyota patens J . Ag . z Dawson , 1 957 .

D ictyota p in nat if ida K i i tz . : Tay lor , 1 950b; Dawson , 1957 .

Lobophor a papenfus s i i (Tay lor ) Womer s ley .Pocockiella papenfuss i i Tay lor : Tay lo r , 1 950b; Dawson , 1957 .

Lobophor a var iegata (Lamx ) Womer s ley : Sm ith and Mar sh , 1973 .

Pocockiella var iegata (Lamx ) Papenfu ss : Tay lor , 1950b; Dawson , 1957; Gi lmar tin , 1960 .

Zonar ia var iegata (Lamx ) Mer tens : Odum and Odum , 1955 .

Padina aus t ralis H auck : Dawson , 1 957 .

Padina tenuis Bo r y .Padina commer son i i Bo r y : Taylor , 1 950b; Dawson , 1 957 ; Gi lmar t in , 1 960 , 1 966 .

Spatog loss um schroeder i (Mer t . ) J . Ag . z Gi lmar tin , 1960 .

Order SCYTOSiPHONALESFam i ly SCYTOSiPH ONACEAEH ydroclathr u s clathr atus (Bo r y) H owe: Tay lor , 1950b; Dawson , 1957.

Rosenv ingea fas t igiata (Zanard ) Boerg . : Tay lor , 1 950b; Dawson , 1 957 .

Rosenvingea in t r icata (J . Ag . ) Boerg . : Tay lor , 1 950b; Dawson , 1 957.

Rosen vingea or ien talis (J . Ag . ) Boerg . : Gi lmart in , 1 960 .

Order DiCTYOSiPH ONAL ESFam i ly CHNOOSPORACEAEChnoospora imp lexa (H er ing) J . Ag . z Dawson , 1 957

Order FUCALESFam i ly SARGASSACEAE

“

Tu rbinari a om ata (Tur n ) J . Ag . z Tay lor , 1 950b; Dawson , 1 957 , 1 964 .

Di v is ion RH ODOPH YTA (red algae)Class BANG lOPH YCEAEOrder BANG lALESFam i ly ERYTH ROPEL TIDACEAEErythrotr ichia cornea (Di llwyn) J . Ag . z Tay lor , 1950b; Dawson , 1 957.

E rythrot r ichia par ieta lis Tanaka: Dawson , 1957 .

Order PORPH YRlD lALESFam i ly GONiOTRiCHACEAEAs terocys ti s om ata (C. Ag . ) H amel : Taylor , 1950b; Dawson , 1 957 .

Gon iotr ichum als idi i (Zanard ) H owe: Tay lor , 1 950b.

Gon io tr ichum elegans (Chauv in) Zanard. : Dawson , 1 957 .

‘On ly specimen r ecorded from M icr ones ia. (Th is table “con ti nued on next page. )

TSUDA

TABLE 1 (con t ’d)Clas s FLORIDEOPH YCEAEOrder NEMALlALESFam i ly ACROCH AETIACEAEAcr ochaet ium gr aci le Boerg . : Dawson , 1 957 .

Acrochaet ium r obus tum Boerg . : Dawson , 1 957 ; Gi lmar t in , 1 960 .

‘Kyl in ia cr as s ipes lBoer g ) Ky l in : Dawson , 1 957 .

‘Kyl in ia secunda (L yngb) Papenfuss : Dawson , 1 957 .

Fam i ly BONNEMA lSON lACEAEAspar agop s is tax iform is (Del i le) Co l l ins and H er vey : Dawson , 1 957 ; Gi lmar t in , 1960 ; Ger ber and Marshall ,F alkenberg ia h i llebr andi i (Bor net) Falkenberg : Dawson , 1 959 ; Gi lmar t in , 1960 .

Fam i ly CH AETANG lACEAEGalaxau ra fas t ig iata Deca i sne: Dawson , 1 957 .

Galaxau r a filamen tosa Chou : Dawson , 1 957 .

Fam i ly GEL iD lACEAEG el idiella bo r net i i (W . v . Bosse) Feldmann and H amel : Dawson , 1 957 .

Gel idiella tenu is s ima Feldmann and H amel : Dawson , 1 957

Ge l idium cr inale (Tur n ) Lamx Dawson ,

Ge l idium pus i l lum (Stackh) LeJoli s : Dawson , 1 957

Wu r demann ia m in iata (Lmk . and D . C . ) Feldmann and H amel : Dawson , 1 957 .

Fam i ly H ELMiNTH OCLADiACEAEL iagora far inosa Lamx . : Dawson , 1 957 .

L iagor a hawai iana Butter s : Dawson , 1 957 .

L iago ra or ien talis J . Ag . z Dawson , 1 957 .

L iagora p in nata H ar v . : Dawson , 1 957 .

L iagora r obus ta Yamada: Dawson , 1 957 .

Order CRYPTONEMIALESFam i ly CORALL INACEAEF os l iella far inosa (Lam x ) H owe: Dawson , 1 957 ; Lawr ence and Dawes , 1969 .

H eteroderma m inu tu la Fos l ie: Dawson , 1 957 .

H eteroderma s ubt i l is s ima Fos l ie: Dawson , 1 957 .

Jan ia cap i llacea H ar v . : Tay lor , 1 950b ; Dawson , 1 957 ; G i lmar tin , 1 960 ; Sm i th and Mar sh , 1 973 .

Jan ia decus sato -dicho toma (Yendo) Yendo : Dawson , 1 957 ; G i lmar t in , 1960 .

Jan ia m icr an throd ia Lamx . : Dawson , 1 957 .

Jan ia tene l la Kil tz . : Dawson , 1 957 .

Porol ithon cr aspedium (Fos l ie) Fos lie : Tay lor , 1 950b; Dawson , 1 957 .

Po r ol i thon gardiher i (Fos l ie) Fos lie: Tay lor , 1 950b; Dawson , 1957 .

Por ol ithon onkodes (H eyd ) Fos l ie: Tay lor , 1 950b; Mar sh 1 970 ; Sm ith and Mar sh , 1 973 .

L ithotham n ion sp . : Tay lor , 1950b .

Fam i ly CRYPTONEMIACEAEG rateloup ia f i l icina (Wulfen ) C . Ag . z Dawson , 1 957 .

Fam i ly PEYSSONEL lACEAECr uor iella dubyi (Cr ouan and Crouan ) Schm idt : Dawson , 1 957 .

Peys sonel ia rubra (Cr ev ) J . Ag . z Dawson , 1 957

Order GIGART INALESFam i ly DiCRANEMACEAED icranema r osal iae Setch . and Gard : Daws on , 1 957 .

Fam i ly H YPNEACEAEH yp nea esper i Bo r y : Dawson , 1957 ; Gi lmar t in , 1 960 .

H yp nea n idu lan s Setch . : Daws on , 1 957 .

H ypnea pannosa J . Ag . z Dawson , 1 957 .

H ypnea sp ine l la (C . Ag . ) K ijt z . : Tay lor , 1950b .

O rder RH ODYMEN IALESFam i ly CH AMPIACEAEChamp ia par vu la (Ag . ) J . Ag . z Dawson ,

1 957 ; Gi lmar t in , 1 960 .

Champ ia viei lardi i KLi tz . : Dawson , 1957 .

Lomen tar ia hakodatens is Yendo : Dawson , 1 957 .

Fam i ly RH ODYMEN IACEAEBot ryoc ladia skottsberg i i (Boerg ) Lev r ing : Dawson , 1 957 ; G i lmar tin , 1 960 .

Coe lo thr ix ll r egu lar is (H ar v ) Boerg . : Dawson , 1 957 .

Rhodymen ia anas tomosons W . v . Bo sse: Dawson , 1 957 ; G i lmar tin , 1 960 .

‘On ly specimen recorded fr om M icr ones ia . (Thi s table continued on next page. )

MARINE BENTH IC ALGAE

TABLE 1 (con t'd)Order CERAMIALESFam i ly CERAMIACEAEAn t ithamn ion lherm in ier i (Crouan and Cr ouan ) Nasr : Dawson , 1 957; G i lmar t in , 1960 .

‘i'An t i thamn ion per cu r r ens Dawson : Dawson , 1 957 .

Ant itham n ionella br evir amosa (Dawson) Wo llaston .

An t ithamn ion breviramous Dawson : Dawson , 1957.

"l’

Call ithamn ion mar shallens is Dawson : Dawson , 1957.

Cen tr oceras apicu latum Yamada: Dawson , 1957 .

Cen tr oceras clavu latum (C. Ag . ) Mont . : Dawson , 1957; Gi lmar tin , 1 960 .

Cen t roceras m in imum Yamada: Dawson , 1957 .

Ceram ium clar ionense Setch . and Gardn . : Dawson , 1957 .

Cer amium graci llimum Gr i ff . and H arv . : Dawson , 1957; G i lmar t in , 1960 .

‘TCeram ium mar shallense Dawson : Dawson , 1 957 .

Ceram ium mazat lanense Dawson : Dawson , 1957; Gi lmar tin , 1 960 .

Ceram ium serpens Setch . and Gardn . : Dawson , 1 957.

TCeram ium sympodiale Dawson : Dawson , 1957 .

‘Ceram ium taylor i Dawson : Dawson , 1957 .

TCeram ium vagabunde Dawson : Dawson , 1 957 .

‘Corynospora pedicel lata (Sm ith) J . Ag .

Neomonospo ra pedice l lata v . tenuis Feldm .

-Mazoyer : Dawson , 1 957 .

Crouan ia attenuata (Bonnemaison) J . Ag . z G i lmar tin , 1 960 .

‘G r iffiths ia ova lis H ar vey : Dawson , 1957 .

G r iff iths ia tenuis C. Ag .z Dawson , 1957 ; Gi lmar t in , 1960 .

H alop legma duper reyi Montagne: G i lmar tin , 1960 .

‘H erpos iphon ia nuda H o llenberg : H o l lenberg , 1 968c.

H erpos iphon ia pacif ica H o l lenberg : H o llenberg , 1 968c .

H erpos iphon ia parca Setch . : H o llenberg , l 968c .

H erpos iphon ia secunda (C. Ag . ) Ambronn : Dawson , 1957.

H erpos iphon ia tenel la (C. Ag . ) Naegel i : Dawson , 1 957 ; H o l lenberg , 1 968c .

H eter s iphon ia wu rdemann i (Bai ley) Falkenberg : Dawson , 1 957 ; G i lmar t in , 1960 .

Lophos iphon ia bermudens is Col l ins and H er v . : Dawson ,1957 .

‘Lophos iphon i

’

a cr is tata Falkenberg : H ol lenberg , l 968d .

Lophos iphon ia obscur a (Ag . ) Falkenber g : Dawson , 1 957 ; Gi lmar t in , 1 960 .

‘Lophos iphon ia vi llum (J . Ag . ) Setch . and Gard : Gi lmar tin , 1 960 .

‘Mesothamn ion car ibaeum Boerg . : G i lmar t in , 1 960 .

Po lys iphon ia coacta Tseng: Dawson , 1957 .

Po lys iphon ia ex i l is H arv . : H o l lenber g , 1968b.

Po lys iphon ia flaccidis s ima H o llenberg : H o l lenberg , 1968a.

“

l'Po lys iphon ia pen tamera H o l lenberg : H ol lenberg , l 968b .

Po lys iphon ia poko H o l lenberg : H o l lenberg , 1 968a.

Po lys iphon ia setacea H o l lenberg : H o llenberg , 1968a.

Po lys iphon ia subt i l iss ima Montagne: Dawson , 1 957 ; Gi lmar t in , 1960 .

Po lys iphon ia tongatens is H ar vey : Dawson , 1 957; Gi lmar tin , 1 960 .

Spermothamn ion saccor h iz a (Setch . and Gardn . ) Feldmann -Mazoyer ; Gi lmar tin , 1960 .

Spyr idia f i lamen tosa (Wulf) H ar v . : Tay lor , 1 950b; Dawson , 1 957 ; Gi lmar t in , 1960 .

Taen ioma perpus i llum J . Ag . z Dawson , 1 957 .

Tolyp iocladia calodictyon (H ar v ) Si lva.

Roschera calod ictyon (H ar v ) W . v . Bosse: Tay lor , 1950b; Dawson , 1 957 ; Gi lmar t in , 1 960 .

Wr angel ia anas tomosons Yamada: Dawson , 1 957 .

Wr angel ia argus (Mont ) Montagne: Dawson , 1 957 .

Wrangel ia pen ici llata C. Ag . : Tay lor , 1950b; Dawson , 1 957 .

Fam i ly DASYACEAEDasya adhaerens Yamada: Tay lor , 1 950b; Dawson , 1 957 .

Dasya cor ymbifera J . Ag . : Gi lmar tin , 1 960 .

Dasya iyengar i i Boerg . : Dawson , 1957; Gi lmar t in , 1 960 .

‘Dasya mo l lis H ar v . : Dawson , 1957 ; Gi lmar t in , 1 960 .

Dasyop s is gepp ii W . v . Bosse: Dawson , 1 957 .

D ictyu r us purpurascens Bor y : Dawson , 1957 .

‘On ly specimen recorded fr om M icr ones ia.

TNew species ; ho lotype based on Enewetak specimens . (Th is table continued on next page. )

TSUDA

TABLE 1 (con t’d)

Fam i ly DELESSER IACEAEH ypog loss um m in imum Yamada: Dawson , 1 957 ; Gi lmar t in , 1 960 .

Mar ten s ia f r ag i lis H ar v .

H em i t rema f rag i lis (H ar v ) Dawson : Dawson , 1957 .

Fam i ly RHODOMELACEAE‘Chondr ia ba i leyana (Montagne) H ar v Gi lmar t in , 1 960 .

‘Chondr ia m inu tu la W . v . Bosse: Dawson , 1957 .

‘Chondr ia po lyr h iz a Co l l ins and H er vey : Dawson , 1957 .

Chondr ia repens Boerg . : Dawson , 1957; Gi lmar tin , 1 960 .

‘Dawson iella bu lbo r hiz a H o l lenber g : H o l lenberg , 1 967 .

Laurencia mar iannens is Yamada: Taylor , 1 950b; Dawson , 1 957 .

Laurencia nana H owe : Dawson , 1 957 .

Laurencia par vipap i llata Tseng : Dawson , 1 957 .

‘On ly specimen recorded from Micr ones ia .

TH o lo type based on Enewetak specimens .

spec ies (n ine Chlorophyta, three Phaeophyta, and 28 Rho

dophyta) or 1696 of the species repor ted from Enewetakrepresen t the on l y co l lect ions known fr om the geogr aph icr egion of M icr ones ia (pr eceded by an as ter isk in Tablei t is un l ikel y that these species are un ique to Enewetak ;fu r ther in tens ive co l lect ions fr om other ar eas in M icrones iaw i l l no doubt pr ov ide add i t ional r ecor ds . To date,

ther ehas been no recor d of a seagr ass on Enewetak . Thalas s iahemp r ich i i (Ehr enb ) Ascher s is the on ly spec ies of

seagras s r epor ted from the Mar shal l i s lands , and i t has

been found on Uji lang Ato l l , Jalu i t Ato l l , and A i l inglapalapAto l l (T suda et ai . , Unt i l fu r ther in tens ive co l leet ions are made from other is lands in M icr ones ia, i t i s di ff i

cult to make a comparat ive anal ys is w i th the mar ine florafrom other is lands . A lthough i t is har dly fai r to compar ethe Enewetak mar ine flor a w i th that of Guam , a h ighis land located far ther wes t , some infer ences can be made.

As can be seen in Table 2 , ther e are s l ightly more generaknown fr om Guam (1 1 8 gener a) than Enewetak (1 06 genera), but ther e ar e mor e spec ies r epor ted fr om Enewetak .

Taylor ( 1 950a) descr ibed the dom inan t algae on the ato l land found i t was di ff icult to compar e the Mar shal l mar ineflora w i th those of other i s lands because of the pauc i ty of

TABLE 2

Compar i son of the Mar i ne Ben th i cF lor a of Enewetak and Guam

Gener a Spec iesD i v i s ion E

‘GT E

‘G'

l‘

Cyanophyta 1 2 1 0 1 6 13

Chlo rophyta 29 30 89 69

Phaeophyta 1 2 15 24 27

Rhodophyta 53 63 109 1 1

Total 106 1 1 8 238 1 20

co l lect ions from other i s lands . H e did men t ion thatSargas sum was no t iceably absen t on Enewetak . Th is genushas yet to be found on any of the ato l ls in the Mar shalli s lands (Doty , 1 954 ; Tsuda, Clear ly , the phycolog ical s tud ies conducted on Enewetak Ato l l have pr ov idedmuch of the info rmat ion on the mar ine ben th ic algaew ith in M icr ones ia .

Locat ions of the ear ly co l lect ion s i tes ar e avai lable in

the Depar tments of Botany and inver tebr ate Zoo log y atthe B . P . B ishop Museum .

REF ERENCES

Bakus , G . J. , 1 967 , The Feeding H abi ts of Fishes and Pr imar yProduction at En iwetok, Mar shal l i s lands , M icr ones ica, 3 :

1 35- 1 49 .

Calver t , H . E . , C . J . Dawes , and M . A . Bo row i tzka, 1 976 , Phylogenetic Relat ion sh ips oi Cau lerpa (Chlor ophyta) Based on

Comparat ive Chlor oplas t U ltrastructure, J. Phycol . , 1 2:

1 49- 1 62 .

Dawson , E . Y. , 1 957, An Annotated L i st of Mar ine A lgae fr omEn iwetok Ato l l , Mar shal l Is lands , Pac . Sci . , 1 1 : 92- 132 .

Doty , M . S. , 1 954 , D istr ibut ion of the A lgal Genera Rhip i l ia and

Sargassum in the Centr al Pacific , Pac . Sci . , 8 : 367-368 .

Drouet , F . , 1 968 , Rev is ion of the Class ification of the Osci l lator iaceae,

Monogr . Nat . Acad . Sci . Ph i la , 1 5: 1 -334 .

1 973 , Revis ion of the N os tocaceae with Cy lindr ical Tr ichomes , H afner Pr ess , N . Y .

and W . A . Dai ly , 1956 ,Rev i s ion of the Cocco id Myxophy

ceae, But ler Un iv . Bot . Stud , 1 2: 21 8 .

Ger ber , R . P. , and N . Mar shal l , 1 974 , Ingest ion of Detr itus by theLagoon Pelag ic Commun ity of En iwetok Atol l , L im nol .

Oceanog r .,19: 8 1 5-824 .

G i lmar t in , M . , 1960 , The Eco log ical D istr ibution of the DeepWater A lgae of En iwetok Ato l l , Eco logy , 41 : 2 1 0-221 .

1 966 , Eco logy and Mo rpho logy of Tydeman ia expedi t ion is , a

T r opical Deep Water Siphonous Gr een A lgae, J . Phycol . , 2:1 00 - 105 .

H i l l is , L . W . , 1 959 , A Rev is ion of the Genus H alimeda (Or derSiphonales ) , Pub l . i ns t . Mar . Sc i . Un iv . Texas , 6 : 321

-403 .

H i l l is -Col invaux , L . , 1977 , H a limeda and Tydeman ia: D istr ibut ion ,

D iver s ity and Product iv ity at Enewetak , Th ird i n ter nat ionalCor al Reef Sympos ium Proceedings (M iam i) , pp . 365-370 .

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Chapter 2

F ung i of Enewetak Ato ll

PAUL H . DUNN‘AND DON

‘R iver s ide F i re Labo r ator y , USDA F ores t Ser viceR iver s ide, Cal ifor n ia 92507

TNatu r al H is tor y Museum , Los Angeles Coun tyLos Ange les , Cal ifor n ia 90007

i N TRODUCTlON

The m icr ofung i of Enewetak Atol l are typical tr op icalforms and are not r es tr icted to that atol l . Because of the

extr eme isolation of Enewetak , however , i t seems l ikelythat the fung i there are indigenous to the atol l and are not

introduct ions by man . Whether the or ig in is ter res tr ial orfrom the beaches r emains a ques tion .

The fir s t s tudy of m icr ofung i on Enewetak (Spar row ,

1948) isolated un icel lular chytr ids of thr ee di fferent fam il ies fr om so i l samples (Table but no fi lamentous chy

TABLE 1

Check l i s t of Enewetak A toll Fungi

K ingdom MYCETEAEClass CH YTRIDIOMYCETINAOrder CH YTR ID IALESFam i ly OLPlDlACEAEO lp idium rhiz ophlyct idis Spar r ow: Spar r ow, 1948 .

Fam i ly PH LYCT IDIACEAERhiz Ophydium mar shallense Spar row : Spar row, 1948 .

Fam i ly RH IZIDIACEAERh iz ophlyct is r osea (DeBary and Woron in) Fischer : Spar r ow, 1948 .

Class ZYGOMYCOT INAOr der MUCORALESFam i ly PIPTOCEPH AL IDACEAE

‘Syncephalas t r un racemosum Cohn ex Schr oter , 1 886 .

Fam i ly TH AMN ID IACEAE‘H el icos tylum g lomeratum (van T ieghem and Le Monn ier ) van T ieghem , 1 876 .

Fam i ly MUCORACEAEAbs idia corymbifera (Cohn) Saccardo and T r otter : Dunn , 1 973 .

Mucor sp . : Dunn , 1 973 .

Rh iz opus n igr icans Ehrenberg : Dunn , 1 973 .

Order ENTOMOPH THORALESFam i ly ENTOMOPH TH ORACEAECon idiobo lus sp . : Dunn , 1973 .

‘New Enewetak Ato l l r ecord . (Th is table continued on next page . )

tr ids or Oomycetes were iso lated . An extens ive s tudy(Dunn , 1973) i so lated 1 0 1 spec imens from the beaches of

Enewetak . An add i t ional ei ght newly r ecor ded taxa are

l is ted in Table 1 . The specimens were iso lated by theWar cup plat ing method (War cup , wh ich es tabl ishedon ly that v iable propagules were pr esen t . Whether the fungal pr opagules of the Enewetak beaches ar e dorman t oract ive was not defined . Fungal propagules have beenpreser ved for long per iods when s tor ed in d is ti l led water(Campbel l and Stewar t , i t can be assumed , ther efor e, that nutr ient-poo r seawater could pr ov ide a hab itat inwh ich those propagules that to lerate sal in i ty could remainv iable.

Repo r ts fr om the atol l beaches (Dunn , 1973 ; Steele,

1 967) are based on i so lations fr om enr ichmen t culture.

Mar ine fungi have been defined as those obser ved grow ingon a subs tr ate in a mar ine habi tat (Kohlmeyer and

DUNN AND REYNOLDS

TABLE 1 ( con t'd )

Class ASCOMYCOT INAOrder SPH AER IALESFam i ly CH AETOMIACEAEAs cotr icha guamens is Arnes : Dunn , 1973 .

Chaetom ium globosum Kunze ex Fr ies : Dunn , 1 973 .

Fam i ly MELANOSPORACEAE‘Melanospo ra sp .

Order PLEOSPORALESFam i ly SPORORMIACEAESpororm ia sp . : Dunn , 1 973 .

Order EUROT IAL ESFam i ly GYMNOASCACEAEEmer ice l lops is m in ima Sto lk : Dunn , 1 973 .

Gymnoascus r ees i i Bar anet sky: Dunn , 1973 .

Or der MICROASCALESFam i ly MICROASCACEAEM icroascus cinereus (Em i le-Wei]and Gaudin ) Cur zi : Dunn , 1 973 .

Microascus t r igonospo r us Emmons and Dodge: Dunn , 1 973 .

Class DEUTEROMYCOT INAOrder SPH AEROPSIDALESFam i ly SPH AERIOIDACEAELas iod ip lodia theobr omae (Pat ) Gr i ff . and Maubl . : Dunn , 1973 .

Order MELANCON IALESFam i ly MELANCONACEAEPes talo t ia sp . : Dunn , 1 973 .

Order MON IL IALESFam i ly MON IL IACEAEAcremon ium baci ll iSpo r um (On ions and Bar r on) W . Gai n s (Teleomorph: Chaetom ium Dunn , 1973 .

‘Acremon ium char t icola: (L indau) W . Gams , 1 971 .

Acr emon ium p ter idi i W . Gams and Fr ank land : Dunn , 1973 .

Acr emon ium ter r ico la (M i ller et ai . ) W . Gams : Dunn , 1973 .

Aspergi l lus acu leatus i i zuka: Dunn , 1 973 .

Aspergi l lus car neus (v . T iegh) B lochwi tz : Dunn , 1973 .

Aspergi l lus conjunctus Kwon and Fennel l : Dunn , 1973 .

Aspergi l lus flavipes (Bain . and Sar t . ) Thorn and Church : Du nn , 1 973 .

Aspergi l lus f lavus L ink : Dunn , 1973 .

Aspergi l lus ochr aceus W i lhelm ; Dunn , 1973 .

Aspergi l lus or yzae (Ahlbu rg) Chon : Dunn , 1973 .

Aspergi l lus recu r vatus Raper and Fennell : Dunn , 1 973 .

Aspergi‘l lus ter r eus Thom : Dunn , 1 973 .

Aspergi l lus ter r ico la? Marchal : Dunn , 1 973 .

Aspergi l lus tubingens is (Schober ) Mosseray: Dunn , 1 973 .

Aspergi l lus us tus (Bain ier ) Thom and Church : Dunn , 1973 .

Aspergi l lus var iecolor (Ber k . and Br . ) Thorn and Raper : Dunn , 1 973 .

Aspergi llus ver s ico lor (Vu i ll ) T i r aboschi : Dunn , 1973 .

Aspergi l lus wen t i i gr oup : Dunn , 1 973 .

Emmons ia sp . : Dunn , 1 973 .

Geotr ichum sp . : Du nn , 1 973 .

H an sfordia oval ispo ra H ughes : Dunn , 1 973 .

Nodu l ispo r ium sp . : Dunn , 1 973 .

Oidiodendron sp . : Dunn , 1 973 .

Paeci lom yces sp . : Dunn , 1 973 .

Paeci lomyces l i lacinum (Thorn ) Samson : Du nn , 1 973 .

Pen ici l lium sp . (monovert ici l late, with sclerotia) : Du nn , 1 973 .

Pen ici l lium sp . (polyver t ici llate): Dunn , 1 973 .

Pen ici l lium canescens Sopp . : Dunn , 1 973 .

Pen ici l lium citr eo -vir ide B iourge: Dunn , 1973 .

Pen ici l lium corylophi lum Dierckx : Dunn , 1973 .

Pen ici l lium f requen tans West l ing : Dunn , 1 973 .

‘New Enewetak Ato l l record . (Th is table continued on next page

F UNG I

TABLE 1 ( con t ’d )

Fam i ly MON IL IACEAE (cont ’d)Pen ici l lium fun ico los um Thorn : Du nn , 1973 .

Pen ici l lium jan thinellum B iourge: Dunn , 1 973 .

Pen ici llium ver r ucu losum Peyr onel : Dunn , 1973 .

Ph ialophora sp . : Dunn , 1 973 .

Spo r o th r ix sp . : Dunn , 1 973 .

Spo r o tr ichum sp . : Dunn , 1973 .

‘Tr ichoderma aur eovir ide R ifai , 1969 .

‘Tr ichoderma har z ian um R ifai , 1 969 .

‘Tr ichoderma p seudokongi i R ifai , 1 969 .

Tr i t i r ach ium sp . : Dunn , 1 973 .

Vert ici l l ium lecan i i (Zimm ) V iegas : Dunn , 1 973 .

Yeast : Dunn , 1 973 .

Fam i ly DEMAT IACEAEAcr ophialophor a fus ispo ra (Saksena) M . B . El l is : Dunn , 1 973 .

Aureobas idium pu l lu lon s (DeBary) Am aud : Dunn , 1 973 .

Cercospo ra sp . : Dunn , 1973 .

C ladospo r ium cladospor io ides (Fr esen ) de V r ies : Dunn , 1973 .

Cladospo r ium her bar um (Per s ) L ink ex S. F . Gray : Du nn , 1973 .

Cladospo r ium sphaerosperm um Pen z . : Du nn , 1 973 .

Cur vu lar ia sp . : Du nn , 1 973 .

Cur vu lar ia clavata Jain : Dunn , 1 973 .

Cur vu lar ia lunata (Wakker ) Boedijn : Du nn , 1973 .

Cur vu lar ia or yzae Bugn icour t : Du nn , 1 973 .

Cur vu lar ia senegalen s is (Speg ) Subram . : Dunn , 1 973 .

Cur vular ia tuber cu lata Jain : Dunn , 1 973 .

D rechs lera aus t ral iens is (Bugn icour t) Subr am . and Jain Ex M . B . El l is : Du nn , 1973 .

D rechs lera halodes (Drechs ler ) Subr am . and Jain : Dunn , 1973 .

D r echs lera hawai iens is (Bugn icourt ) Subr am . and Jain Ex M . B . El l is : Dunn , 1973 .

D r echs lera papendorfii (van der Aa) M . B . El l i s : Dunn , 1 973 .

G l iomas t ix m u rorum (Corda) H ughes : Du nn , 1 973 .

G l iomas t ix m ur or um (Corda) H ughes var . felina (Marchal) H ughes : Du nn , 1 973 .

G l iomas t ix m u ro r um (Corda) H ughes var . polychroma (van Beyma) Dickinson : Dunn , 1 973 .

H um ico la fuscoat ra T r aaen : Dunn , 1 973 .

H um icola sp . : Dunn , 1973 .

Lep tog raphium sp . : Dunn , 1 973 .

Mammar ia sp . : Dunn , 1 973 .

Memnon iella ech inata (R iv ) Ga l loway : Dunn , 1973 .

N igr ospo r a sphaer ica (Sacc ) Mason : Dunn , 1 973 .

N igr ospora state of Khuskia o r yzae H udson : Dunn , 1973 .

Rh inoc ladiel la cellar is (Per s . ex S. F . Gr ay) M . B . El l is : Dunn , 1 973 .

Scolecobas idium hum icola Bar r on and Busch : Dunn , 1 973 .

Scolecobas idium var iabi le Bar r on and Busch : Bunn , l 973 .

Scopu lar iops is baar nen s is Mor ton and Sm i th : Dunn , 1 973 .

Scopu lar iops is br ump t i i : Dunn , 1973 .

Scopu lar iops is brevicau l is (Sacc) Ba in ier : Dunn , 1973 .

Scopu lar iops is char tar um (Sm ith) Mor ton and Sm ith : Dunn , 1 973 .

Scopular iops is f im ico la (Cos t . and Mai r .) Vu i ll . : Dunn , 1 973 .

Scopular iops is sphaerospo r a Zach : Dunn , 1 973 .

Scopu lar iops is sp . : Dunn , 1973 .

Stachybo t rys at r a Corda: Dunn , 1 973 .

Stachybo t rys state of Melanop samma pom iform is (Per s . ex F r . ) Sacc . : Dunn , 1 973 .

Stemphyl ium ves icar ium (Wall r ) Simmons : Dunn , 1 973 .

Tr ichocladium canadense H ughes : Dunn , 1973 .

‘U loc ladium char tar um (Pr euss ) Simmons , 1 967Fam i ly STILBACEAEGraph ium put redin is (Corda) H ughes : Dunn , 1 973 .

i sar ia sp . : Dunn , 1 973 .

‘New Enewetak Ato ll r ecord . (Th is table continued on next page . )

1 4 DUNN AND REYNOLDS

TABLE 1 ( con t’d)

Fam i ly TUBERCULAR IACEAECy lindrocarpon sp . : Dunn , 1 973 .

Fusar ium ep isphaer ia (Tode) F r . : Dunn , 1 973 .

Fusar ium so lan i (Mar t ) Sacc . : Dunn , 1973 .

Fusar ium sp . : Dunn , 1 973 .

Metar r hiz ium an isop l iae (Metschn ) Sor ok Dunn , 1973 .

Myr othecium r or idum Tode ex Steudel : Du nn , 1973 .

Myr othecium s tr iat iSpo r um Pr eston : Dunn , 1973 .

Class BASID IOMYCOT INAUn identi fied ai r i so late.

‘New Enewetak Ato l l record .

Kohlmeyer , None of the fung i isolated fromEnewetak , ther efor e, fi ts the defin i tion of a mar ine fungu s .

inh ib i tor s in seawater (K i r k , 1 980) and in so i l (Lockwood ,1 977) may pr event fungal spor es fr om germ inat ing unti lchanges occur in the endogenous or exogenous nutr ien ts tatus or un t i l the exogenous spore inh ibi tor s are removed .

The exogenous inh ibi tor s of nonmar i ne fu ngi can be par t lyremoved or deactivated by autoclav ing seawater ; mos tmar ine fu ngi can germ inate in r aw seawater . Addi tion of

g lucose can also coun teract the inh ibi tion (Ki r k ,Few s tudies have been done on decompos i t ion of sub

t idal subs tr ate. in a s tudy of seagras s , Thalas s ia tes tudin umKon ig , Newel l and Fel l (1 980) showed that although nonmar ine fungi wer e pr esen t , they played on ly a m inor rolein decompos i t ion of T . tes tudinum leaves un t i l the leaveswer e depos i ted in the inter tidal . The ter r es tr ial fung i areactive in the inter t idal zone , but thei r s ign i ficance in relat ion to mar ine fungi i s unknown (Newel l and Fel l , 1 980 ,

On the bas is of the presen t d iscus s ion , i t can be

sugges ted that the fungal propagules in Enewetak Ato l lbeaches are also dorman t un ti l they come in contact w i th apr oper subs tr ate.

Fung i fr om the beaches of Enewetak Atol l have beencompar ed w i th those fr om other locat ions . When the

specimens iso lated by Dunn (1 973) were compared w i thfung i plated fr om sediment of a coas tal es tuary of Nor thCar ol ina w i th sal in i ties fr om 0 to (Bo r ut and Johnson , a D ice s imi lar i ty quot ient of 1 796 was found ,indicating a d is t inct biota for these two ver y d i ffer entmar ine habi tats .

In a compar i son of the fung i iso lated by War cup p lating fr om beaches at Enewetak i s land and the H awai ianIs lands , a D ice s im i lar i ty quot ien t of 43% was found (Dunn ,

The beaches in the H awai ian Is lands ar e bo th volcan ic and car bonate, wher eas the beaches at Enewetaki s land ar e car bonate. Sim i lar i ty is gr eater between the

mycobiota of the car bonate beaches in Enewetak and

H awai i than between the car bonate and volcan icbeaches in H awai i The pr incipal d i ffer ence betweenthe mycobiota of the car bonate beaches sampled in

Enewetak and those in H awai i was the abundance of

species pr esen t in bo th beaches r elat ive to the total i solates . Typ i cal of the patter n ar e Gymnoas cus r ees i i

Baranetsky and two species in the genus M icr oas cus ,wh ich wer e common at Enewetak i s land beach but werein fr equen tly i so lated fr om Kahala Beach , Oahu , H awai i .

CH YTRID IOMYCOTIN A

The i so lates of th i s class were all un icel lu lar .

Rhizophyct is rosea (DeBary and Wor on in) Fischer was themos t common iso late, and i t is the mos t w idespr ead of allchytr ids (Spar row , Un icellular chyt r ids were foundin the beach areas by Dunn ( 1973 ) but wer e not iden ti fied .Spar r ow (1 948) cons i s tently found on ly the same threespecies of th i s class on Enewetak and on thr ee otherMar shal l i s land ato l ls . The fi lamen tous species of the classwer e consp icuous ly absent .

OOMYCOTIN A

A l though many spec ies in th is class are common plan tpathogens and would be expected to be assoc iated w i ththe vegetat ion , none has been repor ted fr om thi s atoll .They may be too sens i t ive to sal in i ty to do wel l in beachhabi tats (TeSt r ake, 1 959 ; Lee and Baker , al thoughsome have been iso lated from other beach ar eas .

ZYGOMYCOTIN ACon idiobo lus sp . was iso lated fr om subt idal areas on

Enewetak . Species in this genus are normally assoc iatedw i th dung of insect-eat ing an imals or decay ing vegetat ion .

Th is spec ies was isolated on ly fr om seawater sour ces and

seems to r equ i r e sal in i ty . It does not r esemble any

descr ibed species and does not las t many generat ions incultur e. Member s of the or der Mucorales are common isolates fr om beaches at Enewetak but have sal in i ty opt imamor e appropr iate for ter res tr ial species (Du nn ,

ASCOMYCOTINAMany ascomycetes are known to produce two types of

spor es : the teleomorph , or sexual s tate, and the

anamorphs , or asexual s tates . The teleomorph may be

assoc iated wi th one or more anamorphs . The teleomorphgenus r egular ly assoc iated wi th the isolated Scopular iop s is

FUNG I

anamorph , for example, is wel l documen ted but was not

iso lated in the s tudies . Ascot r icha guamens is Ames prev iou sly was isolated fr om Guam , New Guinea , ind ia, and

mor e recently , fr om so i l at Enewetak . M icroas cus cinereus(Emele-Wei l and Gaudin) Gu r z i , the teleomorph , and i ts

as soc iated anamorph were among the mos t frequently isolated fungi in the beach s tudy . These wer e r arely iso latedfr om car bonate beaches in H awai i . Gym noascus rees i i is

typ ical of the sal ine habi tats all thr ough the Paci fi c bas in(Baker and Meeker ,

DEUTEROMYCOT IN AAt Enewetak Atol l , the major i ty of the Deuter o

mycot ina genera i so lated fr om beaches was compr ised ofmember s of the or der Mon i liales (94 taxa), w i th the

major i ty of species even ly div ided between the fam i l iesDemat iaceae and Mon i l iaceae. Acr emon ium baci ll ispor um

(On ions and Bar r on) W . Garns was a pr ominent member ofthe car bonate sand commun i ty at bo th Enewetak and

Kahala Beach , Oahu , H awai i (Dunn , The asper gi l liwer e common and the pen ici l lia were r ar e in beaches at

both loca l i t ies (Dunn , These patter ns co incide w i thother data ind i cating that the asperg i l l i ar e mor e commonin the subtropi cs , and the pen ici l l ia are mor e common in

temper ate cl imates . Gr aph ium putredin is (Cor da) H ugheswas found common ly in the anaerobic zone in deeper sediments off Enewetak Ato l l . F usar ium solan i (Mar t ) Sacc .

was common in beaches at Enewetak but mos tly wasfound near the land zone. The mon i l iaceous genusD reschlera is r epr esented on Enewetak Ato l l by fourspecies (Table All ar e known fr om the tropics and

souther nmos t Afr i ca . Two of the species of

C ladospor ium, C . cladospor io ides , and C . sphaerosper

m um are found in tr op ical and subtr opical cl imatesin sal ine habi tats . A thi r d species , C . her bar um , was

not found fr equen tly at Enewetak and i s cosmopo l i tan in

di s tr ibut ion .

BASIDIOMYCOTINAThe mos t common isolate fr om ai r at Enewetak Ato l l is

a bas idiomycete iden ti fiable by large clamp connections on

the hyphae (Dunn , I t did not spor ulate in cu l tu r e.

Other bas idiomycetes in the Mar shal l i s lands , mos tly associated w i th decay ing wood , were collected by Roger s( 1 947) and Taylor They did not co l lect fung i onEnewetak Ato l l , and fungi fr om decay ing wood have not

been s tudied on Enewetak .

CONCLUSIONThe myco log ica l data fr om Enewetak Atol l r eflect two

pr i ncip les of fungal biogeogr aphy : s im i lar r eg ions haves imi lar biotas , and d is tr ibut ional data ar e oft en due to biascol lect ion act iv i ty of mycologi s ts (Baker and Meeker ,

For example, the genus Myr othecium was in i t ial lyrepor ted in the Paci fic fr om New Zea land . Th is genus was

1 5

then repor ted from Moor ea, Society i s land (Peter son ,

As data fr om other Paci fic is lands became avai lable,

the d is tr ibution was en lar ged to include Tonga, H awai i ,New Caledon ia (Baker and Meeker , and Enewetak(Baker and Meeker , 1972 ; Dunn , A s im i lar expan

s ion thr ough the tr op ical Pac i fic may be expected for mos tof the spec ies r epo r ted here.

ACKNOWLEDGMENTSThe author s thank G . E . Baker for help dur ing the

beach s tud ies and acknowledge the financial suppor t of theU . S. Nuclear Regulator y Comm iss ion Gran t AT -2)-229

and the use of i ts M id-Paci fic Resear ch Labo r ator y .

REF ERENCES

Baker , G . E . , and J . A . Meeker , 1972 , Ecosystems , Myco logi s tand the Geo log ica l Di str i bution of Fungi in the Centr al Pacifi c,Pac . Sci . , 26 : 4 18-432 .

Bo rut , S. Y. , and T . W . Johnson , Jr . , 1962, Some B io log ica lObser vations on Fungi in Es tuar ine Sediments , Myco logi’c , 54 :

1 8 1 - 1 93 .

Campbe l l , M . C. , and J . L . Stewar t , 1 980 , The Medical Myco logyH andbook , John W i ley and Sons , New York .

Dunn , P . H . , 1973 , The Eco logy of Fi lamentous H eterotrophicM icroo rgan i sms in Subtr opical and T ropica l Mar i ne PsammenH abitats , Ph .D . disser tation , Un iver s ity of H awai i .

Ki rk , P . W . , Jr . , 1 980 , The Mycos tatic Effect of Seawater on

Spor es of Ter r es tr ial and Mar ine H igher Fungi , Bo t . Mar . , 23 :

233 -238 .

Kohlmeyer , J.

, and E . Kohlmeyer , 1 979, Mar ine Myco logy, TheH igher Fungi , Academ ic Pres s , New York .

Lee, B . K . H . , and G . E . Baker , 1972, An Eco logica l Study ofthe So i l Micr ofungi in a H awai ian Mangr ove Swamp, Pac .

Sci . , 26 : 1 — 1 0 .

Lockwood , J . L . , 1977, Fungistas is in Soi ls , B io l . Rev . (Comb.

Ph i los . Soc) , 52 : 1 -43 .

Newel l , S. Y. , and J . W . Fell , 1 980 , Mycoflor a of Tur t legrass

(Tha las s ia tes tudinum Kon ig) is Recorded Aft er Seawater Ineubat ion , Bo t . Mar . , 23 : 265-275 .

1 982 , Surface Ster i l i zation and the Active Mycoflora of

Leaves of a Seagrass , Bo t . Mar . , 25 : 339-346 .

Peter son , R . H . , 1960 , Some So i l and Copr oph i lous Fungi fr omthe South Pacific Area, Mycologia, 52: 552-556 .

Roger s , D . P . , 1 947 , Fungi of the Mar shal l Is lands , Cen tral Paci ficOcean , Pac . Sc i . , 1 : 92- 107 .

Spar r ow, F . K . , 1 948 , So i l Phycom ycetes fr om B ikin i , En iwetok ,Ronger ik , and Rongelap Ato l ls , Myco logic , 40 : 445-453 .

1 960 , Aquat ic Phycom ycetes , 2ad edition , Un iver s i ty of

Mich igan Pr ess , Ann Ar bo r .

Steele, C . W . , 1 967, Fungu s Populations in Mar ine Water s and

Coas tal Sands of H awai ian , L ine, and Phoen ix i s lands , Pac .

Sci . , 21 : 3 17—33 1 .

Tay lo r , W . R . , 1 950 ,P lan ts of B ikin i and other Nor ther n Mar shal l

Is lands , Un iver s i ty of M ich igan Pr ess , Ann Ar bor .

TeSt r ake, D . , 1959 , Es tuar i ne Di s tr i but ion and Sal ine To leranceof Some Sapr o legn iaceae, Phyton , 1 2 : 1 47- 1 52 .

War cup , J . H . , 1 950 , The So i l P late Method for the i so lation of

Fungi fr om the So i l , Nature 166 : 1 1 7- 1 1 8 .

https://www.forgottenbooks.com/join

18 LAMBERSON

TABLE 1

Check l i s t of Ter r es tr i al Vascu lar Plan ts of Enewetak A toll‘

Di v i s ion TRACH EOPH YTAClass PSILOPSIDA

Order PSILOTALESFam i ly PSILOTACEAETPs i lotum nudum (L ) Beauvais : Lamber son , 1982; Fosberg et ai . , 1982 .

Class PTEROPSIDAOrder FIL ICALESFam i ly POLYPOD IACEAETPolypod ium sco lopendn

‘

a Bu rmann f .Phymatodes sco lopendr io (Burmann f . ) Ch ing : St . John , 1960 .

Po lypodium phymatodes L inn : Br yan , 1944 .

Class ANG IOSPERMAESubc lass MONOCOTYLEDONEAEFami ly PANDANACEAE¢Pandanus sp . : B r yan , 1944 ; Tay lor , 1 950 ; Woodbur y , 1962 ; Fal l et ai . , 1971 ; Kot anda et ai . , 1973 ; Lamber son , 1982 .

TPandanus tector ius Par k : Fosberg , 1 955; Welander et ai . , 1 966 .

‘i'Pandun us brachypodus Kaneh i r a: Kaneh i r a, 1935 ; St . John , 1960 .

‘l'Pandan us enchabiens is Kaneh i r a: Kaneh i ra, 1935 ; St . John , 1960 .

"( Pandanus kor r or en s is Kanehi ra: St . John , 1960 .

TPandan us odoratiss im us L . f . var . novocaledon icus (Mar tell i ) St . John : St . John , 1 960 .

TPandanus odorat is s im us L . f . var . novogu ineens is (Mar tel l i ) St . John : St . John , 1960 .

TPandanus pu lposus (War b ) Mar tel l i : St . John , 1960 .

f Pandanus rectanu latus Kanehi ra: Kaneh i ra, 1935 ; St . John , 1960 .

TPandanus r hombocarp u s Kaneh i ra: Kaneh i ra, 1 935 ; St . John , 1 960 .

“

( Pandanus u t iyamai Kaneh i ra: St . John , 1 960 .

Fam i ly GRAMINAECenchrus brown i i Roemer and Schultes : Tay lor , 1950 ; St . John , 1960 .

Cenchrus echinatus L . : Br yan , 1 944 ; St . John , 1 950 ; Tay lor , 1 950 ; Fosberg , 1 955 ; St John , 1960 ; Welander et ai . , 1 966 ;Lamber son , 1 982 .

Cenchr u s sp . : Fall et ai . , 1971 .

Ch lor is inf lata L ink : St . John , 1 950 ; Fosberg , 1955; St . John , 1960 ; Welander et ai . , 1966 ; Lamber son , 1982 .

Cynodon dacty lon (L ) Per soon : B ryan , 1944 ; Fosberg , 1955 ; St . John , 1960 ; Welander et ai . , 1966 ; Koranda et ai . , 1973 ;

Lamber son , 1 982 .

Dacty locten ium aegyp t icum (L ) Wi lldenow : St . John , 1 960 ; Welander et ai . , 1 966 ; Lamber son , 1 982.

TD igi tar ia set igera Roemer and Schultes : St . John , 1960 ; Lamber son , 1 982 .

D igi tar ia p r u ri ens (T r in ius ) Buese: Taylor , 1 950 .

D igi tar a m icrobachne (Pr es l) H enrard: Fosber g , 1 955 .

E leus ine indica (L ) Gaer tner : B r yan , 1944 ; Tay lor , 1950 ; Fosberg , 1955 ; St . John , 1 960 ; Welander et ai . , 1966 ;Lamber son , 1982 .

Eragr os tis scabr ifol ia Swallen : Lamber son , 1 982 .

Eragros tis tenel la (L ) Roemer and Schultes : Lamber son , 1982 .

Eragr ost is ambi l is (L ) W ight and Ar nott : Tay lor , 1 950 , Fosberg , 1955 ; St . John , 1 960 ; Welander et ai . , 1966 .

E ragr os ti s sp . : Lane, 1960 .

Lep tu ropet ium mar shallense Fosberg and Sachet , 1 982 .

Lep tu r u s repen s (For ster f . ) R . Brown var . occiden talis : St . John , 1 960 .

Lep tu r u s r epens (For s t . f . ) R . B rown : Br yan , 1944 ; Tay lor , 1 950 ; Fosberg , 1 955 ; Palumbo , 1 962; Woodbury , 1962 ;Welander et ai . , 1 966 ; Fal l et ai . , 1971 ; Koranda et ai . , 1 973 ; Lamber son , 1 982 .

Lep tur us sp . t Lane, 1960 ; Carpenter et ai . , 1 968 .

l lLep tur us repens (For s ter f . ) R . B rown var . subu latus Fosberg .

B r ach iar ia mut ica (For sskal) Stapf . : Lamber son , 1982 .

Setar ia ver t ici l lata (L ) Beauvai s : B r yan , 1 944 ; Tay lor , 1950 ; Fosberg , 1955 ; St . John , 1 960 .

TThuarea in vo luta (For ster f . ) Roemer and Schultes : Tay lor , 1950 ; Fosberg , 1 955 ; St . John, 1960 ; Lamber son , 1982 .

Tr icachne insu lar is (L ) Nees : Fosberg , 1 955 ; St . John , 1960 ; Welander et ai . , 1966 ; Lamber son , 1982 .

Tr icho laena rosea Nees

Tr icho laena repen s (Wi lldenow) H i tchcock : Tay lor , 1 950 ; Lane, 1 960 ; St . John , 1 960 .

‘Excludes plant species r epo r ted on ly fr om gardens , wh ich ar e cons ider ed to be tr ans ient components of the flora.

“

( Species cons idered indigenous to M icrones ia.

taxonomy of the genus Pandanus is disputed . Some taxonom ists would cons ider most of the species l isted here as form s of thespecies Pandanus tector ius .

l lNew Enewetak record ; see Table 4 . (Th is table cont inued on next page. )

H ISTORY OF TERRESTR IAL VASCULAR PLANTS 19

TABLE 1 (con t'd)

Fam i ly CYPERACEAECyperus comp ress us L . : Lamber son , 1 982 .

TCyper us javan icus H ou tt uyn : St . John , 1 960 .

Cyperus fer ox L . C . R ichardCyper us odoratus L . : St . John , 1 960 .

Cyperus r o tundus L . : Lamber son , 1982 .

TF imbr is tylus cymosa R . B r own ssp . Spathacea (Roth) Koyama, 1964 .

F imbri s tylus annua Roemer and Schultes forma diphyll ia (Retz) Kukenth : Bryan , 1 944 .

F imbr is tylus cymosa R . Br own : Tay lor , 1950 .

F imbr is tylus atollens is St . John : Lane, 1960 ; St . John ,1960 ; Welander et ai . , 1966 ; Fall et ai . , 1 971 ;

Koranda et ai . , 1 973 ; Kot anda et ai . , 1 978 ; Lamber son , 1 982 .

F imbr is tylus sp . : Woodbur y , 1962 .

Fam i ly PALMAECocos nucifer a L . : B r yan ,

1 944 ; St . John , 1950 ; Tay lor , 1950 ; Fosberg , 1 955 ; St . John , 1960 ; Palumbo , 1962; Woodbur y ,1962 ; Welander et al . , 1 966 ; Carpenter et al . , 1968 ; Fal l et ai . , 1 971 ; Kar anda et ai . , 1 973 ; Lamber son , 1982 .

Fam i ly AMARYLL IDACEAETCr inum as iat icum L . : B r yan , 1944 ; St . John , 1 960 ; Lamber son , 1 982 .

Pancrat ium littorale Jacquin : Lamber son , 1982 .

Fam i ly TACCACEAE .

TTacca leon topetalo ides (L ) O . Kuntze: Taylor , 1 950 ; Fosberg , 1 955 ; St . John , 1960 ; Welander , et ai . , 1966 ; Kor anda et ai . ,

1 973 ; Lamber son , 1982 .

Tacca p innat if i'da For ster : B r yan , 1 944 .

Subc las s DICOTYLEDONEAEFam i ly MORACEAEA r tocarpus alt i l is (Park inson) Fosberg : Fosber g et ai . , 1979 .

A r tocarpus incisus (Thunberg) L . f . : St . John , 1 960 .

TA r tocarp us mar iannens is T r ecu l : Fosberg et al . , 1 979

Fam i ly URT ICACEAETLapor tea r uderalis (For s ter f .) W . L . Chew : Fosberg et ai . , 1 979; Lamber son , 1 982 .

F leu rya r uderal is (For ster f . ) Gaudichaud: St . John , 1 950 ; Tay lor , 1950 ; Fosberg , 1 955 ; Lane, 1960 ; St . John ; 1 960 ;Welander et ai . , 1 966 .

Fam i ly OLACACEAETXimen ia amer icana L . : Tay lor , 1 950 ; Fosberg , 1 955 ; St . John ,

'1960 ; Fosberg et ai . , 1979.

Fam i ly POLYGANACEAECocco loba uvifera (L ) L . : Lam ber son , 1982 .

Fam i ly AMARANTH ACEAETAchyran thes aspe ra L . : Fosberg , 1 955 ; St . John , 1960 ; Fosberg et ai 1979.

TAchyran thes canescens R . B r own : St . John , 1 960 .

TAmaran thus dubius Mar tel]: Fosberg , 1 955 ; St . John , 1 960 ; Fosberg et ai . , 1 979 .

Amar an thus viri dis L . : Fosber g , 1 955 ; St . John , 1960 ; Fosberg et ai . , 1979 .

Fam i ly NYCTAGINACEAE“

( Boe r haviaalbiflor a var . powell iae Fosberg : Fosberg et ai . , 1 979 .

TBoer havia repens L .

Boer havia diffusa L . var . eudiffusa: St . John , 1 950 .

Boer havia diffusa L . : Tay lor , 1 950 ; Fosberg , 1 955 ; Lane, 1960 ; Welander et ai . , 1 966 ; Fall et ai . , 1 971 .

Boer havia diffuse L . var . diffusa: St . John , 1 960 .

‘

l’Boer havia tet r andr a For ster : Tay lor , 1950 ; Fosberg , 1 955 ; Palumbo , 1 962; Woodbur y , 1962 ; Welander et ai . , 1966 .

Boer havia diffusa var . tetrandra (For ster f . ) H eimer l : Br yan , 1 944 ; St . John , 1 960 .

Boer havia spp . : Lamber son , 1 982 .

TP ison ia gr andis R . Br own : Br yan , 1 944 ; Tay lor , 1 950 ; Fos berg , 1 955 ; Lane, 1960 ; St . John , 1960 ; Welander et ai . , 1 966 ;Carpen ter et ai . , 1 960 ; Fal l et ai . , 1 971 ; Koranda et ai . , 1 973 ; Koranda et ai . , 1978; Fosberg et ai . , 1979; Lamber son ,

1 982 .

Pison ia sp . : Woodbury , 1 962; Lee and Lee, 1 978 .

Fam i ly PORTULACACEAETPor tu laca aus tr a lis Endl . : Fosberg et ai . , 1 979.

Por tu laca quadr ifida L . : B ryan , 1 944 ; Tay lor , 1950 ; Welander et ai . , 1 966 .

Por tu laca samoens is v . Poelln . : Fosberg , 1 955: Lane, 1 960 ; St . John , 1 960 ; Lamber son, 1982 .

TPo r tu laca lutea So lander ex For ster f . : Tay lor , 1 950 ; Fosberg , 1 955 ; St . John , 1960 ; Welander et ai . , 1966 ; Fos berg ci a]1 979 ; Lamber son , 1 982 .

“

(Species cons idered indigenous to Micr ones ia. (Thi s table continued on next page

20 LAMBERSON

TABLE 1 (con t’d)

Fam i ly PORTULACACEAE (con t ’d)Por tu laca o ler acea L . : Br yan , 1944 ; St . John , 1 950 ; B iddulph and Cory , 1952 ; Fosberg , 1955 ; Lane, 1960 ; St . John , 1960 ;Palumbo , 1 962; Welander et ai . , 1 966 ; Fos berg et ai . , 1 979 ; Lamber son , 1982 .

Fam i ly LAURACEAECassytha f i l iform is L . : Br yan , 1944; Tay lor , 1 950 ; Fosberg , 1955; St . John , 1 960 ; Woodbury , 1 962; Welander et ai . , 1966 ;

Koranda et ai . , 1 973 ; Fosberg et ai . , 1 979 ; Lamber son , 1 982 .

TCythya f i liform is : Koranda et ai . , 1 978 .

Fam i ly FABACEAE“

(Canaval ia cathar t ica Thouar s : Fosberg et ai . , 1979 ; Lamber son , 1 982.

Canava lia m icrocarpa (DeCandolle) Piper : Fosber g , 1955 ; St . John , 1 960 ; Welander et al . , 1966 .

Desmod ium incanum DeCandolle.

Desmod ium adscendens (Sw ) DeCandolle: Lam ber son , 1 982 .

Leucaena leucocephala (Lamar ck) Dewi tz Lamber son , 1982 .

TVigna mar ina (Burmann ) Mer r i ll : B r yan , 1 944 ; St . John , 1960 .

Fam i ly ZYGOPH YL LACEAETr ibu lus cis to ides L . : Fosberg , 1 955 ; St . John , 1 960 ; Fosberg et al 1979 .

Fam i ly RUTACEAEllCitr u s au ran t ifol ia (Chr is tm ) Swing le.

Fam i ly SURIANACEAESur iana mar i t ima L . : Tay lor , 1 950 ; Fos berg , 1 955 ; Lane, 1 960 ; St . John , 1 960 ; Woodbury , 1962 ; Welander et ai . , 1966 ;

Kot anda et ai . , 1 973 ; Fosberg et ai . , 1 979 ; Lamber son , 1 982 .

Fam i ly SIMAROUBACEAE‘I'Sou lamea amar a Lamar ck : Welander et ai . , 1966 .

Fam i ly EUPH ORB IACEAETEuphor bia cham is son is (K lot sch and Gat oke) Bo iss ier : Fosber g , 1 955 ; St . John , 1 960 ; Fos berg et ai . , 1 979 ; Lamber son , 1982 .

Euphor bia ato to For ster : Welander et ai . , 1966 .

Euphor bia h ir ta L . : Fosberg , 1955 ; St . John , 1960 ; Welander et ai . , 1 966 ; Fosberg et ai . , 1 979 ; Lamber son , 1982 .

Euphor bia thym ifo l ia L . : St . John , 1960 ; Welander et ai . , 1966 ; Lamber son , 1 982 .

Phy l lan thus amarus Schum . and Thonn . : St . John , 1960 ; Fosberg et ai . , 1979 ; Lamber son , 1 982 .

R icinus commun is L . : Fosberg , 1955 ; St . John , 1 960 ; Fosber g et ai . , 1 979 .

Fam i ly T IL IACEAE‘l'Tr iumfetta pr ocum bens For ster : B r yan , 1 944; Tay lor , 1 950 ; Fosberg , 1955; Lane, 1 960 ; St . John , 1960 ; Palumbo , 1962;Woodbur y , 1962 ; Welander et ai . , 1 966 ; Carpenter et ai . , 1 968 ; Fal l et ai . , 1 971 ; Kot anda et ai . , 1973 ; Fosberg et ai . ,1 979 ; Lam ber son , 1 982 .

Fam i ly MALVACEAE“

( H ibiscus t i l iaceus L . : Lam ber son , 1 982 .

Ma lvas t rum co romandel ian um (L ) Gar cke: St . John , 1960 .

‘l‘Sida fa l lax Walper s : B r yan , 1 944 ; Tay lo r , 1950 ; Fosberg , 1 955 ; St . John , 1 960 ; Welander et ai . , 1 966 ; Kot anda et ai . , 1 973 ;

Fosberg et ai . , 1 979 ; Lam ber son , 1982 .

Fam i ly STERCUL IACEAEWalther ia indica L . : Lamber son , 1 982 .

Fam i ly LYTH RACEAE“

(Pemphis acidu la For s ter : Tay lor , 1 950 ; Fosber g , 1 955 ; Lane, 1 960 ; St . John , 1 960 ; Welander et ai . , 1 966 ; Kor anda et ai . ,

1 973 ; Fosberg et ai . , 1 979 ; Lamber son , 1 982 .

Pemphis sp . : Carpenter et ai . , 1 968 .

Fam i ly COMBRETACEAE‘l‘Term inalia samoens is Rechinger : Fosberg , 1 955 ; St . John , 1 960 ; Welander et ai . , 1 966 ; Fal l et ai . , 1 971 ; Fosberg et ai . ,

1 979 .

Term inalia litor a lis Seeman : Tay lor , 1 950 ; Lane, 1960 ; Lam ber son , 1 982 .

Term ina l ia sp . : Woodbur y , 1 962 .

Fam i ly LOGAN IACEAEPolyp rem um procum bens L . : Lam ber son , 1982 .

Fam i ly APOCYANACEAETNeisosperma Oppos it ifo l ia (Lamar ck) Fosber g and Sachet : Fosberg et al . , 1 979 ; Lamber son , 1 982 .

Ochr os ia par vif lora (For ster ) H ens low : B r yan , 1944 ; Tay lor , 1950 .

Ochr os ia oppos it ifo l ia (Lamarck) K . Schumann : Fosberg , 1 955 ; Lane, 1 960 ; St . John , 1960 ; Welander et ai . , 1966 ;Fal l et ai . , 1 971 ; Koranda et ai . , 1 973 .

Ochros ia sp . : Woodbury , 1 962 .

TSpec ies cons idered indigenous to Micrones ia.

llNew Enewetak record ; see Table 4 . (Th i s table continued on next page . )

H ISTORY OF TERRESTRIAL VASCULAR PLANTS 2 1

TABLE 1 (con t’d)

Fam i ly APOCYANACEAE (cont ’d)llPlumer ia obtusa L . , 1 753 .

llP lumer ia rubra L . , 1 753 .

Fam i ly CONVOLVULACEAETipomoea macr an tha Roemer and Schultes : Fosberg et ai 1979 ; Lamber son , 1 982 .

Ipomoea gr andif lor a (Cho isy) H al l ier : Bryan , 1 944 .

ipomoea tuba (Schlechtendahl) G . Don : St . John , 1950 ; B iddulph and B iddulph , 1 953 ; Fosberg , 1955 ; Lane, 1960 ; St .

John , 1 960 ; Woodbur y , 1 962 ; Welander et ai . , 1 966 .

ipomoea alba L . : Taylor , 1 950 .

Tlpomoea pes -caprae s sp . bras i lien s is (L ) v . Oos ts tr oom : St . John , 1950 ; Fosberg , 1 955 ; Lane, 1960 ; St . John , 1 960 ; Welanderet ai . , 1 966 ; Koranda et ai . , 1 973 ; Koranda et ai . , 1978 ; Fosberg ci ai . , 1 979 .

ipomoea bras i l iens is (L ) Sweet : Lamber son , 1 982 .

Ipomoea sp . : Carpenter et ai . , 1 968 ; Fal l et ai . , 1 971 .

Fam i ly BORAGINACEAETCordia subcordata Lamarck : Tay lor , 1950 ; Fosberg , 1 955 ; Lane, 1 960 ; St . John ,

1 960 ; Welander et ai . , 1 966 ; Fal l et ai . ,1 971 ; Koranda et ai . , 1 973 ; Fosberg et ai . , 1 979; Lamber son , 1982 .

Cordia sp . : Woodbury , 1 962; Car penter et ai . , 1968 .

TH el iotrop ium procum ben s var . depressum (Cham isso) Fosberg and Sachet .H eliot ropium anomalum H ooker and Ar nott : Welander et ai . , 1 966 ; Lamber son , 1982 .

Tour nefor t ia argen tea L . f . : Tay lor , 1 950 : Carpenter et ai . , 1 968 ; Fall et ai . , 1971 ; Fosberg et ai 1979 .

Mes ser s chm idia argen tea Johnston : B r yan , 1 944; St . John , 1950 ; Fosberg , 1 955 ; Lane, 1960 ; St . John , 1960 ;Woodbur y , 1 962 ; Palumbo , 1962 ; We lander et ai . , 1966 ; Jackson and Carpenter , 1967; Koranda et al . , 1 973 ; Koranda et

ai . , 1 978 ; Lamber son , 1 982 .

Mes ser schm idia sp . : Lee and Lee, 1 978 .

Fam i ly VERBENACEAETClerodendr um inerme var . ocean icum A . Gr ay : Fosberg et ai . , 1 979 .

L ipp ia nodiflor a (L ) R ich : Lamber son , 1 982 .

TP r em na obtus ifo lia R . B r own : Lam ber son , 1 982 .

Stachytarpheta jamaicens is (L ) Vahl .Stachytarpheta u r t icaefol ia (Sal isb) Sim s : Lamber son , 1 982 .

llVitex t r ifo lia L .

Fam i ly SOLANACEAEN icot iana g lauca Graham : St . John , 1 960 .

Physal is angu lata L . var . angu lata St . John , 1960 ; Welander et ai . , 1 966 ; Lamber son , 1 982.

Fam i ly ACANTH ACEAEPseuder an themum car r u ther s u (Seemann) G u i llaum in var . car r u ther s u: Lamber son , 1 982 .

Pseuderan themum car r u ther s u (Seemann) G u i llaumin var . at ropurpureum (Bul l ) Fosberg ; Lamber son , 1 982 .

Fam i ly RUB IACEAETG uettarda speciosa L . : Br yan , 1 944 ; St . John , 1950 ; Tay lor , 1 950 ; Lane, 1 960 ; St . John , 1 960 ; Palumbo , 1 962;

Welander et ai . , 1 966 ; Fall et al . , 1 971 ; Kor anda et ai . , 1 973 ; Fosberg et ai . , 1979; Lamber son , 1 982 .

G uettarda sp . : Woodbur y , 1 962 .

Mor inda cit r ifol ia L . : B r yan , 1 944 ; St . John , 1 950 ; Tay lor , 1 950 ; Fosberg , 1 955 ; St . John, 1 960 ; Welander et ai . , 1 966 ;Fal l et ai . , 1 971 ; Koranda et al . , 1 973 ; Fosber g et ai . , 1 979 ; Lamber son , 1 982 .

Mor inda sp . : Woodbur y , 1962 .

Fam i ly GOODEN IACEAEScaevo la taccada (Gaer tner ) Roxburgh : Lane, 1 960 ; Jackson and Carpenter , 1967 ; Carpenter et ai . , 1 968 ; Fal l c i ai . , 1 971 ;Fosberg et ai . , 1 979 ; Lamber son , 1 982 .

Scaevo la fr utescens (M i l ler ) Kr ause: B r yan , 1 944 ; St . John , 1 950 ; Taylor , 1 950 ; Fosberg , 1955 ; St . John , 1960 ;

Woodbur y , 1962 ; Welander et ai . , 1 966 ; Kor anda et ai . , 1 973 ; Kot anda et ai . , 1978.

Scaevo la ser iacea Vahl : Palumbo , 1 962 .

Scaevo la sp . : Lee and Lee, 1 978 .

Fam i ly COMPOSITAEB idens p i losa L . var . m inor (B l ) Sher ff : Lamber son , 1 982 .

Conyza bonar iens is (L . ) Cr onquist : Fosberg et ai . , 1 979 ; Lam ber son , 1 982 .

Er igeron bonar iens is L . : St . John , 1 960 .

Ecl ipta alba (L ) H asskar l : Lamber son , 1 982 .

TPluchea indica (L ) Less ing : St . John , 1 960 ; Fosberg et ai . , 1 979 ; Lamber son , 1 982.

“

(Species cons idered indigenous to M icr ones ia .

l lNew Enewetak record ; see Table 4 . (Th is table cont inued on next page . )

22 LAMBERSON

TABLE 1 (con t'd)

Fam i ly COMPOSITAE (cont ’d)P luchea symphyt ifol ia (M i l ler ) Gi ll is : Lamber son , 1 982 .

P luchea odorata (L . ) Cass in i : Fosberg , 1 955; St . John , 1 960 ; Welander et ai . , 1966 ; Koranda et ai . , 1 973 ;

Kor anda et ai . , 1 978 .

Tr idax pr ocumbens L . : Lamber son , 1982 .

Ver on ia cinerea (L ) Less ing : Fosberg , 1955; St . John , 1 960 ; Welander et ai . , 1 966 ; Fosberg , 1979 ; Lamber son , 1 982 .

Wedel ia tr i lobata (L ) H i tchcock : Fosberg et ai . , 1 979 ; Lamber son , 1 982 .

TWollas ton ia bif lor a (L ) DeCandolle.

Wedel ia biflo ra (L ) DeCandolle: Br yan , 1 944 ; Fosberg , 1 955 ; St . John , 1 960 ; Welander et al . , 1966 ; Lamber son , 1982.

TSpecies cons ider ed indigenous to M icr ones ia.

TABLE 2

Garden Plan ts Repo r t ed fr om Enewetak A tol l ,Con s ider ed to Be T r an s ien t Mem ber s of the F lor a

D iv is ion TRACH EOPH YTAClass ANGIOSPERMAESubc lass MONOCOTYLEDONEAEFam i ly GRAMINAESo rghum bico lor (L . ) Moench var . techn icum (Koer n ) Fior i and Pao l i , broom corn Fosberg , 1955 . So rghum vu lgare (L ) Per soon :Tay lor , 1 950 .

Zea mays L . ,

“cor n St . John , 1960 .

Fam i ly AGAVACEAE‘Cordy line term ina lis (L ) Kunth , “

t i .”

Fam i ly L IL IACEAEA l lium cepa L . , on ion B r yan , 1 944 ; St . John , 1 960 .

Fam i ly MUSACEAE‘Musa sap ien tum L . , banana .

Fam i ly ARACEAE‘Ep ipremnum p innatum (L ) Eng ler cv . Aur eum .

Subc lass DICOTYLEDONEAEFam i ly NYCTAG INACEAEM irabi lis jalapa L . ,

“ four -o ’clock Fosberg , 1 955; St John , 1 960 ; Fosberg et ai . , 1 979 .

Fam i ly CRUCIFERAEB ras s ica o leracea L . cap itata, cabbage B r yan , 1 944 ; St . John , 1 960 .

B rass ica pekinens is (Lour ) Rupr . ,

“

Ch inese cabbage”: B ryan , 1 944 ; St . John , 1 960 .

B ras s ica sp . : B r yan , 1944 ; Fosber g , 1955 ; St . John , 1 960 ; Fos berg et ai . , 1 979 .

Raphanus sat ivus L . ,

“

rad ish": B r yan , 1 944 ; St . John , 1 960 .

Fam i ly FABACEAEPhaseo lus sp . ,

“ beans B r yan , 1944 ; St . John , 1960 .

Fam i ly MALVACEAE‘H ibiscus hybr id (H . r osa—s inen s is x

“ h ibiscus .

Fam i ly CAR ICACEAECar ica papaya L . ,

“papaya St . John , 1 960 .

Fam i ly CONVOLVULACEAEipomoea purpurea (L ) Roth : St . John , 1 960 .

Fam i ly SOLANACEAESo lpnum lycoper s icum L . , tomato Br yan , 1944 ; Fosberg , 1955 ; St John , 1 960 ; Fosberg et ai . , 1 979 .

Fam i ly CUCURB ITACEAECit ru l lus lanatus var . caffror um (A lef ) Fosberg , watermelon Fosberg , et ai . , 1979 .

Cit ru l lus vu lgar is Schrader , “watermelon": Bryan , 1 944 ; Fosberg , 1 955 ; St . John , 1 960 .

Cucum is melo L . , canteloupe Fosber g , 1 955 ; St . John ,

'

1 960 ; Fosberg et ai . , 1979 .

Cucur bita max ima Duchesne,

“

squash": B r yan , 1 944 ; Fosber g , 1 955 ; St . John , 1 960 ; Fosberg et ai . , 1979.

Fam i ly COMPOSITAELactuca sat iva L . ,

“ lettuce Br yan , 1 944 ; St . John , 1 960 .

‘New Enewetak record ; see Table 4 .

H ISTORY OF TERRESTRIAL VASCULAR PLANTS

TABLE 3

Plan ts Repor ted fr om Enewetak A tol l bu t FoundOn ly as D r i ft Seeds on Beaches

Div is ion TRACH EOPH YTAClass ANGIOSPERMAESubc lass D ICOTYLEDONEAEFam i ly H ERNAND IACEAEH er nandia sonor a L . : St . John , 1960

Fam i ly FABACEAECaesalp ina bonduc (L ) Roxbury : St . John , 1960 .

D ioclea ref lexa H ook . f . : St . John , 1960

En tada phaseo lo ides (L ) Mer r i ll : St . John , 1 960 .

Mucuna urens (L ) Medicus : St . John , 1 960 .

Fam i ly EUPH ORB IACEAEA leur i tes moluccana (L ) Wi lldenow : St . John , 1960 .

Fami ly SAPINDACEAESap indus saponar ia L . : St . John , 1 960 .

TABLE 4

Col lect i on Da ta of N ew ly Recorded Ter r es tr i alVascu lar Plan ts fr om Enewetak‘

Fam i ly GRAMINAELep tur us r epens (For st . f . ) R . B rown var . s ubu latu s Fos berg .

Enewetak Is let : Mar . 1976 ; Enjebi i s let : co ll . byR . Warner (MPRL ) .

Fam i ly MUSACEAEMusa sap ien tum L . Enjebi Is let : P lanted in Lawrence L ivermoreLabor atory exper imental garden . Japtan Is let : P lanted nearhomes .

Not co llected .

Fam i ly ARACEAEEp ipremn

'um p innatum (L ) Eng ler cv . Aureum Enewetak Is let :Mar . 3 1 975, under ai r conditioner (MPRL ) .

Fam i ly AGAVACEAECor dy line term inalis (L ) Kunth . Enewetak Is let : Feb. 1 2, 1 975,

near bui ldings , co ll . by E . H . Br yan , Jr . (BPBM) .Fam i ly RUTACEAECi tr us au ran tifol ia (Chr isti n ) Swingle. Enjebi is let : P lanted inLawr ence L ivermore Labor atory exper imental garden .

Not co l lected .Fam i ly MALVACEAEH ibiscus sp . (hybr id, H . r osa-s inens is x

Enewetak Is let : Mar . 1 975, near chape l (MPRL ) .Fam i ly APOCYANACEAEP lumer ia obtusa L . Enewetak Is let : Feb . 1975, near

bui ldings (MPRL ) .P lumer ia rubra L . Enewetak Is let : Feb. 1 975, near bui ldings ,co ll . by E. H . Bryan , Jr . (BPBM) .

Fam i ly VERBENACEAEVitex tr ifo lia L . Enewetak Is let : Feb . 1975, near bui ldings atsouthwes t end of runway ; also co llected Feb. 1 2, 1 975 byE. H . Bryan , Jr . (MPRL , BPBM) .

‘Taxa are l isted in the same order as in Tables 1 thr ough 3 .

Specimens were co l lected by J . O . Lamber son , 1 975 thr ough1 977, except as noted, were determ ined by H . St . John and

F . R . Fosberg , and are depos i ted in the U . S. National MuseumH er bar ium (USNM ). the Bi shop Museum H er bar ium (BPBM), andthe M id-Paci fic Research La boratory Refer ence Co llecti on(MPRL ) .

23

the Pter idophyta cor relates w i th that g iven by Fosberg et

ai . (1 979,

Th i r ty-s ix spec ies of the total) are introducedweeds , and 30 spec ies ar e introduced or namen talsor food p lan ts . N ineteen of these species occur on ly undercult ivat ion and are cons idered tr ans ient mem ber s of the

flo r a (Table Seven spec ies have been r epor tedon ly as dr i ft seeds found on beaches (Table New

r ecor ds for the ato l l r epor ted in th is chapter (Table 4)br ing the total num ber of species recor ded from the ato l lto 1 28 .

Enewetak Ato l l averages 1 25 to 150 cm of rain annually and is much dr ier than ato l ls far ther south , which arecloser to the tr op ical convergence zone jus t nor th of theequator . Obser vat ions dur ing the per iod 1975 to 1 977

and meteor o log i ca l r ecor ds show that there is a defin i teseasonal i ty on Enewetak : dry and w indy fr om Novemberthr ough Apr i l , w i th more rain and less wind the other halfof the year .

The plants of Enewetak reflect th is seasonal i ty , producing mor e flower s and fr u i t in summer and shedding someor mos t of thei r leaves in w inter , depending on the

species . P ison ia for es ts become open and l ight in the

w inter because P ison ia tr ees shed mos t of thei r leaves .

Th is species flowered abundan tl y in Januar y and Febr uary1 976 , and the flower s wer e fol lowed by the s ticky fr u i tsand new growth of leaves in Mar ch and Apr i l . Term inal iatr ees shed almost all of thei r leaves , wh ich tur n yel low and

dr op in the dry season . Cordia, Tour nefor t ia, G uettarda,

and Scaevo la shed some of thei r leaves dur ing the dry sea

son and produce more flower s and fr u i t dur ing the summermon ths . P r emna and H ibiscus t i l iaceus shed many of thei rleaves in w inter , and new gr owth is produced in Apr i l orMay. Pemphis , Sur iana, Cocos , and Mor inda produceflower s and fr u i t year r ound , though the plants may

appear more w ind-blown and dry in w inter , and the seedsusual ly germ inate in ear ly summer .

Many of the ground plan ts and weedy species alsoreflect the seasonal i ty . Annuals such as Con yza, Physal is ,and some of the gr asses ar e seen mos tl y in summer , as

are Lapo r tea and Por tu laca. Ipomoea spec ies ar e pr esentyear r ound but flower in summer . Tacca d ies down in

October or Novem ber and gr ows fr om the under gr oundtuber s in Apr i l or May, pr oduc ing flower s in June and

seeds in July . Cyper us species flower in summer , althoughF imbr is tylus is in bloom year round . Euphor bia species arepr esen t year r ound , but Phyl lan thus i s , much moreabundant in summer . Walther ia also d ies down in winter ,and new gr owth is seen in Apr i l or May.

T r op ica l s torms , though uncommon in the Mar shal li s lands , do occas ionally occur in the v icin i ty of Enewetak,and s torm waves somet imes wash over the is lands . The

dense shr ub growth at the top of mos t of the i s landbeaches affords some pr otection to plants far ther inland ,but s torms may wash away par ts of the is lands or damagethe larger trees , espec ial ly P ison ia, wh ich is soft and eas i lybr oken . A typ hoon appr oached the atol l in the summer of1 977 , caus ing some damage to the P ison ia on the leewar d

24 LAMBERSON

i s lands and send ing s torm waves over the nor theas t end of

Enewetak Is land to a depth of more than 1 m .

H ISTORY AND PREVIOUSOBSERVATION S

Du r ing the late 1 8005 and ear ly 1 9005 , the people of

Enewetak Atol l cult ivated Tacca, br eadfr u i t , coconut ,Pandanus , and other plants on the ato l l and traded copr ato the Germans . Some of the coconut tr ees plan ted in neatrows du r ing German t imes s t i ll sur v ive and producecoconuts . German so i l used as bal las t in sh ips was de

pos i ted on the is land of Jap tan to pr omote coconutgr owth . Plants and plan t par ts wer e ut i l i zed extens ivel y inthe Mar shal l i s lands for food , dr ink , bu i ld ing mater ials (forhouses and canoes) , utens i ls , cloth ing , toys , medic ine, and

other uses (Anon , 1951 ; B r yan ,

in 1 91 4 ,Japan acqu i r ed Enewetak as a r esult of a

Wor ld War l mandate, and a Japanese tr ader and hi s as s istants r es ided on the ato ll . The fi r s t known botan ical co l leet ion from Enewetak was one of Pandanus made by Ohbain 1 934 and publ ished by Kaneh i ra In the ear ly1 9405 , the Japanese cleared par ts of some of the is landsfor cons tr uct ion of m i l i tar y fo r t i ficat ions , includ ing an ai r

field on Enjebi .In 1 944 , the atol l was attacked and captur ed by

Un i ted States for ces . The heavy shel l ing dur ing the batt leand subsequent clear ing and gr ad ing r esulted in large-

scaledes tr uct ion of the vegetat ion on Enjebi , Medr en (thencal led Par r y) , and Enewetak i s lands . B r yan ( 1 944)descr ibed the atol l ’s vegetat ion at that t ime and noted thatMedr en resembled a plowed field w i th almos t no plan ts at

all . B r yan ’

s r epor t and 1 954 Army Map Ser v ice mapsbased on aer ial photog raphs taken in 1 944 ind icate, how

ever , that some coconut trees were st i l l pr esen t on sever alis lands . in 1 946 , Fosberg v is i ted the ato l l and r epo r ted ther esults of his botan ical sur vey and co l lect ion of Enewetakplan ts w i th those from o ther ato l ls in the Mar shal l i s lands(Fosber g , Taylor (1 950) descr ibed the mar ine and

ter r es tr ial vegetat ion and the plant spec ies he encoun ter edwhen he v is ited the ato ll in 1 946 . At that t ime, P ison ia

had been cut on lku r en , Jedr ol , and Alembel ; m i l i tar yact iv i ty had also des troyed some of the vegetat ion on

Lujor , Alembel , Run i t , Japtan , and Jedrol ; and the is landsof Enewetak , Med i en , and Enjebi had been reduced to“was tes of wh i te sand between bu i ld ings w i th a few

weeds'

and scr ubby coconuts . Run i t , however , was com

p letely occup ied by a formal coconut grove, except fo r a

m i l i tar y ins tal lat ion at the south end.

Aer ial photogr aphs taken in 1 956 show that the flor aof almos t all the is lands was affected by m i l i tar y act iv i ty .

dur ing the nuclear tes t pr og r am conducted fr om 1 948 to

1958 . The i s lands of L ouj thr ough Boken (i rene) wer ealmos t bare, as wer e Run i t , Boko , and Bokandr etok .

Enewetak , Medr en , and Enjebi wer e covered w i th bu i ldings , and on ly a few plan ts grew on these i s lands amongthe s tr uctur es and r oadways . Mos t of the other is lands sup

por ted scr ub vegetat ion or were par t iall y cleared , and on lythe leewar d i s lands fr om Iku ren thr ough B iken wer e r elat ively und is tur bed .

The effects of the nuclear tes t prog ram on the b iotaand the env i r onment at Enewetak and B ik in i wereextens ively s tud ied by the Appl ied F i sher ies Labor ator y ofthe Un iver s i ty of Wash ington , later r enamed the Laborator y of Rad iat ion B iology . Thei r r esults were r epor ted inUWFL ser ies , in publ i shed paper s , and in jour nali s t ic formin the book P r oving Gr ound (H ines ,

An ar t icle by St . John ( 1 960) pr ov ides a comprehen

s ive r ecor d of the land p lants found on the atol l up to thatt ime . i t g ives a sys temat ic account of spec ies r ecor ded ,w i th notes on is lands , co l lector s , and dates . A lso includedar e a key to spec ies , index to genera , and other data helpful to an under s tand ing of the flor a of the ato l l .

The long-term effects on the vegetat ion because of the

nuclear tes t pr ogr am were pr imar i l y those r esult ing fr omdes tr uct ion of habi tats and the r emoval of so i l and vegetat ion fr om the i s lands (St . John , 1 950 ; B iddulph and Cor y ,1 952 ; Palumbo , Some abnormal i t ies , includ ingtumor s , calc ium deficiency , gr owth i r r egular i ties , abnormalfr u i ts and flower s , co lor abnormal i t ies , and s ter i l i ty , werefound in plants near tes t s i tes (St . John ,

1 950 ; B iddulph ,1 950 ; B iddulph and Cor y , 1 952 ; B iddu lph and B iddulph ,

H owever , hab i tat d is tur bance and r emoval of theso i l w i th loss of o rgan ic matter , inorgan ic nutr ients , and

water retent ion capab i l i ty appar ently had more las tingeffects on the vegetat ion than the rad ioact iv i ty . W i th thedes tr uct ion of pr otect ive hab i tats , exposur e to adver seenv i ronmental factor s such as salt spr ay and dr y ing w indswas gr eater .

Fo l low ing the conclus ion of nuclear tes t ing on

Enewetak in 1 958 , the ato ll was used for var ious otherm i l i tar y and sem i -m i l i tar y pu rposes dur ing the 1 9605 and

ear ly 1 9705 . The Pac i fic Cr ater ing Exper iment (PACE) pr ogram bulldozed Aomon to the water table level anddetonated nonnuclear explos ives wh ich left several smal lcr ater s on the is land .

The Enewetak Mar ine B io logical Labo r ator y (EMBL )was es tabl ished in 1 954 . An extens ive r efer ence col lect ionof ato l l biota,

includ ing mar ine and ter res tr ial p lants , hasbeen gr adual ly gathered , and the labor ator y has pr ov idedfac i l i t ies for a lar ge number of v is i t ing scient is ts . Spec imensfor the r efer ence co l lect ion wer e con tr ibuted byG . R . Baker , E . H . B r yan , P . Co l invaux , V . Fr ey ,M

.Gi lmar t in ,

J . Lam ber son , P . Lam ber son , l . E . Lane,

M . A . Lee, H . St . John , and R . War ner . Some botan ica ls tud ies have been made through the laborator y and are

descr ibed in the labor ator y annual repor ts (B r yan and Lee,

1 975 ; Lee and Lee,

In 1 957 to 1 958 , the Un iver s i ty of H awai i , the U . S.

Weather Bur eau , and the Jo in t Task For ce Seven con

ducted m icr ocl imat ic weather obser vat ions at Enewetak .Dur ing that s tudy , Lane made botan ical co l lect ions and

no tes and con tr ibuted a detai led section , wh ich is includedin that r epor t , on the vegetat ion of some of the is lands(Lane,

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H ISTORY OF TERRESTR IAL VASCULAR PLANTS 27

plan ts such as Por tulaca, Boer havia, Lapo r tea, and Sida

appear . P ison ia may become a component of the youngm ixed for es t , and v ines ar e abundant . Cas sytha may helpto open up the for es t by k i l l ing some tr ees . Lar ger plantspr esent include Tour nefor tia, Scaevo la, Cocos , Pandanus ,

Cordia, Term ina lia, Mor inda, G uettarda,Pison ia, Sur iana,

and Pemphis . Th is s tage was found on the m iddle sectionof lku r en , on Mut , on K idrenen , and on the ocean s ide of

B iken . It was also seen on Anan ij, Van , the cen tral por t ionof Alembel , and Bokenelab (F ig .

F ig 3,

Stage 111, young m ixed for es t. Plan ts shown include Scaevola, Tour nefort ia, Guet tarda, Pemph is , andLep tu r us .

28 LAMBERSON

In Stage IV , P ison ia and Cocos dominate the o lderm ixed fores t , and Scaevola is near ly shaded ou t . Mor inda,

Tour nefor t ia, G uettarda, and Term inal ia reach for sun , and

Neisosperma may appear . The who le fores t i s mo re open ,

and the undergr owt h cons is ts of smal l Cocos and P ison iaw i th . gr ound plan ts of Lapor tea, Boer havia, Lep tu r us , and

Por tu laca. Sida and Scaevo la are r es tr icted to open ar eas ,

Sur iana and Pemphis occu r mos tly at the for es t margins ,and v ines become fewer , usual ly grow ing on ly in the sun

in clear ings and at the outer marg ins . Th i s s tage pr edominated on the main por t ions of Iku ren and Anan ij

In Stage V , P ison ia takes over . Cocos trees becomeunproductive and gr adual ly die ou t ; other species of treesar e confined to the edges of the for es t . There is l i tt le

F i g . 4 Stage W , older m ixed for es t. Plan ts i l lus tr ated ar e pr edom inan tl y Cocos and P ison ia.

H ISTORY OF TERRESTRIAL VASCULAR PLANTS 29

gr ound cover other than Pison ia seedl ings and r egener ated Stage III and common in Stage IV , become less abundant .branches . Nes t ing noddy ter ns fi l l the br anches of the Stage V was seen on B iken and Kidrenen (F ig .

P ison ia tr ees , and the for es t is open and easy to walk Por tions of Japtan wer e between Stages IV and Vthr ough . Coconut cr abs , wh ich are often present in

F i g . 5 Stage V ,P ison ia for es t. On l y P ison ia i s seen i n th i s photogr aph .

30 LAMBERSON

TH E ISLANDS. 1975 TO 1 977

Between Januar y 1 975 and May 1977, the authorv is i ted mos t of the atol l is lands to recor d vegetat ive data.

An i s land-by-is land summar y of plant species co l lected orobser ved is pr esented in Table 5 .

The nor thern.

is lands fr om Bokoluo to Boken wer eextens ively dis tur bed by the nuclear tes t prog r am . Theseis lands had an open vegetat ion of s cattered Scaevo la and

Toum efor t ia to 6 m in height wi th some Cordia, Guettarda, Cocos , and Pandanus wi th v ines ; a gr ound cover ofLep tu r us and Fimbr is tyl is ; and some Boerhavia, Cas s ytha,Tr iumfetta, Sida, ipomoea, and Euphorbia (Stages Ithrough II I) .

Th i r ty-four plan t species , other than those p lanted inthe LLL gar den pr oject , were found on Enjebi . The general vegetat ion (was open to dense s cr ub of Toum efor t iaand Scaevo la to,6 m in height , w i th occas ional Pandanus ,Pemphis , M

'

oii ri da, and Cocos (mod i fied Stage Ii to I I I) .

Cenchr us (repor ted ext inct on th is is land in

Lep tu r u s , F imbr is tyl is , Cassytha, Euphor bia spp . , Phyllan thus , i pomoea spp . , P luchea spp . , Stachytarpheta,

H eliotr op ium , and Tr idax were common , especial ly in the

LLL gar den . Th is is land was once planted wi th coconutsand was the home of the people of Enjebi , but the coconutplantation was des tr oyed by Japanese and Amer ican m i l itar y act iv i t ies . Ten nuclear tes ts were conducted on or

near the i s land , and du r ing the tes t days the vegetat ioncon s is ted of on ly gr ass , v ines , and low shrubs . By 1964,

however , Toum efor t ia tr ees wer e up to 6 m in height , andScaevo la almos t to 5 m (Welander et ai . , SomePison ia was r epor ted in 1 946 (Fosberg , 1955) and 1 964

(Welander et ai . , 1966) but was not seen in 1975 to

1 977 .

The is lands of M ijikadr ek thr ough Lujor suppor ted on l ya scr ub vegetat ion , cons is t ing predominately of Scaevo laand Tour nefor tia (Stage I I to II I) . Coconut tr ees werepr esen t on Bokenelab; a few sprouts had appear ed on

Lujor ; and a grove of coconut trees w i th P ison ia occur r edon Elle. P ison ia was also pr esent on M ijikadr ek , and i t s

s t icky fr u i ts may even tual l y be tr anspo r ted by b i r ds fr omther e to other nor ther n is lands . Lep tu r us , Cas s ytha, F im

br is tylus , Tr iumfetta, Sida, and Ipomoea wer e fai r ly common , and there was some G uettarda . Lujor had muchMor inda, and El le had a gr ove of Term inal ia on the southend, ocean s ide. The is land of Taiwei or Per cy , betweenK idrenen and Bokenelab, has been bar r en at leas t s ince1 944 , and s torm waves somet imes wash over i t . It has a

bo r der of beachr ock and may once have been vegetated ,but the on ly ev idence of plant l i fe seen there fr om 1 975 to1977 was a spr outed coconut wh ich had d ied .

The is lands of Eleler on thr ough B iji le were once jo inedby causeways , though Eleler on was d iv ided by a nucleartes t in 1 956 , and par t of i t is now separate, although par ti s s t i l l jo ined to Aomon by a causeway . Aomon was

bulldozed and pockmar ked w i th smal l cr ater s dur ing the

PACE progr am . Ther e was an ai r s tr ip on B iji le,which was

joined by a br idge to Lojwa, the s i te of the nor ther n camp

dur ing the 1 977 and 1978 cleanup oper ations . All of theseis lands had a mixed scr ub vegetation of Tour nefor t ia and

Scaevo la, w i th some Cocos , Pandanus , P ison ia (on B iji le),Sur iana, Mor inda, and Term inal ia (on Aomon ), Cordia, and

G uet tarda (Stage I I to II I) . Pluchea was common , as wereLep tu r us , F imbr is tylus , Euphor bia, Tr iumfetta, ipomoea,

and H el iot rop ium . Stachytar pheta, L ipp ia, Eragr os t is , Por tu

laca, Cass ytha, and Sida wer e pr esent but uncommon .

Aomon was occupied by the Mar shal lese peop le dur ing thewar and pos twar per iod , and in 1977 Tacca s t i l l gr ew on

the ocean s ide, near s tands of H ibis cus t i l iaceus and

Wollas ton ia bif lor a. In 1977, wedge-tai led shearwater swer e found nes t ing in bur r ows near P ison ia tr ees on B iji le,

and Lapor tea and Tacca wer e seen near the nes t ing ar ea .

Alembel was r elatively undis tur bed and had a cen tr alm ixed for es t of P ison ia, Cocos , Scaevo la, Tour nefo r tia,P remna, Mo r inda, and G uettarda (Stage II I) . Cordia Tvaspresent on the lagoon s ide, and two large Pandanustrees —

one bear ing fr u i t— wer e at the nor ther nmos t end.

The outer margin was a th ick scr ub growth , pr imar i ly ofTour nefor t ia, G uettarda, and Scaevo la, w i th scr ub P ison iaand Pemphis on the ocean s ide, and Sur iana on the lagoons ide (Stage II) .

Run i t , wh ich was extens ively di s tur bed du r ing the

nuclear tes t prog r am and i s now a depos i tor y for radioac

t ive debr i s fr om other is lands , had an open vegetat ion of

scatter ed large Tour nefo r tia and Scaevo la bushes and a few

coconut tr ees (mod i fied Stage I I) . There was a gr oundcover of Cass ytha, Conyza, H e l iotrop ium, Lep tu r us , F im

br is tylis , Eup hor bia, Tr iumfetta, and ipomoea pescap r ae. A

few ind iv iduals of Sur iana, Pemphis , Mor inda, G uettarda,

and Pluchea wer e also seen . Th is i s land was bulldozeddur ing the nuclear tes t days and was the s i te of 1 7 tes ts ,includ ing Lacr osse (1 956) and Cactus ( 1 958) wh ich leftfai r ly large cr ater s at the nor th end.

The smal l is lands of Boko , Mun jor , and Inedr ol sup

por ted a l im i ted vegetat ion (Stage I) . Inedrol had on l yLep tu r us and Tour nefor tia, and the other s had thosespecies plus Scaevola. Inedr al and Van had mor e species ,includ ing P ison ia,

Boerhavia,ipomoea macr an tha, and

G uettarda. Inedral also had Sur iana, and Van had a youngm ixed for es t (Stage I I I) of coconut , Tr iumfetta, Cordia, and

Canava lia.

Anan ij had a Cocos — P ison ia for es t (Stage IV) in the

m iddle of the w ide po r t ion of the is land , w i th Lep tu r us andBoer havia beneath the tr ees and Scaevo la and Tour nefor tiaar ound the edges of the old r unway (Stage II) . There wereseveral Term inalia tr ees , p lus a few Cordia, Sur iana, and

G uettarda. The conso l idated r ock bar extending oceanwar dper pend icular to the is land was covered w i th Pemphis .

Jin im i is an unusual l i tt le is land wh ich was coveredw i th a dense mat of Boer havia w i th s cattered scr ubP ison ia, Tour nefor tia, and Scaevo la bushes . Por tu laca and

F im br is tylis wer e also common . The is land was somet imesthe s i te of a large nes t ing co lony of sooty ter ns , wh ichwere also found nes t ing on Aomon .

Japtan was par t ial ly clear ed du r ing the war and tes tdays and in 1 977 became the home of about 80

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30 LAMBERSON

TH E ISLANDS, 1975 TO 1977

Between Januar y 1975 and May 1977 , the authorv is i ted mos t of the atol l is lands to recor d vegetat ive data.

An is land-by-is land summar y of p lant species col lected orobser ved is presented in Table 5 .

The nor t hern i s lands fr om Bokoluo to Boken wereextens ivel y di s tur bed by the nuclear tes t program . Theseis lands had an open vegetation of scat tered Scaevo la and

Tour nefor ti a to 6 m in height wi th some Cordia,Guet

tarda, Cocos , and Pandanus wi th v ines ; a ground cover of

Lep tu r u s and Fimbr is tyl is ; and some Boerhavia, Cass ytha,

Tri umfetta, Sida, i pomoea, and Euphor bia (Stages Ithrough I I I) .

Th i r ty-four plant species , other than those plan ted inthe LLL gar den project , were found on Enjebi . The general vegetat ion (was open to dense s cr u b of Tour nefor tiaand Scaevo la to ;6 m in height , w i th occas ional Pandanus ,Pemphis , Maf inda, and Cocos (mod i fied Stage II to I II) .Cenchr u s (repor ted ext inct on th is i s land in

Lep tu r us , F imbr is tyl is , Cassytha, Euphor bia spp . , Phyllan thus

,ipomoea spp . , P luchea spp . , Stachytarpheta,

H e l iotr op ium , and Tr idax were common , especial ly in the

LLL gar den . Thi s i s land was once p lanted w i th coconutsand was the home of the peop le of Enjebi , but the coconutplantat ion was des tr oyed by Japanese and Amer i can mi l itar y activ i ties . Ten nuclear tes ts wer e conducted on or

near the i s land , and du r ing the tes t days the vegetationcons is ted of on ly gr ass , v ines , and low shr ubs . By 1 964 ,

however , Tour nefor tia tr ees were up to 6 m in height , andScaevo la almos t to 5 m (Welander et ai . , SomeP ison ia was r epor ted in 1 946 (Fosberg , 1955) and 1 964

(Welander et ai . , 1966) but was not seen in 1975 to

1977 .

The is lands of M ijikadr ek through Lujor suppor ted on lya scr ub vegetat ion , cons is ting predom inately of Scaevo laand Tour nefor tia (Stage Ii to I I I) . Coconut tr ees werepresent on Bokenelab; a few sprouts had appeared on

L ujor ; and a grove of coconut tr ees w i th Pison ia occur r edon El le. P ison ia was also presen t on M ijikadr ek , and i t s

s ticky fr u i ts may even tual ly be transpor ted by bi r ds fr omther e to other nor ther n is lands . Lep tur us , Cassytha,

F im

br is tylus , Tr iumfetta, Sida, and Ipomoea wer e fai r ly com

mon , and ther e was some G uettarda. Lujor had muchMor inda, and El le had a grove of Term inal ia on the southend, ocean s ide. The is land of Taiwel or Per cy , betweenK idr enen and Bokenelab, has been bar ren at leas t s ince1 944 , and s torm waves somet imes wash over i t . It has a

bor der of beachr ock and may once have been vegetated ,but the on ly ev idence of plant li fe seen there from 1 975 to1977 was a sprouted coconut wh ich had died .

The is lands of Eleleron thr ough B iji le wer e once jo inedby causeways , though Eleler on was d iv ided by a nucleartes t in 1956 , and par t of i t is now separate, although par tis s t i l l jo ined to Acmon by a causeway . Aomon was

bulldozed and pockmar ked w i th smal l crater s dur ing the

PACE prog ram . There was an ai r s tr ip on B iji le,wh ich was

jo ined by a br idge to Lojwa, the s i te of the nor ther n camp

du r ing the 1 977 and 1978 cleanup operat ions . A ll of theseis lands had a m ixed scr ub vegetation of Tour nefor tia and

Scaevo la, w i th some Cocos , Pandanus , P ison ia (on B iji le) ,Sur iana, Mor inda

, and Term inal ia (on Aomon) , Cordia, and

G uettarda (Stage I I to Il l) . Pluchea was common , as wer eLep tu r us , F imbr is tylus , Euphor bia, Tr iumfetta, ipomoea,

and H e l iotr op ium . Stachytarpheta, L ipp ia, E rag r os tis , Por tulaca, Cas s ytha, and Sida wer e pr esen t but uncommon .

Aomon was occup ied by the Mar shal lese people dur ing thewar and pos twar per iod , and in 1977 Tacca s t i l l grew on

the ocean s ide, near s tands of H ibis cus t i l iaceus and

Wo llas ton ia biflor a . In 1 977, wedge-tai led shearwater swere found nes t ing in bur r ows near P ison ia t r ees on B iji le,

and Lapor tea and Tacca wer e seen near the nes t ing ar ea .

Alembel was r elat ively undis tur bed and had a centr alm ixed fores t of P ison ia, Cocos , Scaevo la, Tour nefor tia,P remna

,Mor inda, and G uettarda (Stage Cordia Twas

pr esen t on the lagoon s ide, and two large Pandanustrees —

one bear ing fr u i t — wer e at the nor ther nmos t end.

The outer mar g in was a th ick scr ub gr owth , pr imar i ly ofTour nefo r t ia, G uettarda, and Scaevo la, w i th scr ub P ison iaand Pemph is on the ocean s ide, and Sur iana on the lagoons ide (Stage l l ) .

Ru n i t , wh ich was extens ively dis tur bed dur ing the

nuclear tes t prog r am and is now a depos i tor y for r adioac

t ive debr i s fr om other i s lands , had an open vegetat ion of

scatter ed lar ge Tou r nefor tia and Scaevo la bushes and a few

coconut tr ees (mod i fied Stage Ii ). There was a gr oundcover of Cas s ytha, Con yza,

H eliotr op ium , Lep tu r us , F im

br is tylis , Eup hor bia, Tr iumfetta, and Ipomoea pescap rae. A

few ind iv iduals of Sur iana, Pemphis , Mor inda, G uettarda,

and P luchea wer e also seen . Th is is land was bulldozeddur ing the nuclear tes t days and was the s i te of 1 7 tes ts ,includ ing Lacr osse (1 956) and Cactus (1 958) which leftfai r ly large crater s at the nor th end.

The smal l is lands of Boko, Munjor , and Inedrol sup

po r ted a l im i ted vegetat ion (Stage I) . inedr ol had on lyLep tu r us and Tour nefor tia, and the other s had thosespecies plus Scaevo la . lnedr al and Van had more species ,

includ ing P ison ia,Boer havia, Ipomoea macr an tha, and

G uettarda. lnedr al also had Sur iana, and Van had a youngm ixed for es t (Stage III) of coconut , Tr iumfetta, Cordia, and

Canava l ia.

Anan ij had a Cocos — P ison ia fores t (Stage IV) in the

m iddle of the w ide por t ion of the is land , w i th Lep tu r us and

Boer havia beneath the tr ees and Scaevo la and Tour nefor t iaaround the edges of the old r unway (Stage II) . Ther e wer esever al Term inalia trees , plus a few Cordia, Sur iana, and

G uettarda. The conso l idated rock bar extend ing oceanwar dper pend icular to the is land was cover ed w i th Pemph is .

Jin im i is an unusual l i ttle is land wh ich was cover edw i th a dense mat of Boer havia w i th scatter ed scr ubP ison ia, Tour nefo r t ia, and Scaevo la bushes . Por tulaca and

F imbr is ty l is were also common . The is land was somet imesthe s i te of a lar ge nes t ing colon y of soo ty ter ns , wh ichwer e also found nes t ing on Aomon .

Japtan was par t ial l y clear ed dur ing the war and tes tdays and in 1 977 became the home of about 80

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32 LAMBERSON

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34 LAMBERSON

Mar shal lese people. I t had a pr oduct ive coconut plan tat ion(es tab l ished dur ing the t ime of the German occupat ion )wh ich was somewhat overgr own and a field of Tacca. The

ocean s ide of the i s land was over grown w i th scr ubScaevo la and Tour nefor t ia (Stage I I) , and the clear ed areahad a gr ound cover of Cench r us and other grasses , sedges ,Euphor bia spp . , Physalis , Tr iumfetta, Por tu laca, Phy llan thus , Con yza, Ipomoea, Stachytarpheta, H e l io tr op ium ,

and Tr idax . P luchea was notably absent . The r es iden tsclear ed the under br ush from the area around the bu i ldingsand plan ted Pandanus and bananas . Ther e was also a

mi xed fores t on the southwes t th i r d of the i s land dominated by P ison ia, w i th N eisosperma, Cocos , Mor inda,

G uettarda, Co rdia, and Pandanus , w i th Boer havia near the

marg ins (Stage IV to V) .Jedr ol , a smal l is land ins ide the deep pass , set back

fr om the atol l r im , had a mature P ison ia for es t w i th no

Cocos but w i th Scaevola, Tour nefor t ia,Term inal ia, and

G uettarda (Stage IV to V) . F imbr is ty l is , Lep tu r us , and

Ipomoea macran tha were common , and Por tu laca and

Boer havia wer e pr esen t but less common . Some so i l wasr emoved fr om th i s i s land in 1 975 for the Enjebi gar den .

Medr en and Enewetak wer e completely clear ed dur ingthe war and nuclear tes t days for cons tr uct ion of hous ingand other bu i ld ings . Medr en was abandoned in 1 969 , and

the bu i ld ings and equ ipmen t left ther e became overgr ownw i th plan ts (mod i fied Stage I I) . Large shr ubs and tr ees of

Tour nefor t ia, Scaevo la, and Cocos were gr ow ing , and

Pluchea spp . were abundant . Ther e wer e remains of gar

dens contain ing Cocco loba, Cr inum, and Leucaena. Ther ewas a gr ound cover of var ious gr asses (not includ ingCench r us ) , sedges , Port u laca, Cas s ytha, Eupho r bia spp . ,

Phy l lan thus , Tr iumfetta, and ipomoea and in tr oduced species such as Tr idax , L ipp ia, Stachytarpheta, Con yza, Ver

non ia,H el iot r op ium , and Walther ia. Enewetak was less

over grown because i t had been con t inual l y occup ied buthad even mor e intr oduced spec ies , includ ing the weedsPolyp rem um , Desmod ium , and B idens . Or namen tals in

cluded Cr inum , Wedel ia, Pseuder an themum , Cocco loba,

Plumer ia, and H ibiscus , and gar dens con tained many mor etr ans ien t spec ies of food plants and flower s not r ecor dedfor th is r epor t .

Bokandr etok i s a smal l i s let jus t nor th of Enewetakwh ich may be r eached fr om there on foo t at low tide. Itwas bar ren and occupied by tr oops in the 1 9505 but in1977 suppor ted a dense cover of Scaevo la and Tournefor t ia ,

w i th Lep tu r us and Tr iumfetta ar ound the edges(Stage II) .

The leewar d is lands , Iku r en through R ibewon , were li tt le affected by the war and tes t days and were not

invo lved in the c leanup and r ehab i l i tation . There was a

weather s tat ion on Kidr enen , and there wer e a few derel ictbar ges and boats left along the s hor es , but the vegetat ionin 1 977 was Stage I I I to IV m ixed for es t . Coconuts wer eabundant on lku r en , Mut , and K idr enen , but those on Mutwer e old and seemed to be dy ing ou t (Stage IV to V) .P ison ia tr ees wer e pr esen t on all of the leewar d is lands ,and ther e wer e some near l y pur e s tands of th i s spec ies

ACKNOWLEDGMEN TSTh is r epor t i s based in par t on obser vat ions made by

the author du r ing employmen t at the M id-Pac i fic Mar ineLabor ator y fr om Januar y 1 975 thr ough May 1 977 .

Spec ies of p lants r epor ted as new r ecor ds for Enewetakwere iden t i fied by H aro ld St . John of the Ber n ice P .

B i shop Museum , H ono lulu , H awai i , and by F . R . Fosber gof the Nat ional Museum of Natu r al H is tor y . The M id

Pac i fic Resear ch Laborator y is suppor ted by the D iv is ionof B iomed ical and Env i r onmen ta l Resear ch of the Depar tmen t of Ener gy under Con tract No . AT (26- 1 )-628 to the

U n iver s i ty of H awai i . The autho r ’

s s incere thanks are

extended to Edw in H . Br yan , Jr . , H ar old St . John , and

F . R . Fosber g fo r botan ical informat ion and help in ident if icat ion , for thei r interes t and suppor t dur ing the per iod ofobser vat ion , and in the wr i t ing of th i s r epor t . Appr ec iat ionalso i s extended to S. H . Sohmer and Dar r el H er bs t formany helpful commen ts in thei r r ev iew of the manuscr iptand to Peggy Gaines for typ ing i t .

(Stage V) and of Pemphis on R ibewon and Boken . The

success ional s tages des cr ibed ear l ier were wel l i l lus tr atedon these is lands .

B iken , iso lated 20 km along the reef southwes t ofBokoluo , in 1 977 had a for es t dom inated by P ison ia w i thsome old coconut tr ees and open under neath (Stage V) .Ther e was a scr ub Scaevo la— Tour nefor t ia bo r der along thesouth s ide,

w i th a few G uettarda and Mo r inda . Cench r us ,Sida, and Ipomoea wer e pr esen t but uncommon on the

lagoon s ide, and on the nor th s ide ther e were Pandanusand a gr oup of Term inalia tr ees . A gr ove of old Cor diatr ees was found at the en tr ance to an over gr own road onthe nor thwes t s ide. The vegetat ion on th is is land was damaged by Oak tes t to the nor th in 1958 . It was descr ibed in1 964 as th ick and overgr own w i th Ipomoea macr an thamak ing tr ans i t d i ff icult (Welander et ai . , but in1 975 thr ough 1 977 i t was open and easy to walk thr ough .ipomoea and Cas sytha wer e s t i l l present but not abundant .The gr ound cover outs ide the P ison ia ar ea includedBoe r havia, Lep tu r us , and some Por tu laca and Tr iumfetta .

Ther e wer e a num ber of small cr ater s on the is land and

r eef flat dat ing fr om Wor ld War I I when the i s land wasused as a tar get for bomb ing pr act ice . Coconut cr abs wer eabundant , and r oos t ing immatu r e fr igate b i rds were oftenseen , as wer e many other spec ies of bi rds .

REFERENCES

Anonymous , 1 95 1 , The Agr iculture of Ar no Ato l l , Mar shal lIs lands , A to l l Res . Bul l . NO . 6, Pacific Science Board, Washlugton , D . C .

B iddu lph , O . , 1 950 , Sect ion VI of Radiobio log ical Sur vey of B ikin i ,En iwetok and L ikiep A to l ls , July— August , 1 949 , UWFL - 23 ,

Appl . Fish . Lab . Un iv . Wash .and R . Cor y , 1 952 , The Relat ionsh ip Between C0 45 , TotalCalcium and F is s ion Product Radioact iv i ty in P lan ts of Por tulaca o ler acea Growing in the Vicin ity of the A tom Bomb Tes t

Si tes on En iwetok A to l l , UWFL — 3 1 , Appl . F ish . Lab . Un iv .

Wash .

H ISTORY OF TERRESTRIAL VASCULAR PLANTS

B iddulph , S. F . , and O . B iddu lph , 1953 , A Descr ipt ion of Tumo r s

on Ipomoea Tuba fr om the A -Bomb Tes t Si tes on En iwetok

Ato l l, UWFL — 23 (App ) , Appl . Fish . Lab . U n iv . Wash .B r yan , E . H . , Jr . , 1 944 , Revegetat ion of Cer tain Mar shal l Is lands ,

u npubl . repor t , Cent . Pac . Base Comm .

1 972 , L ife in the Mar shal l Is lands , Pacific Science Informat ion Center , Ber n ice P . B i shop Museum , H ono lulu .

and M . A . Lee , 1 975 , Sur vey of Ter restr ial P lants on

Enewetak Ato l l , M id-Pacific Mar ine Labo r ator y Annua l Repo r t ,1 974— 1 975 , NVO O628

-T l l .

Buddemeier , R . W . , S. V . Sm ith , and R . A . Kinz ie, 1 975 , H o locene W indward Reef-Flat H istor y , Enewetak Ato l l , Geo l . Soc .

Amer . Bul l . , 86: 1 58 1 - 1 584 .

Carpenter , M . L . , W . B . Jackson , and M . W . Fal l , 1 968 , B i rdPopulation s at En iwetok Ato l l , M icr ones ica, 4: 295-307 .

Emer y , K . O . ,J . I . T racey , and H . S . Ladd , 1 954 , Pt . 1 , Geo l

ogy,Geo logy of B ik in i and Near by Ato l ls (Mar shal l Is lands ),

U . S. Geo l . Su r vey , Prof Paper 260 -A .

Fal l , M . W . , A . B . Medina, and W . B . Jackson , 1 971 , FeedingPatter ns of Ratt us r at tus and Rattu s ex u lans on En iwetokAto l l , Mar shal l Is lands , J . Mamma l . , 52: 69-76 .

Fosberg , F . R . , 1 955 , Nor ther n Mar shal l Is lands Expedition ,

1 951 — 1952 . Land B iota: Vascular P lants , Ato l l Res . Bu l l . ,No . 39,

Pacific Science Board , Wash ington , D . C .

1 960 , The Vegetation of M icrones ia, Bu l l . Amer . Mus . N at .

H is t , 1 1 9: 1-76 .

and M . H . Sachet, 1982 , M icr ones ian Poaceae: Cr i t ica l andD istr ibutional Notes , M icr ones ica, 1 8: 45

- 1 02 .

M . H . Sachet, and R . O l iver , 1 979 , A Geog raph ical Checkl is t of the Micr ones ian D icotyledoneae, M icr ones ica, 1 5:

41 -295 .

M . H . Sachet , and R . O l iver , 1 982 , Geog r aph ica l Check l istof the M icr ones ian Pter idophyta and Gymnospermae,

M icr ones ica, 1 8: 23 -82 .

H ines , N . O . , 1962 , Pr ov ing Ground . An Accoun t of the

Radiobio logical Studies in the Pacif ic, 1 946— 1 961 , Un iver s i tyof Wash ington Pr ess , Seatt le.

Jackson , W . B . , 1 969 , Sur v ival of Rats at En iwetok Ato l l , Pac .

Sc i . , 23 : 265-275 .

and M . L . Carpenter , 1 967 , Radio isotope Cycl ing in Terr estr ial Commun i ties at En iwetok Ato l l , in Pr oceedings of theSecond Nat ional Sympos ium on Radioeco logy , Ann A r bor ,May 1 5— 1 7, 1 967, Atom ic Energy Comm is s ion , Oak R idge,

Tennessee, pp . 644 -651 .

Kanehi r a , R . , 1 935 , An Enumer at ion of M icr ones ian Plants ,Kyushu U n iv . Dep . Agr ic . J. , 4: 237-464 .

Koranda , J . J. , J . R . Mar t in , S. E . Thompson , Jr . , M . L . Stuar t ,

35

D . R . Mclntyr e , and G . Potter , 1973 , Enewetak Radio logicalSur vey, NVO l 40 , U . S. Atom ic Energy Comm i ss ion , Las

Vegas , Nevada, pp . 225-235 .

W . L . Robison , S. E . Thompson , and M . L . Stuar t , 1 978 ,

Enewetak Radioeco logy Resear ch Prog ram : I . Eco log ica lStudies on Enjebi Is land 1 975 — 1976 , Lawr ence L ivermore

Labor ator y Report UCRL -52409-1 , U n iver s i ty of Ca l i for n ia,

L ivermor e.

Koyama, T . , 1 964 , The Cyperaceae of M icrones ia, M icr ones ica,

1 : 59- 1 1 7

Lamber son , J . O . , 1982 , A Guide to Ter res t r ia l P lan ts ofEnewe tak A to l l , Paci fic Scient i fic Informat ion Center , Ber n iceP . B ishop Museum , H ono lu lu .

Lane, I . E . , 1 960 , in M icroc l imatic Obser vations at En iwetok ,D . I . B lumenstock and D . F . Rex , A to l l Resear ch Bu l l . , No .

71 , Paci fic Science Board , Wash ington , D . C. , pp . 1 5 19 .

Lee , M . A . , and J . S. Lee, 1 978 , Ter r es tr ial Nutr i ent Cycl ing :Ana lys is of Compar tments , M id-Pacif ic Mar ine Labo r ator yAn nua l Repo r t

, 1 975— 1 976,NVO 0628 2 , U n iver s ity of

H awai i , H ono lu lu .

Leopo ld , E . B . , 1 969 , M iocene Po l len and Spo r e F lo ra of

En iwetok Ato l l , Mar shal l Is lands , U . S . Geo l . Sur v . Pr of . Pap .

260 -11.

Morat, P . , 1 981 , Note sur les Gram inées de las Nouve l leCa ledon ie. VI: Lep tu r opet ium , genre nouveau endem ique,

Adan son ia , 20: 377-38 1 .

Palumbo , R . F . , 1 962, Recover y of the Land P lants at En iwetokAto l l Fo l lowing a Nuclear Detonation , Radiation B io l . , 1 :

1 82- 189 .

St . John , H . , 1 950 , F lora of Engebi , Aomon -B rui r e, and Run it

Is lands , Rep . UWFL -23 , Sect . V , Appl . Fish . Lab . Un iv .

Wash .1 960 , F lora of En iwetok Ato l l , Pac . Sc i . , 14: 3 13 -335 .

Tay lor , W . R . , 1 950 ,P lan ts of B ikin i and Other Nor ther n

Mar shal l Is lands , Un iver s ity of M ich igan Pr ess , Ann Ar bo r .

T r acey , J . I . , Jr ., and H . S. Ladd , 1 974 ,

Quater nar y H is tor y ofEn iwetok and B ikin i Ato l ls , Mar shal l Is lands , in Proceedings ofSecond In ter nat iona l Sympos ium on Coral Reefs , Great Barr ier Reef Comm i ttee, B r i sbane, Austral ia,

pp . 537-550 .

Welander , A . D . , K . Bonham , L . R . Donaldson , R . F . Palumbo ,

S. P . Gessel , F . G . Lowman , and W . B . Jackson , 1 966 ,

B ikin i -En iwetok Studies, 1 964 , Rep . UWFL -93 , Lab. Radiation

B iol . U n iv . of Wash .Woodbur y , A . M . , 1 962 , A Rev iew of the Eco logy of En iwetok

A to l l, Pacif ic Ocean , Contr . No . DA-42-007-403 -CML -427,

Un iv . U tah In st . of Env . B io l . Res , U . S. Arm y Chem .

Corps , Dugway Pr ov ing Gr ound .

Chapter 4

R ecen t F oramin ifera and N onp lank ton ic P rotozoans"

E . H . CH AVET andDENN IS M . DEVANEY

1'Un iver s ity of H awa i i , H awa i i Under seaResear ch Labo r ator y, H ono lu lu, H awai i 96822i Ber n ice P . B ishop Museum , H ono lu lu, H awai i 968 1 7

FORAMIN IF ERA

In 1 946 ,dur i ng Oper at ion Cr oss roads , 45 samples

contain ing Recen t Foram in i fer a wer e obtained in the

Enewetak Lagoon fr om the shor e to depths of 62 m (204feet) and on the outer s lopes at depths from 44 to 914 m( 1 44 to 3000 feet) (Cushman et al . , Of the 204

species of foram in i fera r ecor ded at Enewetak , appr oxi

mately 60% were also co l lected fr om the r eef flats ,lagoons , and outer s lopes of other Mar shal l Is land atol ls(Ronger ik , Rongelap , B ik in i) and Sylvan ia Guyot . Four teenof the 1 8 new species and two new var ieties descr ibed atthat time fr om the Mar shal l Is lands wer e found at

Enewetak .In the 1 954 s tudy , the Mar shall Is land for amin i feral

fauna was composed of near ly 85% species character is ticof the tr opi cal Paci fic , 696 new to s cience, and the balanceundetermined . No s ign i fi cant d i fferences in the for am in if

er al compos i tion were ev ident between the four atol ls sam

pled dur ing the s tudy . Compared w i th other unspec i fiedPaci fic is lands , the absence of Bacu logyps ina spp . ,

Calcar ina calcar , and medium-to-Iarge Rotalia, as wel l asthe pr esence of a greater var iety of m i liolid species , wasappar ent (Cushman et al . ,

Gener al features of speci fic biotopes (reef flat , lagoon ,

outer s lope, and deep water s) w i th thei r dominan t or re

s t r icted for amin i feral componen ts were g iven , togetherwi th speci fic s tation loca l i ties (op .

For amin i fera from Enewetak were also r ecor ded fr ombo r ings at two dr i ll s i tes on the ato l l (Co le, 1 957 ; Toddand Low, These s i tes include a large number of

extant species . In fact , Co le (op . ci t . ) repor ted that samples‘W i th the exception of plankton ic for am in iferans , Which are

included in th is chapter , other plankton ic protozoans (dinoflagellates , r adio lar ians , and t int inn ids ) are presented in Chapter 20(th is vo lume) .

fr om the sur face to about 1 86 m (6 10 feet) con tained on lyRecen t species . The pr esent l is t of Recent EnewetakForam in i fera r ecor ded fr om these two paper s includes 23addi t ional spec ies that wer e l is ted among Recen t Mar shal lls land spec ies by Cushman et al .

Todd ( 1960) included Enewetak among local i ties discus sed in the d is tr ibution of Ca lcar ina spengler i and

absence of member s of the genus Bacu logyp s ina.

Lack ing a r epresentative iden t i fied co l lect ion of Foram in ifer a and other pro tozoans at Enewetak , M id-Pac i ficResear ch Laborator y (MPRL ) contr acted for such a collec

tion to be made in 1956 . Approximately 1 40 for am in i fer alspecies wer e determ ined from the 1 42 samples obtainedfrom shal low water throughout the ato l l . The seawar d r eefand r eef flat genera were compar ed w i th those found inbeach sands and plankton tows (H i r shfield et al . ,

Th i s wor k also compared the for amin i feral gener a of

Enewetak w i th that from smal ler samples taken in H awai i ,Kwajalein , and the So lomon and Car ol ine Is lands . The

species l is t (unpubl ished r epor t to MPRL by H i r shfield,

H elson , and Charmatz , 1 957) included sever al add i tions tothe Enewetak fauna, s ix of whi ch were also found byChave in 1 981 .

Subsequen t wor k on Foram in i fer a at Enewetakincluded s tud ies on the fine s tr ucture and morpho logy of

Car ter ina sp icu lo tes ta (L ipps and Enr ico , 1 973 ; Deutschand L ipps , 1976 ; Conger et al . , Foram in i feran pr edation by fishes and the methods of feeding by foram in i fer ans were also inves tigated at Enewetak (L ipps and

Delaca , The species , Amphis tegi na les son i i and A .

obifera, r ev iewed by H alloch and Lar sen ( 1979) includedmater ial fr om Enewetak . Shower s and Atk inson (1 979)wor ked at the ato l l on the di fferent forms of Rosalinag lobu lar is .

The checkl is t of Recent Foramin i fera and nonp lank

ton ic pr otozoans fr om Enewetak is pr esented in Table 1

and contains 280 species . The above-men t ioned publ icat ions l is t 233 of these species . For ty-seven new Enewetakr ecor ds have been added to th is l is t by Chave (TableFour teen of these species wer e l is ted by Cushman et al .

( 1954) fr om other Mar shal l ls land ato l ls ; 33 species ar e

new to Enewetak and the Mar shal l Is lands .

CHAVE AND DEVANEY

TABLE 1

Checkl i s t of Recen t For am i n i fer a and Nonp lan k ton ic Pr otozoans‘

Phy lum SARCODINAClass RETICULAREASubc lass GRANULORETICULOSIAOrder FORAMIN IFERIDAFam i ly SACCAMMIN IDAESaccamm ina sp .

P roteon ina sp . : Cushman et al . , 1 954 .

Fam i ly H YPERAMMINIDAESagen ina f r ondescens (Br ady) : Todd and Low , 1960 .

Fam i ly NOURIIDAETNou r ia po lymorphino ides H er on

-Al len and Ear land: Cushman et al . , 1954 .

Fam i ly LITUOL IDAE‘Placop s i l ina bradyi Cushman and McCullouch , 1 939 .

Fam i ly TEXTULARIIDAEB igener ina nodosar ia d

’

Or bigny: Todd and Low , 1960 .

TB igener ina sp . : Cushman et al . , 1954 .

Clavu l ina angu lar is d’

Orbigny: Todd and Low , 1 960 .

Clavu l ina pacifica Cushman : Cushman et al . , 1954 .

Text u lar ia agglu t inans d’

Or bigny: Cushman et al . , 1954 ; Todd and Low , 1 960 .

Tex tu lar ia candeiana d’

Or bigny: Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Tex tu lar ia con ica d’

Or bigny: Cushman et al . , 1954 ; Todd and Low , 1960 .

Tex tu lar ia fol iacea H eron -A l len and Ear land: Cushman et al . , 1954 .

Textu lar ia fol iacea var . ocean ica Cushman : Cushman et al . , 1954 ; Todd and Low , 1 960 .

Textu lar ia ker imbaens is Said : Todd and Low , 1960 .

Textu lar ia m i llett i Cushman : Cushman et al . , 1 954 .

‘Tex tu lar ia sagi tu la var . fis tu losa Brady , 1 884 .

Textu lar ia sem ialata Cushman : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Valvu l ina davidiana Chapman : Cushman et al . , 1 954 ; Todd and Low , 1960 .

Fam i ly TROCH AMMIN IDAE‘Trochamm ina n it ida B rady , 1 884 .

TTrochamm ina cf . r otal iform ia Wr ight : Cushman et al . , 1954 .

Fam i ly ATAXOPH RAGMIIDAETG audryina cf . pauperata Ear land: Cushman et al . , 1954 .

G audryina'tr iangu lar is angu lata Cushman : Todd and Low , 1 960 .

G . t r iangu lan’

s var . angu lata Cushman : Cushman et al . , 1954 .

G audryinatr u ll is sata Todd : Cushman et al . , 1954 .

Gaudryina (Siphogaudryina) rugu losa Chapman : Cushman et al . , 1 954 : Todd and Low , 1 960 .

Gaudryina (Siphogaudryina) s iphon ifera B r ady : Todd and Low , 1 960 .

Kar r er iella br adyi (Cushman) : Cushman et al . , 1 954 .

Fam i ly F ISCH ERINIDAECom usp i ra p lanor bis Schultze: Cushman et al . , 1954 ; Todd and Low , 1 960 .

P lan isp i r inella ex igua (Br ady) .P lan isp ir in ia ex igua (B r ady): Cushman et al . , 1954 .

Fam i ly NU BECULAR IIDAENebecu lar ia lacunens is Chapman : Cushman et al . , 1954 .

‘N ubecu l ina divar icata (B rady ,

‘P tychom i l iola separ an s Eimer and Ficker t , 1921 .

TSp i rolocu l ina acescata Cushman : Cushman et al . , 1 954 .

Sp i rolocu l ina angu lata Cushman : Cushman et al . , 1954; Todd and Low , 1 960 .

Sp i r olocu l ina clar a Cushman : Cushman et al . , 1954 ; Todd and Low , 1960 .

Sp ir olocu l ina clara var . li rata Cushman : Cushman et al . , 1 954 ; Todd and Low , 1960 .

Sp i r o locu l ina commun is Cushman and Todd : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Sp i r o locu l ina cor r ugata Cushman and Todd : Cushman et al . , 1954; Todd and Low , 1 960 .

Sp i r olocu l ina jo l ium Todd : Todd and Low , 1 960 .

Sp i rolocu l ina sp . A : Cushman et al . , 1954 .

‘Sp i rolocu l ina gr ateloup i d

’

Or bigny, 1839 .

Sp i r olocu l ina mar shallana Todd : Cushman et al . , 1 954 ; Todd and Low , 1960 .

‘New Enewetak r ecord .

‘

I‘Not found in 1 98 1 Enewetak co l lections . (Th i s table continued on next page. )

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CHAVE AND DEVANEY

TABLE 1 (con t'd)

Fam i ly NODOSARIIDAEBol ivinella [o lia (Parker and Jones ) : Todd and Low , 1960 .

Bol ivinella fo l ia var . om ata Cushman : Cushman et al . , 1 954 .

‘Den tal ina fil iform is (d’

or bigny ,“

l'F r ondicu lar ia sp . A of Boomgaar t : Cushman et al . , 1 954 .

Lagena g lobosa (Montagu) : Cushman et al . , 1954 .

'l'Lagena Sp i r alis Brady : Cushman et al . , 1954 .

I'Lagena s t ri ato -punctata Parker and Jones : Cushman et al . , 1954 .

I'Lagena wi ll iamson i (Alcock) : Cushman et al . , 1954 .

Nodosar ia pauci locu lata Cushman : Cushman et al . , 1954 .

Fam i ly VAG INUL INIDAE‘As tacolus pacifica (Cushman and H anazawa,

°

As tacolus p lanu lata Gal loway and Whi ss ler , 1927 .

Robu lus gi bbus Cushman et al . , 1 954.

Robu lus l imbos us (Reuss) : Cushman et al . , 1954 .

°

Robu lus or bicu lar i s (d’

Orbigny,

Robu lus sp . A . : Cushman et al . , 1 954 .

Fam i ly POLYMORPH INIDAE°

Roma lino g lobu l ifera Br ady, 1 879 .

Fam i ly GLANDUL IN IDAEF is s u r ina ci r cu lar i s Todd : Cushman et al . , 1 954 ; Todd and Low , 1960 .

‘

I'F is su r ina formosa (Schwager ) : Cushman et al . , 1 960 ; Todd and Low , 1 960 .

TF iss u r ina lacunata (Bur r ows and H o lland) : Cushman et al . , 1 954; Todd and Low , 1960 .

F is s u r ina marginato-perforata (Seguenza) : Todd and Low , 1 960 .

F is s u r ina sp . B : Cushman et al . , 1954 .

F is s u r ina m i llett i Todd : Cushman et al . , 1954 .

F is s u r ina or bignyana flin t i (Cushman) : Todd and Low , 1 960 .

F is su r ina sp . C: Cushman et al . , 1 954 .

l F is s u r ina sp . A : Cushman et al . , 1 954 .

Fam i ly SPIRIL L IN IDAEA lanwoodia campanaeform is (B r ady ,Con iCOSp ir i ll ina sem i - invo luta Cushman : Todd and Low , 1960 .

P lan iSp ir i ll ina den t icu logr anu lata (Chapman ) .Sp i r i ll ina den t icu lo-

gr anu lata Chapman : Todd and Low , 1 960 .

Plan iSp i r i llina tuber cu lato l imbata (Chapman ) .Sp i r i ll ina tuber cu lato - limbato Chapman : Cus hman et al . , 1 954 ; Todd and Low , 1960 .

Sp ir i ll ina decorate Br ady : Todd and Low , 1 960 .

Sp i r i ll ina decor ata B r ady var . of Sidebo ttom : Cushman et al . , 1954 .

‘Sp in

‘

ll ina inaegual is B rady , 1 879 .

TSp in‘

ll ina sp in igera Chapman : Todd and Low, 1960 .

Sp i ri ll ina v ivipara var . densepunctata Cushman : Cushman et al . , 1954 ; Todd and Low , 1960 .

Sp ir i ll ina vivipara var . rever tens Rhumbler : Cus hman et al . , 1954 .

Fam i ly CERATOBULIMINIDAE‘Ceratobu l im ina pacifica Cus hman and H ar r ls , 1927 .

H oeg lundina flin t i (Gal loway and Wh iss ler ,H oeg lundina elegans d’

Or bigny: Cushman et al . , 1954 .

'Lamar ckina scabra (Br ady ,°

M iss iss ipp ina concen tr ica (Par ker and Jones ,Fam i ly ROBERTIN IDAE

‘All iati na excen tr ica (di Napo l i All iata,

Fam i ly TURRILIN IDAE‘I‘

Bu l im inella elegan ti ss ima Cushman et al . , 1954 .

Bu l im inella m i llett i Cushman : Cushman et al . . 1 954 .

Bu l im ino ides wi ll iamson ianus (B r ady) : Cushman et al . , 1954 .

Fam i ly SPHAEROID IN IDAETSphaero idina bu ll io ides d

’

Or bigny: Cus hman et al . , 1 954 .

Fam i ly BOLIVIN ITIDAE‘Bo l ivina abbreviata H eron-Al len and Ear land, 1 924 .

Bo l ivina compacta Sidebottom : Cus hman et al . , 1954 ; Todd and Low , 1 960 .

‘New Enewetak r ecord .TNot found in 198 1 Enewetak co llections . (This table conti nued on next page.)

FORAMINIFERA AND NONPLANKTONIC PROTOZOANS 4 1

TABLE 1 (con t’d)Fami ly BOLIVINITIDAE (cont ’d)Bolivina pseudopygmaea Cushman : Cushman et al . , 1954 ; Todd and Low , 1 960 .

Bolivina r homboidalis (M i llett) : Cushman et al . , 1954 ; Todd and Low , 1 960 .

Bolivina r obus ta Brady : Cus hman et al . , 1 954 .

‘Bol ivina spinescens Cushman , 191 1 .

Bol ivina s tr iatu la Cushman : Cushman et al . , 1 954 ; Todd and Low , 1960 .

Bolivina subexcavata Cushman and Wickenden : Cushman et al . , 1954.

°

Bol ivina vadascens Cushman , 1 933 .

Rectobol ivina dimorpha var . pacifica (Cushman)Siphogener ina dimorpha var . pac ifica Cushman : Cushman et al . , 1 954 .

Fam i ly ISLANDIEL LIDAE‘Cass idu lino ides sp .

Fam i ly BULIMIN IDAEBullm ina fijiens is Cushman : Cushman et al . , 1 954 .

F ijiella s imp lex (Cushman ,

Reussella s imp lex (Cushman): Cushman et al . , 1 954 , Todd and Low , 1960 .

F ijiella sp inu losa (Reuss ,Reussella sp . A : (Cushman) : Cushman et al . , 1954 ; Todd and Low , 1960 .

M imos ina pacifica Cushman : Cushman et al . , 1954 .

Fam i ly UVIGERIN IDAESiphogener ina raphana (Parker and Jones ) : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Tr ifar ina albat r os s i var . om ata (Cushman) .Angu loger ina albat r os s i var . om ata Cushman : Cushman et al 1 954 .

Tr ifar ina car inata (Cushman ,

Tr ifar ina bradyi Cushman : as figu r ed by Cushman et al . , 1 954 .

‘Tn

‘

far ina bradyi Cushman , 1942 .

‘Tr ifar ina scrobicu lata (Cushman ,

Uviger ina por r ecta Br ady : Cushman et al . , 1954 .

“

I'Uviger ina pr oboscidea Schwager : Cushman et al . , 1954 .

l gen'

na pr obosc idea var . vadescens Cushman : Cushman et al . , 1954 .

Fam i ly DISCORBIDAED iscorbinella s ubber thelot i (Cushman) .D iscor bis subber theloti Cushman : Cushman et al . , 1954 .

Ep is tom inella pulchra (Cushman) : Cushman et al . , 1954 ; Todd and Low , 1 960 .

Ep is tom inella sp . : Cushman et al . , 1 954 .

Neoconor bina cr us tata (Cushman) .D iscor bis cr us tata Cushman : Cushman et al . , 1954 .

Neoconorbina tuberocap itata (Chapman) .D iscor bis tuberocap itata (Chapman) : Cushman et al . , 1 954 .

°

Patell inella car inata Co ll ins , 1 958 .

°

Patellinella f ijiana Cushman , 1933 .

‘

l’

PatelIinella inconspicua (B rady): Cushman et al . , 1954 .

Rosa l ina concinna (B rady).D iscorbis concinna (Br ady) : Cushman et al . , 1 954 .

Rosa lina flor idana (Cushman ,

D iscor bis op ima Cushman : Cushman et al . , 1 954 .

'Rosal ina globu lar is d

’

Or bigny, 1 826 .

Rosa lina m icens (Cushman) .D iscor bis m icens Cushman : Cushman et al . , 1954 .

°

Ros alina vi llardeboana d’

Orbigny, 1 839 .

°

Rotor binella lobatu lus Par r , 1950 .

Rugidia cor ticata (H eron-Al len and Ear land) : Cushman et al . , 1954 .

Rugidia sp inosa Cushman : Cushman et al . , 1 954 .

Tretomphalu s bu llo ides 1 839.

Tretomphalus p lanus Cushman : Cushman et al . , 1954 .

‘Tretomphalus m i llett i i H eron

-Al len and Ear land, 1915 .

Valvu l inen‘

a rugosaD iscor bis rugosa Cushman et al . , 1954 .

Rosa lina r ugosa d’

Or bigny: Todd and Low , 1960 .

‘New Enewetak r ecord .found in 1 981 Enewetak co llections . (T h is table continued on next page. )

42 CHAVE AND DEVANEY

TABLE 1 (con t'd)Fam i ly EPONIDAE

‘Epon ides ber thelot ianus (d

'Or bigny,

Epon ides Iateralis (Terquem ) : Todd and Low , 1 960 .

Epon ides r epandus (F ichtel and Mo ll ,Por epon ides cr ibro repandu s Asano and U icho : Todd and Low , 1960 .

Fam i ly GLABRATELL IDAEG labratella patell iform is (Brady) .D isco r bis patell i/o rm is (B r ady) : Cushman et al . , 1954 .

Neoconor bina tabem acu lar is (B r ady) : Todd and Low , 1960 .

H eronallen ia sp . : Cushman et al . , 1 954 .

Fam i ly PEG IDIIDAEPegidia dubia Todd and Low , 1960 .

Fam i ly EPISTOMARIIDAEEp is tomar o ides po lys tomello ides (Parker and Jones ) : Cushman et al . , 1 954 ; Todd and Low, 1 960 .

Fam i ly SIPH ONIN IDAESiphon ina tu bu los a Cushman : Cushman et al . , 1954 .

Siphon ino ides ech inata (B rady) : Cushman et al . , 1 954; Todd and Low , 1 960 .

Fam i ly AMPH ISTEG IN IDAEAmphis tegina bicir cu lata Lar sen , 1 976 .

Amphis tegi na radiata var . venosa (F ichtel and Mo ll) : Cushman et al . , 1 954 .

Amphis teg ina madagascar iens is d’

Or bigny: Cushman et al . , 1954 (in par t) .Amphis teg ina lesson i i H alloch and Lar sen , 1 979 .

Amphis tegi na madagascar iens is d’

Orbigny: Cushman et al . , 1954 (in par t) : Todd and Low , 1 960 (in par t ?)Amphis teg ina lobifera Lar sen : H alloch and Lar sen , 1 979 .

Amphis teg ina madagas car iens is d’

Or bigny: Cushman et al . , 1954 (in par t) ; Todd and Low , 1960 (inAmphis tegina pap i l losa Said .Amphis tegina radiata var . pap i l losa Said : Cushman et al . , 1 954 .

Amphis tegi na sp .

Amphis teg ina radiata (F ichtel and Mol l) : Cushman et al . , 1 954 .

Amphis tegina radiata var . : Cushman et al . , 1954 .

Fam i ly CIB ICID IDAETCibicides cicatn

'

cos us (Schwager ) : Cushman et al . , 1 954 .

Cibicides lobatu lus (Walker and Jacob) : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Cibicides mayo r i (Cushman) : Todd and Low , 1 960 .

Cibicides refu lgens Mon tfor t : Todd and Low , 1 960 .

Cibicidella var iabi lis Cushman et al . , 1954 ; Todd and Low , 1 960 .

‘Planu l ina ar im inens is d

’

Or bigny, 1 826 .

P lanor bu lino ides r et inacu latus (Parker and Jones) : Todd and Low , 1 960 .

Fam i ly PLANORBUL IN IDAEPlanor bu l ina acer val is B r ady : Cushman et al . , 1954 ; Todd and Low , 1960 .

Planor bu l inella lar vata (Parker and Jones ) : Todd and Low , 1960 .

Fam i ly ACERVUL IN IDAEAcer vu l ina inhaerens Schul tze: Cushman et al . , 1 954 ; Todd and Low , 1960 .

Gyp s ina g lobu lo (Reuss ) : Cushman et al . , 1954 ; Todd and Low , 1960 .

G yp s ina p lana Car ter : Todd and Low , 1960 .

Gyp s ina ves icu lan‘

s (Parker and Jones ) Cushman et al . , 1 954 ; Co le, 1957 , Todd and Low , 1960 .

‘Planogyp s ina squam iform is (Chapman ,

Fam i ly CYMBALOPORIDAE‘Cymbalopo rella tabellaeform is (B r ady ,Cymbalopo r etta br adyi (Cushman) : Cus hman et al . , 1 954 ; Todd and Low , 1 960 .

Cymbalopo r etta squammosa Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Pe p i lus r otundatus Cushman : Cushman et al . , 1954 ; Todd and Low , 1 960 .

Fam i ly HOMOTREMIDAEB iar r i tz ina mon t icu lar is (Car ter ) .Carpen ten

'

a mon ticu lar is Car ter : Todd and Low , 1960 .

H omotrema rubrum (Lamarck) : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

M in iacina m in iacea (Pal las ) : Cushman et al . , 1 954 .

i Fam i ly GLOBOROTAL IIDAE

G lobo rotalia cras saform is Ga l loway and Whiss ler , 1 927 .

‘New Enewetak r ecord .

TNot found in 198 1 Enewetak co llections .

ZtSpecies names of fam i ly updated but not studied in 1 98 1 . (Th is table con ti nued on next page .)

FORAM INIFERA AND NONPLANKTONIC PROTOZOANS

TABLE 1 (con t’d)

Fam i ly GLOBOROTAL IIDAE (cont ’d)G loborotal ia cras sa Cushman et al . , 1 954 .

G lobo rotal ia cf . h i r suta Cushman et al . , 1 954 .

G loborotal ia menardi i Cushman et al . , 1 954 ; Todd and Low , 1960 .

G lobo rotal ia tr uncatu l ino ides Cushman et al . , 1954 .

G lobo r o tal ia tum ida (B r ady) : Cushman et al . , 1 954 .

t Fam i ly H ANKENTIN IDAE

H as t iger ina s iphon ifera (d’

Or bigny,

G lobogen‘

nella aequ i late ral is (Br ady) : Cushman et al . , 1 954 .

¢Fam i ly GLOB IGERIN IDAECandeina n it ida d

'Orbigny: Cushman et al . , 1 954 .

G lobigen‘

na bu llo ides d’

Or bigny: Cushman et al . , 1 954 ; Todd and Low , 1960 .

G lobigen’

na eggen‘

Rhumbler : Todd and Low , 1 960 .

G loboger ina s ubcretacea Lomn icki : Cushman et al . , 1 954.

G lobiger ina? sp . : Cushman et al . , 1 954 .

G lobigen'

no ides conglobatus (B r ady) : Cushman et al . , 1 954 ; Todd and Low , 1960 .

G lobiger inoides ruber Cushman et al . , 1954 ; Todd and Low, 1 960 .

G lobiger ino ides saccu l ifer (B rady): Cushman et al . , 1954 ; Todd and Low , 1 960 .

H as t iger inella digi tata (Rhumbler ) : Cushman et al . , 1 954 .

Or bul ina un iver sa d’

Or bigny: Cushman et al . , 1954 ; Todd and Low , 1960 .

Pu llen iat ina obl iqu i locu lata (Par ker and Jones ) : Cushman et al . , 1 954 ; Todd and Low , 1960 .

Sphaero idinella dehiscen s (Parker and Jones ): Cushman et al . , 1 954 .

Fam i ly ROTAL IIDAEAmmon ia beccar i i tep ida (Cushman) .Rotalia cf . beccar i i var . tep ida Cushman : Cushman et al . , 1 954 .

St reblus beccar i i tep ida (Cushman) : Todd and Low , 1 960 .

‘Carpen ter ia balan iform is Gr ay , 1 858 .

Carpen ten‘

a u tr icu lan'

s (Carter ,Carpen ten

‘

a proteiform is Goes : Todd and Low , 1960 .

Fam i ly CALCARIN IDAECalcar ina h isp ida B rady : Cushman et al . , 1 954 ; Todd and Low , 1 960 ; Todd, 1 960 .

Calcar ina spengler i (Gmel in) : Cushman et al . , 1 954 ; Todd and Low , 1960 ; Todd, 1 960 .

Fam i ly ELPH IDIDAEE lphidium advenum (Cushman) : Cushman et al . , 1954; Todd and Low , 1 960 .

E lphidium advenum var . dispar Cushman : Cushman et al . , 1954 .

Elph idium jen sen i (Cushman) : Cushman et al . , 1954 ; Todd and Low , 1 960 .

‘Elphidium ocean icum Cushman , 1 933 .

Elphidium s imp lex Cushman : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Fam i ly NUMMUL IT IDAE‘Cyc loc lypeus carpen ten

‘

Br ady , 1 856 .

H eteros tegina s ubo r bicu lar is d’

Or bigny: Cushman et al . , 1 954 ; Co le, 1 957; Todd and Low , 1 960 .

Oper cu l ina ammono idea (G ronov ius ) : Cushman et al . , 1 954 .

Fam i ly CAUCASINIDAECas s idella ear landi (Cushman) .Vi rgu l ina ear landi Cushman : Cushman et al . , 1 954 .

Sigmaui rgu l ina to r tuosa (B rady).Bo l ivina tor tuosa Brady : Cushman et al . , 1954 ; Todd and Low , 1 960 .

Fam i ly LOXOSTOMATIDAELoxos tomum convallar ium (M i llett ) : Cushman et al . , 1 954 .

Loxos tom um kar ren‘

anum (B rady): Todd and Low , 1 960 .

Loxos tom um Iimbatum (B rady) : Cushman et al . , 1 954 ; Todd and Low , 1960 .

Loxos tomum mayo r i (Cushman ) : Cushman et al . , 1954.

Fam i ly CASSIDULIN IDAE"( Cass idu lina angu losa Cushman : Cushman et al . , 1 954 .

Cas s idu lina cos tatu la Cushman : Cushman et al . , 1 954 .

‘Cas s idu lina cr as sa d’

Or bigny, 1 839 .

‘Cass idu lina gemma Todd : Todd, 1 954 .

Cass idu lina mar shallana Todd : Cushman et al . , 1 954 .

‘New Enewetak r ecord .

(“Not found in 1 98 1 Enewetak co llections .

zt Species names of fam i ly updated but not studied in 1 98 1 . (Th i s table continued on next page. )

CHAVE AND DEVANEY

TABLE 1 (con t’d)

Fam i ly CASSIDUL INIDAE (cont ’d)Cass idu lina m inuta Cushman : Cushman et al . , 1954 .

‘Cass idu lina m ur r hyna Schwager , 1 866 .

Cas s idu l ina or iangu lata Belford, 1966 .

Cass idu lina s ubg lobosa Br ady : Cushman et al . , 1 954 (in par t) .Cass idu lina patu la Cushman : Cushman et al . , 1954 .

Cas s idul ina su lcata Belford, 1 966 .

Cass idu lina de licata Cushman : Cushman et al . , 1 954.

”

( Cas s idu lina s ubg lobosa Br ady : Cushman et al . , 1 954.

‘

I‘

Cas s idu l ina s ubg lobosa Brady var : Cushman et al . , 1 954 .

Ehr enbergi na pacifica Cushman : Cushman et al . , 1 954 .

‘Ehr enbergina r et icu lata Cushman ,

Fam i ly NONIONIDAE‘As tr onon ion tum idum Cushman and Edwards : Cushman et al . , 1954 .

N on ion pacif icum (Cushman) : Todd and Low , 1960 .

TNon ionella sp . A : Cushman et al . , 1 954 .

Non ionella tr ans lucens Cushman : Cushman et al . , 1954 .

TPu lIen ia sal isbu ryi Stewart and Stewar t : Cushman et al . , 1 954 .

Fam i ly ALABAMIN IDAEA labam ina tubu lifer a (H eron-Al len and Ear land) .Ep is tom inella tubu lifer a (H eron-Al len and Ear land) : Cushman et al . , 1954; Todd and Low, 1960 .

TGyro idina soldan i i d’

Orbigny: Cushman et al . , 1 954 .

TOr idor sal is tenera (Brady) .Epon ides tenera (B rady) : Cushman et al . , 1954 .

Fam i ly GAVELINELLIDAEAnomal ina cf . g labrata Cushman : Cushman et al . , 1954 .

Anomal inella ros tr ata (B r ady) : Cushman et al . , 1 954 ; Todd and Low , 1 960 .

Cibicido ides p seudoungeri anus (Cushman ) .Cibicides cf . p seudounger ianus (Cushman ) : Cushman et al . , 1 954 .

Fam i ly CARTERIN IDAECar ten

‘

na sp icu lotes ta (Car ter ) : Cushman et al . , 1 954 ; L ipps and Enr ico , 1 973 ; Deutsch and L ipps ,Conger et al . , 1 977 .

Phy lum CIL IOPH ORAClass KINETOFRAGMINOPH ORASubclass GYMNOSTOMATAOrder H APTORIDA

Lacrymar ia sp . : H i r shfield et al . , 1 957 .

Or der PLEUROSTOMATIDAEsp . : H i r shfield et al 1 957 .

Subc lass VESTIBUL IF ERAOrder COLPODIDA

Co lpoda sp . : H i r shfi eld et al . , 1957 .

Class OLIGOH YMENOPHORASubc lass H YMENOSTOMATAOrder SCUTICOCIL IATA

Cohn i lembus uerm inus : H i r shfieid et al . , 1957 ; Thompson , 1968 .

Cycl idium sp . : H i r shfield et al . , 1 957 .

Pseudochon i lembus sp . : H i r shfield et al . , 1 957 .

U r onema sp . : H i r shfi eld et al . , 1957.

Subclass PERITRICH AOrder PER ITRICH IDA

Syphidia sp . : H i r shfield et al . , 1 957 .

Vor tice l la sp . : H i r shfield et al . , 1 957 .

Class POLYH YMENOPHORASubc lass SPIROTRICH AOrder H YPOTRICH IDA

Eup lotes sp . : H i r shfield et al . , 1 957 .

Sty lonych ia sp . : H i r shfield et al . , 1 957 .

‘New Enewetak r ecord.TNot found in 1 98 1 Enewetak co llections .

1:Species names of fam i ly updated but not s tudied in 198 1 .

FORAMINIFERA AND NONPLANKTONIC PROTOZOANS

TABLE 2

New Enewetak Recor ds

45

Species

Rosalina vi lardeboana

Ro tor binella lobatu lus

Tr etomphalus m i llett u

Epon ides ber thelot ian us

Planu l ina ar im ines is

Planogyp s ina squam iform is

Locati on‘

L

L

L

OS

OS

0 5

OS

0 5

L

CH

0 5

L

0 5

L

OS

L

0 5

OS

0 8

OS

Form‘l‘

N um ber :

Att ached (brown)

Attached (gray)

Cymbalopo rella tabellaeform is L

Carpen ter ia balan iform is OS Encrus ting (gr ayi sh br own )Elphidium ocean icum L

Cycloclypeus carpen ter i OS Attached (wh ite)Cas s idu lina crassa OS Attached (br own )Cas s idu lina gemma L

0 5

Cas s idu lina mur r hyna OS

Ehrenbergina r et icu lata OS Attached (gr ayi sh brown )Placop s i l ina bradyi OS Encrus ting (tan)Tex tu lar ia sag i tu la var f is tu los a OS Attached (r eddish brown)Trocham ina n iti da OS

N ubecu l ina divar icata OS

Ptychom i l iola separans OS

Sp irolocu l ina gr ateloup i L

Sp irolocu l ina sp . B 0 5

A r ticu l ina sagr a OS

Pseudomas sal ina aus tr a l is var L

CH

Pyrgo fom as in i OS

Qu inquelocu l ina bos ciana L

Qu inquelocu l ina lamar ckiana L

Qu inquelocu l irra po lygona OS

Tr i locu l ina linneiana CH

Den tal ina fil iform is OS Fr ee (r ed)As taco lus pacifica OS

As tacolus p lanu lata OS

Robu lus or bicu lan'

s OS Free (pink)Ramu l ina globu l ifera OS

Sp ir i ll ina inaegual is 0 8

‘L , lagoon ; CH , channel ; OS, outer s lope .

TL iv ing form (fo llowed by co lor of protoplasm ) : encru sting , tes t cemen ted to substrate; attached, ho lding on to substr ate wi th pseudopods ; fr ee, not attached to subs tr ate.

:l:TotaI number of intact specimens in all 42 samples , each con tain ing 1 cc of sand or scr aped mater ial . (Th is table con tinued on next page

46 CHAVE AND DEVANEY

TABLE 2 (Con t ’d)

Species Locat i on‘ Subs t r ate F orm'

l'

N umber tl:

80 to 300 m Crev ices in r ocks Attached (br own )200 m Rocks Encrust ing (reddish br own )300 m Coar se sand400 m Coar se sand

‘L ,lagoon ; CH , channel ; OS, outer s lope.

TL iv ing form (fo l lowed by co lor pr otoplasm ): encrusting , tes t cemented to subs t rate; attached , ho lding on to subs tr ate wi th pseudopods ; free, not att ached to subs tr ate .

i Total num ber of in tact specimen s in al l 42 samples , each contain ing 1 cc of sand or scraped mater ial .

The taxonomy to the gener ic level is based on Loebl ichand Tappan (1 964, 1 974 , Species were iden t i fiedby Chave us ing El l i s and Mess ina ( 1 940 et seq . ) and a

lar ge number of publ icat ions of wh ich Gr aham and M i l itante Bar ker Todd and Belfo r d(1 966) wer e espec ial ly usefu l .

CILIOPH ORA

Sever al mar ine ci l iates wer e co llected at Enewetakfr om reef flat scum (H i r shf ield et al . , In add i t ion ,

Scyphidium sp . was at tached to unspec i fied inver tebrateson the r eef , and Cycl idium sp . was found w i th habi tatunspec i fied (H i r shf ield, op . Obser vat ions made byPatr ick Co l in fr om 1 979 to 1 980 off Run i t Is let r evealedd isco lor ed br own ish ar eas on the lagoon bottom , wh ichwer e found by Devaney to be formed by h igh concentr at ion s of an undeterm ined per i tr ich c i l iate that r esemblesScyph idia .

Dur ing 1 966 , 42 hymenos tomme c i l iates wer e col

lected fr om the ocean and lagoon s ides of n ine i s lets at

Enewetak Ato l l . Permanen t s i lver — impr egnated s l ides weremade of these pro tozoan s from cultures (Thompson and

Sel ler s , but species wer e not l is ted . Subsequen tly ,Thompson (1 968) des cr ibed Cohn i lembus verm in us fr omEnewetak .

Berger ( 1 964) no ted that undeterm ined endocommen

sal c i l iates wer e presen t in sever al unspec i fied r egular sea

u r ch ins but wer e absen t fr om at leas t two spec ies (Eucidar is metu lar ia and Par asal in ia gr at iosa) and fr om i r r egu laru r ch ins and other un speci fied ech inoderm s exam ined . The

class i ficat ion fo r ci l iophoran s fo llows Cor l is s

OTH ER PROTOZOAN S

Fr esh and brack i sh water protozoans at Enewetakr evealed main l y amoebo id types fr om near ly 100 waterand so i l s i tes on n ine i s lets . Add i t ional fi sh , gecko , rat and

bi r d blood , and other t i s sues wer e exam ined for pr otozoanpar as i tes (D i l lon and Kasweck ,

N EW EN EWETAK RECORDS

In 1 981 , 42 samp les wer e s tud ied at Enewetakimmed iately after being co l lected from the lagoon at

depths of fr om 1 to 6 1 m , in the channels fr om 1 to

33 m , and on the outer s lopes fr om 1 to 400 m . Of the

new Enewetak r ecor ds , 23 species wer e represented byone to thr ee spec imen s and 14 species wer e fai r ly common but wer e obtained by methods not used dur ing the

1 946 co l lect ions (har d sur faces wer e scr aped by hand todepths of 33 m , and r ocks were br ought up fr om deepar eas by the submer s ible Mak i l i

’

i) . The final 10 spec ieswer e common in sed iment samples and were not r epor tedby Cus hman et al .

Twen ty-n ine spec ies l is ted by Cushman et al . (1 954)wer e not found by the sen ior author . Twenty-seven of

these spec ies were l i s ted as r ar e or fr om the 91 4 m s ta

t ion (op . Two spec ies , H auer ina ser rata and

As tr onon ion tum idum ,wer e l is ted as common in lagoon

sed imen ts (op .

ACKNOWLEDGMEN TSWe acknowledge and are gr atefu l to H . I . H i r shfield,

New Yor k U n iver s i ty , for perm iss ion to include the recor dsof c i l iates g iven in hi s 1 957 r epor t to MPRL . J . Res ig at

the U n iver s i ty of H awai i made valuable sugges t ions and

cor r ect ion s in the Foram in i fer a par t of th is paper . The

fac i l i t ies at Enewetak and mater ial obtained by the H awai iU nder sea Resear ch Laborator y ’s submer s ible Makal i

’

i

wer e made avai lable to bo th author s through gr an ts to

Patr ick Co l in and John H ar r i son of MPRL fr om NOAA ’

s

Nat ional Under sea Resear ch Prog ram , the Depart ment ofEner gy , and the Defense Nuclear Agency .

REF ERENCES

Barker , R . W . , 1960 , Taxonom ic Notes on the Species Figur edby H . B . Br ady in hi s Repor t on the Foram in ifera Dr edged bythe H MS Chal lenger Dur ing the Year s 1 873 — 1 876 ,

Spec . Pub . 9, Tulsa , Ok lahoma, 238 pp .

Belford , D . J. , 1 966 , M iocene and P l iocene Smal ler Foram in i fera fr om Papua and New Gu inea , Aus tr . Bu r . M in . Res .

Geo l . Geoph . Bu l l . 79, 306 pp .

Berger , J. , 1964 , M id-Pacific Resear ch Labo rator y Annual Rept . ,F Y 1 963 — 64 , 3 pp .

Co le, W . S. , 1 957 , Larger Foram in ifera from En iwetok Ato ll Dr i l lH o les , U . S. Geo l . Sur v . Prof . Pap . 260-V,

pp. 743-784 .

https://www.forgottenbooks.com/join

Porifera of Enewetak A to ll

Com pi led by DEN N IS M . DEVAN EY (deceased)

Ber n ice P . B is hop Museum .

H ono lu lu, H awai i 968 1 7

The fi r s t sponges r epo r ted fr om Enewetak wer e basedon co llect ions made between 1 946 and 1 948 fr om a

number of tr op i cal Paci fic ar eas (de Laubenfels ,

The Pac i fic Science Boar d of the National Resear ch Counci l sponsored the co llect ions pr oject . Six of the 1 3 spec iesr ecor ded from Enewetak were co l lected by dr edg ing near

the center of the lagoon (8 km nor th of the southanchorage; appr ox imately 1 1 ° 29’ N , 1 65

°1 5'E, de Lau

benfels , 1954) at a depth of 35 m . Another species camefrom the r eef flat , and s ix were fr om unr ecor ded local i t ies .

Thr ee new spec ies were r ecor ded from Enewetak , but on l yone (L is sodendoryx calyp ta) was un ique to that ato l l .

Thr ee sponges , each prey for d i fferent species of dor id

nud ibr anchs , have also been recor ded fr om Enewetak(Young , The color , form , and habi tat of thesesponges wer e given .

Beg inn ing in 1 972 , r esear ch was in i t iated on the

biochem is tr y of Enewetak sponges (M iddlebrook et al . ,1 972 ; Scheuer et al . , 1974 ; Rav i and Scheuer , 1 975;Scheuer et al . , 1 975; Chevolot et al . , 1977; Yunker and

Scheuer , 1 978 ; Rav i et al . , A sponge fr om the

lagoon was used as a sour ce of r ad ionuclides concentr atedin i t s s i l iceous spicules (H ur d and Lawson ,

The need for val id ident i fi cat ion of sponges i s clear lyind icated in r esear ch wher e chem i cal s tudies have beenmade on species that r emain undetermined (Chevolot et

al . , 1 977 ; Yunker and Scheuer , 1978 ; Rav i et al . ,Attempts to get Enewetak sponge mater ial iden t ified have

Chapter 5

r esulted in 1 7 add i t ional sponges receiv ing pr el im inar ydeterm inat ions by K laus Ruetzler (per sonal commun i cat ion ,

Examples of some or all of these species ar e de

pos i ted at U . S. National Museum of Natural H is tor y(USNM ) , M id-Pac i fi c Resear ch Laboratory (MPRL ), and theB . P . B ishop Museum (BPBM) . Br ief descr ipt ions (color ,exter nal morphology , co lony form) and habi tat informationon these sponges were pr epar ed by Janet Lamber son forMPRL . The pr esence of several bor i ng clion id sponges in

four selected Enewetak corals has been noted byH ighsm i th The bor i ng organ isms wer e the mos tcommon infaunal assoc iates of the corals studied , wi th86% of the corals show ing sponge bioeros ion effects .

A ca lcar ean ear sponge” w i th foss i l affin i ty co llectedon the ocean s ide of B iken (Leroy) is close to M u r rayona

phaneo lep is but might be a new species (Reu tzler , per sonalcomm un i cat ion ,

I t occur s on the roo fs of small

caves w i th sclerosponges , is flat tened , and is green in

co lor . Another calcarean sponge, Neocoel ia cr ypta, and the

scler osponge, As t roscler a wi lleyana, have also been ident i

fied from Enewetak (Bas i le et al . , in pr es s) . K inzie (1 973)found scler osponges in a large cr evasse that is cons idereda ver y pr om inent featu re on the leeward s ide of EnewetakAto l l . Two species of scler osponges were fi r s t descr i bedfr om Enewetak , as well as other par ts of the tr op icallndo Paci fic , by H ar tman and Goreau ( 1 975,

The Enewetak sponge fauna remains too poor l y knownat th is t ime to reach zoogeogr aphi cal conclus ions and

awai ts the wor k of someone wi th extens ive field and taxonom ic r esear ch exper ience in the r egion to clar i fy thes ys temat ics of th i s fauna.

The class i fi cation scheme herein i s based on Ber qu is tTable 1 i s a checkl is t of Enewetak Por i fer a.

TABLE 1

Checkl i s t of Por i fer a at Enewetak A toll

Class CALCAREASubclas s CALCINEAOrder LEUCELL IDAFami ly LEUCEL LIDAE

‘Leucetta sp .

Subclass PHARETRONIDAOrder INOZOAFam i ly MURRAYONIDAE

‘Mur rayona cf . M . phanolep is Ki r kpatr ick , 1 910 .

‘New Enewetak record. (Thi s table continued on next page . )

DEVANEY

TABLE 1 (con t’d)Or der SPH INCTOZOAFam i ly CRYPTOCOELIIDAE

‘Neocoel ia cr ypta Vacelet , 1 977 .

Class DEMOSPONGIAESubc lass TETRACTINOMORPH AOrder CHORISTIDAFam i ly STELLETIDAEAncor ino acer ous (Bower bank ,H ezekia walker r i de Laubenfels , 1954 .

Fam i ly JASPIDAEJasp is s tellifera (Car ter ) : de Laubenfels , 1 954 .

Order SPIROPH ORIDAFam i ly TETILLIDAE

‘Cinachyr a spl

Order H ADROMERIDAFam i ly SUBER ITIDAETerp ios ap loos de Laubenfels : de Laubenfels , 1 954 . Young , 1 967 .

Fam i ly SPIRASTRELLIDAESp i ras tr ella decumbens R idley : de Laubenfels , 1 954 .

Sp i ras trella vagabunda R idley , 1 884 .

An thos igmella uagabunda (Ridley) : de Laubenfels , 1954 .

Fam i ly CLIONIDAEAka sp . cf . A . diagonoxea Thomas : H ighsm i th , 198 1 .

Aka spp . H ighsm i th , 198 1 .

C liona mucr onata So llas : H ighsmi th , 1 981 .

Cliona sp . cf. C . quadrata H ancock : H ighsm i th , 1 98 1 .

Cliona sp . cf . C . vi r idis Schm idt : H ighsm i th , 1 98 1 .

Cl iothosa hancocki Topsent : H ighsm i th , 198 1 .

Fam i ly TETH YIDAEAap tos un isp icu lus (Car ter ): de Laubenfels , 1954 .

‘Tethya sp .

Fam i ly CHONDROSIIDAEChondr i l la aus tral iens is Car ter : de Laubenfels , 1 954 .

Chondros ia sp .

Chondr os ia chucal la: Ravi et al . , 1979

Order AXINELLIDAFam i ly AGELASIDAEAgelas maur i tiana (Car ter ) : de Laubenfels , 1954 .

Subc las s CERACTINOMORPHAOrder H ALICHONDRIDAFam i ly HYMENIACIDONIDAEPr ianos ph lox de Laubenfels : Young , 1967.

Order POECILOSCLERIDAFam i ly CRELL IDAE

‘G r ayella sp .

‘Yves ia sp inu lata (H entschel ,Fam i ly MYXILLIDAEL is sodendoryx calyp ta de Laubenfels : de Laubenefels , 1 954 .

Fam i ly CLATH RIIDAEClathr ia abiet ina (Lamarck) : de Laubenfels , 1954 .

Order HAPLOSCLERIDAFam i ly H AL ICLONIDAE

‘H al iclona sp . 1 .

‘H al iclona sp . 2 .

‘Toxadoc ia sp .

Fam i ly NIPH ATIDAECr ibrochal ina olemda de Laubenfels , 1954 .

Fam i ly ADOCIIDAEAdoeio sp .

Fam i ly CAL LYSPONG IIDAECallyspongia fis tu lar ia (Topsent ) : de Laubenfels , 1954 .

‘Callyspongia r id leyi Bur ton , 1934 .

‘New Enewetak record . (Th is table con ti nued on next page.)

PORIFERA 5 1

TABLE 1 (con t’d)

Order DICTYOCERATIDAFam i ly DYSIDEIDAE

‘Dys idea arenar ia Bergquis t, 1 965 .

Fam i ly SPONGIIDAEH eteronema erecta Keller , 1 889 .

Thorectop samma mela de Laubenfels : de Laubenfels , 1954 .

‘H eter onema sp .

Ir cin ia halm i/orm is (Lendenfeld) : de Laubenfels , 1954 .

Fami ly DYSIDEIDAEOr der DENDROCERATIDAFam i ly APLYSIL LIDAEAp lys i lla gr acial is de Laubenfels : Young , 1967 .

Order VERONG IDAFam i ly VERONG IIDAEPsammap lys i lla purpurea (Car ter ,Thorectop samma xana de Laubenfels : de Laubenfels , 1954 .

Class SCLEROSPONG IAEOr der CERATOPORELLIDAFam i ly ASTROSCLERIDAE

‘As tr os clera wi lleyana L i ster , 1900 .

Order TABULOSPONG IDAFam i ly ACANTHOCH AETETIDAEAcan thochaetetes wells i H ar tman and Gor eau : H ar tman and Goreau , 1 975 .

Stromatospongia m icr ones ica H ar tman and Goreau : H ar tman and Goreau , 1976 .

‘New Enewetak r ecord.

ACKNOWLEDGMENTSThi s r ev iew is more near ly complete due to Roger Gu f

fey (Penns ylvan ia State Un iver s i ty) and especial ly KlausReu tzler (USNM) who have permi tted the inclus ion of thei rspecies determ inations wh ich form the bas is of new

Enewetak r ecor ds . Janet and Ph i l Lamber son , r es identmanager s at MPRL in the late 1 9705 , wer e ins tr umental incol lect ing fr esh mater ial wi th notes that now permi t f ielddeterm inat ion of at leas t some of the mor e commonsponges at Enewetak .

REFERENCES

Bas i le, L . L . , R . J . Cuffey , and D . F . Kos ich, Sclerosponges ,

Pharetr on ids , and Sphinctozoans (Rel ic Cr yptic H ard-BodiedPor i fera) in the Modern Reefs of Enewetak Ato ll , J . Paleon t . ,

in press .

Ber gquis t , P . R ., 1 978 , Sponges , Un iver s i ty of Cal i forn ia Pr ess ,

Ber keley .Chevolot , L . , S. Padua, B . N . Ravi , P. C. B lyth , and P . J .

Scheuer , 1 977, Iso lation of 1 -Methyl-4 , 5-Dibromopyr role

2-Car boxyl ic Acid and Its 3' (H ydantoyl) Propylam ide (M idpacam ide) fr om a Mar ine Sponge, H eterocyc les , 7 : 891 -894 .

De Laubenfels , M . W . , 1 954 , The Sponges of the Wes t-Cen tr alPaci fi c, Oregon State Monogr . Studies in Zoo l . , No . 7, pp .1 -306 .

H ar tman , W . D . , and T . F . Goreau , 1975, A Paci fic TabulateSponge, L iv ing Repr esentative of a New Order of Sclerosponges , Pos ti l la, 1 67: 1 - 1 3 .

1 976 , A New Cer atoporell id Sponge (Por i fera: Sclerospongiae) fr om the Paci fic, F . W . H ar r i son , and R . R . Cowdew

(Eds ), pp . 329-347, Aspects of Sponge B io logy, Academ icPr ess , N . Y .

H ighsm ith , R . C. , 198 1 , Coral B ioeros ion at Enewetak : Agentsand Dynam ics , In t . Revue Gesamen . H ydrobiol . , 66 : 335 375 .

H urd , D . C . , and D . S. Lawson , 1 975, On the Poss ible Coneent r at ion of Radionucl ides by the Si l iceous Spicu les fr omEnewetak Lagoon , MPML Ann . Rept . F Y 1 975 .

Kin zie, R . A . III , 1 973 , Sclerosponge Eco logy , EMBL Ann . Rept .F Y 1 972— 73 .

Middlebr ook, R . E . , L . W . Wi tt le, and E . D . Scura , 1 972, Iso lation and Pur i fi cation of Naturally Occur r ing Compounds of

Mar i ne Organ i sms , EMEL Ann . Rept ._F Y 1 971 72 .

Rav i , B . N . , and P . J . Scheuer , 1975 , Toxins and B iomedicalAgents of Mar ine Inver tebr ates , MPML Ann . Rept . FY 1 975.

H . Per r y Per zanowski , R . A . Ross , T . R . Erdman , and P . J .Scheuer , 1979 , Recent Research in Mar i ne Natural Pr oducts :the Puupehenones ,

l

Pur e and App l . Chem . , 51 : 1 893-1900 .

Scheuer , P . J. , B . J . Bur reson , and M . G . M i ssakian , 1 974,

Antim icr obial and Anti tumor Consti tuents of Sponges ; etc . ,

EMEL Ann . Rept . F Y 1 973 74 .

R . F . Dietr ich , and F . X . Woo lard, 1975 , Toxins and

B iomedica l Agents of Mar ine Inver tebrates , MPML Ann . Rept .F Y 1 975 .

Young , D . K . , 1967 , New Records of Nud ibranch ia (Gas tr opoda:Op isthobranchia: Nudibranch ia) fr om the Centr al and Wes tCentr al Paci fic, Veliger , 10 : 1 59- 1 73 .

Yunker , M . B . , and P . J . Scheuer , 1 978 , Mokupalides , Thr eeNovel H exapreno ids fr om a Mar ine Sponge, J . Amer . Chem .

Soc . , 100 : 307-309 .

Chapter 6

Sea Anemones of Enewetak A to ll

CH ARLES E . CUTRESS‘and A . CH ARLES ARN ESON '

I'

‘Depar tmen t of Mar ine Sc iences , Un iver s ity of Puer toR ico

, Mayag ii ez ,Puer to R ico 00708

TScr ipp s Ins t i tut ion of Oceanogr aphyLa Jo l la, Califor n ia 92093

IN TRODUCTIONThe sea anemones cover ed in th is chapter wer e col

lected and/ or photog raphed i n par t by Cu t r ess dur ing Mayand June 1 955 and in par t by Ar neson from Septem berthr ough November 1 980 . Col lect ion r ecords ar e shown at

the end of the chapter . Mos t iden t i fications wer e madefr om pr eser ved specimen s , but a few were made fr om pho

TABLE 1

Check l i s t of Enewetak Sea Anemones

Phy lum CN IDAR IAClass ANTHOZOAOrder CORALL IMORPH ARIAFam i ly ACT INODISCIDAEAct inod iscus neg lectus (Fowler , 1 888)Rhodact is howes i i Sav i l le-Kent , 1 893

Order ACT IN IAR IAFam i ly EDWARDSI IDAEEdwar ds ia pudica Klunz inger , 1 877Edwards ia g i lber tens is Ca r lgren , 193 1

Fam i ly BOLOCEROIDIDAEB unodeops is medusoides (Fowler , 1 888)Fam i ly AL ICIIDAETr iact is producto Klunz inger , 1877Fam i ly ACT INIIDAEAn thos tella badia (Ca r lgr en , 1 900)An thop leu ra n ig rescen s (Ver r i l l , 1 928)Physobr achia doug las i Sav i l le-Kent , 1 893 : A l len , 1 972, pp . 91 , 1 66 , 1 67 , 1 69 , 1 73 , 1 77, 179 F ig . 95 , 1 83 ,

229, 252 , 256 , I : En tacmaea quadr ico lo r (Rueppel and Leuckar t ) [pro par te], Dunn , 198 1 ,p . 15 , 27 .

Act in iogeton sesere (H addon and Shack leton ,

Fam i ly ACTINODENDRONIDAEAct inodendron p lumosum H addon , 1 898 : Johannes et al . , p . 542 .

Act inodendron glomeratum H addon , 1 898

Megalact is hemp r ichi i Ehr enberg , 1 834(Th is table cont inued on next page . )

tog r aphs alone. Avai lable photog raphs of Phyman thus

s t randes i and Physobrach ia doug las i are inadequate.

H eterodactyla hemp r ichi was not photogr aphed .

No publi shed taxonomic wor k on Enewetak sea

anemones exis ts . Dunn in a r ev is ion of the clownfish anemones , r efer s by catalog number s to pr eser vedspecimens fr om Enewetak wh ich she exam ined . A l len( 1 972) r efer s to and i l lus trates seven species of hos tanemones that he obser ved at Enewetak fr om 1 968 to

1 971 . Josephson ( 1 966) made phys io log ical obser vationson Calliact is po lypus , and Johannes et al . (1 972) r efer toan un iden t i fied d igg ing anemone subsequently ident i fied asAct inodendron p lumos um . Table 1 prov ides a checkl is t ofEnewetak sea anemones .

54 CUTRESS AND ARNESON

TABLE 1 (con t ’d)

Fam i ly TH ALASSIANTH IDAEH eterodactyla hemp r ichi i Ehr enberg , 1 834Cryp todendr um adhes ivum Klunz inger , 1 877: Dunn , 1 98 1 , p . 13 .

Fam i ly PH YMANTH IDAEPhyman thus s t randes i i Car lgren , 1 900

Fam i ly ST ICH ODACTYLIDAESt ichodactyla mer tens i i B r andt , 1 835: Du nn p. 99 :Sto ichact is gigan teum (For sskal) ; A l len 1 972, pp . 1 02, 1 05 , 1 24 , 1 66 , 1 73 , 1 78 Figs 93-94 , 1 84 , 1 90 , 193 ,

1 94 F ig . 1 02, 1 96 , 200 , 206 F ig 1 05 , 237 F ig . 123 , 252, 254, 255 , 258 , 259 ,I I

H eteract is aurora (Quoy and Gaimard, Dunn , 198 1 , pp . 57 , 66 : Radian thus s imp lex (H addon and Shack leton) ;A l len , 1 972,

pp . 1 24 , 166 , 1 77 , 1 8 1 , 1 90 , 1 91 F ig . 100 , 192 F ig 1 0 1 , 1 93 , 196 , 197, 230 , 250 , 252 , 258 , 259, l l .

H eteract is r itter i (Kw ietn iewsk i , H eteract is magn ifica (Quoy and Gaimard) [pro par te], Du nn , 198 1 , p . 40 :

Radian thus r i tter i (Kwietn iewsk i ) : A llen , 1 972, pp . 1 66 , 169, 1 75 Figs . 90-91 , 1 76 , F ig 92 , 200 , 220 F ig . 1 1 3,222 F ig . 1 14 , 228 F ig . 1 18 , 232 F ig . 120 , 233 , 236 , 250 , 257 , II .

H eteract is macrodactylum (H addon and Shack leton ,

H eteractis cr ispa (Ehr enberg) [pr o parte], Du nn , 198 1 , p . 48 :

Radian thus malu (H addon and Shack leton ) : A l len , 1972, pp . 152 F ig . 75, 1 66 , 168 , 1 69, 1 70 Figs . 85-86 , 1 71 F ig . 87 ,

1 72 F ig . 89, 200 , 209, 225 F ig . 1 15 , 228 , F ig . 1 1 7 , 229 F ig . 1 1 9, 233 F ig . 1 21 , 254, 255 , II .

H eteract is gelam (H addon and Shack leton ,

Macrodactyla doreens is (Quoy and Gaimard) [pr o par te], Dunn p . 29 : Radian thus gelam (H addon and

Shack leton ) : A llen , 1 972 , pp . 1 22 F ig . 56 , 1 66 , 168 , 1 77, 1 86 F ig . 98 , 1 87 F ig . 99, 200 , 252 , I .Fam i ly ISOPH ELL IIDAETelmatact is decora (Ehr enberg ,

Telmatact is verm iform is (H addon ,

Fam i ly HORMATH IIDAECall iact is po lypus (For s skal , Josephson , 1 966 , p . 305 .

Fam i ly SAGARTIIDAEVer r i llact is pagu r i (Ver r i l l ,Fam i ly AIPTASIIDAEA ip tas ia pu lchel la Ca r lgr en , 1 943 .

Order CERIANTH ARIAFam i ly ARACH NANTH IDAEIsarachnan thus bandanens is Car lgr en , 1 924 .

B IOGEOGRAPH YD is cus s ion of the geog raph ic r anges of tropi ca l Paci fi c

shallow-water sea anemones would , at th is t ime, be meaningless . In the many paper s on these anemones , speciesare often impr oper ly or inadequately identi fied and impr oper ly or inadequately synonym i zed .

Two corallimorphar ians , 24 actin iar ians , and one cer ianthar ian are present ly known fr om Enewetak . Of these,

n ine have an lndo-Pac i fic range, and 1 8 are known on lyfr om the tr op i cal Paci fic . Cu t r ess has also collected at

Majuro , Mar shall Is lands ; Fr ench Po lynes ia; and H awai i . AtMajuro ( 1 month), 19 act in iar ians , one corall imo rphar ian ,

KEY TO SPECIES

Cor al l im or phar ia

Po lyps o ften gr egar ious , smal l to med ium s i zed . Co lum n shor t , smoo th . Ten tacles ver y numer ous , ar ranged in r ad ial rows ,somet imes r ud imentar y .

1 . D i s cal ten tacles dendr i t i c , mar g inals s imple . Crown d iameter to 6 cm . Gener al co lor ochre,brown , or

green i sh brown (F ig . 4c) . Rhodact is howes i i Sav i lle-Kent

and one cer ianthar ian were co llected . Al l species wer eiden t ical to those fr om Enewetak . In French Po lynes ia(3 months ) , 1 4 act in iar ians , two cor allimorphar ians , and

one cer ian thar ian were co l lected . All species were iden t ica lto those fr om Enewetak . In H awai i (6 year s ), 25 act in iar i

ans , one corall imorphar ian , and one cer ian thar ian wer e col

lected. Of these, on l y s ix species were ident ical to thosefr om Enewetak .Wi th more co l lect ing at Enewetak , add i tional species of

sea anemones wi l l cer tainly be found . Th is speculation is

based on 10 species that are known fr om the G i lber tIs lands or Kwajalein that have not yet been foun d at

Enewetak .

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56 CUTRESS AND ARNESON

1 2 . Co lumn w i th inflatable, s imp le, or compound ves icles .

Co lumn smooth or w i th so l id war ts .

1 3 . Ves icles s imple, of d i fferent s i zes , scatter ed over pr ox imal half of co lumn . Ten taclesdec iduous . Column d iameter and co lumn length cm . Gener al co lor pa le green i sh br own , co lumnoften w i th alter nate s tr eaks of wh i te, yel low , and v io let (F ig . 4d) . Bunodeop s is meduso ides (Fowler )Ves icles br anched and compound , form ing a r ing ar ound m id-column . Tentacles not deciduous .

Cr own d iameter and co lumn lengt h cm . General colo r dar k brown (char treuse when car r ied byLybia tes se lata) (F ig . 4f) . Tr iact is p roducto Klunz inger

1 4 . Column smooth . Mar g in w i thout ves icles . 1 5

Co lum n w i th or w i thout war ts . Mar g in w i th ves icles . 1 6

1 5 . Tentacles to 4 cm , usual ly w i th subterm inal inflat ions . Crown d iameter to 8 cm . Co lumn off wh i te topale gr een ish br own to pink to v io let . Ten tacles gr een ish br own , t ips o ften wh i te (not i l lus trated) .

Physobr achia doug las i Savi l le-Ken tTen tacles to 4 cm , even ly tapered to po in t . M id-co lumn w i th cincl ides . Crown diameter and co lumnlength to 7 cm . General co lor pale to dar k o live brown (F ig . 1e) . A ip tas ia pulchella Car lgr en

1 6 . Marg inal ves icles pr om inen t . Co lumn w i thout war ts . Crown d iameter and column length cm . Gen

er al co lor r eddi sh br own , larger ves icles wh i te (F ig . 3b) . An thos tella badia Car lgr en

Column w i th r ows of pr om inent adhes ive war ts . 1 7

1 7 . Gener al co lor v io let black , war ts oft en wh i te. Cr own diameter to cm (F ig . 3a) . An thop leu r a n igr es cen s (Ver r i l l)Ten tacles br ight gr een to brown ish green . Column pale pink . Crown diameter to cm (F ig . 3e) .

Act inogeton seser e (H addon and Shackleton)1 8 . Co lumn elongate to verm i form , w i th pr om inent cut icle. 1 9

Co lumn shor t , cut icle inconspicuous . 22

1 9 . Ten tacles number 48 or more. 20

Ten tacles fewer than 48 . 21

20 . Ten tacles po in ted . Co lumn w i th th in sand-covered cut icle, lengt h to 4 cm , d iameter to cm(F ig . 1 c) . Telmatact is verm iform is (H addon )Ten tacles blun t to cap i tate. Co lum n w i th th ick , shaggy cut icle, length and diameter to and cm(F ig . 1a) . Telmatact is decora (Ehr enberg)

21 . Co lum n length and diameter 25 and cm (F ig . 4e) . Edwards ia pudica Klunz ingerCo lumn lengt h and d iameter to and cm (F ig . 4b) . Edwards ia gi lber tens is Car lgr en

22 . Column fi rm , lower par t w i th one or mor e r ings of pr om inent pores through wh ich salmon-co lor edthr eads are frequently extr uded . Co lor of column gr ay to r us ty or ange, often w i th s treaks and blotchesof dull plum r ed. Crown d iameter to 5 cm . L ives w i th herm i t cr abs (F ig . 1d) . Call iact is po lypus (For sskal)Colum n lack ing por es , wh i te to gr ay , oft en w i th fine s tr eaks of r us ty or ange.

often th icker than res t . Crown d iameter to 2 cm . L ives w i th herm i t crabs (F ig . 1b) . Ver r i llact is pagu r i (Ver r i l l)

Cer ian thar ia

So l i tar y , elongate, tube-dwel l ing po lyps , w i th tentacles d iv is ible into shor t labials and long marginals . On ly species r ecor dedfrom Enewetak :

Tentacles num ber ing to 56 ,mor e common l y 48 or 52 . Co lum n length to 1 0 cm , cr own d iameter to 6

cm . Gener al color r edd ish br own (copper ) (F ig . l f) . Isar achnan thus bandanens is Car lgr en

COLLECTION RECORDS

Act inodendr on g lomeratum H addon Act inod is cus neg lectus (Fowler )Enewetak : 60 feet , Sept . 1 980 . Lojwa: Sept . 1980 .

Act inodendr on p lumos um H addon Act inogeton sesere (H addon and Shackleton )Chop Top : Sept . 23 , 1 980 . Anan ij: May 8 , 1955 .

SEA ANEMONES 57

A ip tas ia pu lche l la Car lgrenMedr en : May 1 5 , 1 955 .

An thop leu r a n igr escens (Ver r i l l)Igur in : May 1 1 , 1 955 .

An thos tella badia Car lgren

Anan ij: May 1 1 , 1955 .

Bunodeops is meduso ides (Fowler )Anan ij, Enewetak : June 21 , 1 955 ;Chop Top : lagoon , 40 feet ,Sept . 22 , 1 980 .

Call iact is po lyp us (For s ské l)Medren : May 1 8 , 1 955 .

Cryp todendr um adhes ivum Klunz inger

Medr en : June 25 , 1 955 .

Edwards ia gi lber tens is Car lgrenBokoluo: 45 feet , Sept . 1980 .

Edwar ds ia pud ica Klunz ingerMedr en , Run i t : May 1 8 , 1 955 ;

Medr en : 50 feet , Sept . 1 980 .

H eteract is aur ora (Quoy and Gaimard)Bokoluo : Sept . 1 980 .

H eter act is ge lam (H addon and Shackleton)Medren : May 22 , 1 955 .

H eter act is macrodactylum (H addon and Shackleton)Japtan : Cemen t sh ip , 30 feet , Sept . 1 980 .

H eter act is r i tter i (Kw ietn iewski )Medr en : Sept . 1980 .

Plates 1 -4F ig . l a, Telmat act is decor a (E h r en ber g ). l ive Specimen fr om H awai i ; b, Ver r i l lact is pagu r i (Ver r i ll ), l ive specimen fr om H awai ic, Telmatact is verm iform is (H addon ), pr eser ved specimen fr om Majur o; d, Ca l l iact is po lypus (For ss kal ), l ive specimen fr om H awai ie, A ip tas ia pu lchel la Ca r lgr en , l ive specimen fr om H awai i ; f, Isa r achnan thus bandanens is Car lgr en , l i ve specimen f r om H awai i .

Page 58

F ig . 2a , M ega lact is hemp r ich i i Ehr enber g , l i ve specimen in s i tu , Enewetak , 40 feet; b, H eter act is r i t ter i (Kw ietn iewsk i ). l ive sped

men ln s i tu, Medr en . Enewetak , patch r eef, 10 feet; c, Cryp todendr um adhes ivum Klunz i nger , pr es er ved specimen fr om Enewetakd, H eter act is macr odactylum (H addon and Shack leton ), l i ve specimen i n s i tu, Cemen t s h ip, Enewetak , 30 feet ; e, St ichodactylamer tens i i B r andt, l i ve specim en i n s i tu , H en r y , outs ide r eef Enewetak , 30 feet : f. H eter act i s au r or a (Quoy and Gaimar d), l i ve sped

men in s i tu, Enewetak .Page 59

F ig . 3a, An thop leu r a n igr escens pr eser ved specimen fr om H awai i ; b, An thos tella bad ia Car lgr en , pr eser ved specimen f r omMaj ur o; c, H eter act is gelam (H addon an d Shack let on ), l i ve speci men i n s i tu, wi th Amphr ip r ion t i nctus , Bokoluo, Enewetak patchr eef , 40 feet; d, Act inodendr on p lumosa H addon , l ive speci m en i n s i tu, Chop Top , Enewetak. 30 feet; e, Acti nogeton ses er e (H addon and Shackleton ), l i ve specimen fr om H awai i ; f, Act inodendr on g lomer at um H addon , l ive specimen in s i tu , off Enewetak , 60

feet .Page 60

F i g . 4a, Act i nod iscus neg lectus (Fow ler ), l ive speci men in s i tu , Lajwa, Enewetak , 20 feet ; b, Edwards ia g i lber tens is Car lgr en ,

pr es er ved specimen fr om Maj ur o; c, R hodact is howes i i Savi l le-Ken t, pr eser ved specimen fr om Enewetak ; d, Bunodeops is

meduso ides (Fow ler ), l ive specimen fr om H awai i ; e, Edwards ia pud i ca Kl unz i nger , l ive specimen in s i tu , R un i t , Enewetak , 30 feet ;f. Tr iact is p r oducto K lunz i nger , l i ve specimen fr om H awai i .

Page 61

H eterodactyla hemp r ichi i Ehr enbergNo co llection data now at hand but foundin 1 955 and 1 980 .

Isarachnan thus bandenens is Car lgr en

Medren : May 1 5 , 1 955 ;

Bokoluo : 45 feet , Sept . 1 980 .

Megalact is hemp r ichi i EhrenbergEnewetak , Maggio : Sept . 1980 .

Phyman thus s trandes i Car lgren

Anan ij: June 1 5, 1 955 .

Physobr achia doug las i Sav i l le-Ken tAnan ij: June 1 6 , 1 955 ;

Bokoluo : Sept . 1 980 .

Rhodact is howes i i Sav i l le-KentMedren : May 22, 1 955 .

St ichodactyla mer tensu Br andtH en r y , Enewetak : Sept . 1 980 .

Telmatact is decor a (Ehr enberg)Anan ij: May 8 , 1955 ;

Enewetak : 60 feet , October 1 7 , 1 980 .

Telmatact is verm iform is (H addon)Medr en : May 1 5, 1955.

Tr iact is p roducto Klunz ingerAnan ij: June 8 , 1 955 .

Ver r i llact is pagu r i (Ver r i l l)Medr en : May 1 8 , 1 955 .

REFERENCES

CUTRESS AND ARNESON

A llen , Ger ald R . , 1972 , The Anemone/ ishes : Their C lass ificationand B io logy , Neptune City , N . J. , T . F . H . Pu bl . Inc . L td. ,

288 pp .

Dunn , Daphne Fau t in , 1981 , The Clown f ish Sea Anemones :

St ichodactyl idae (Coe lenterata: Actin iar ia) and Other Sea

Anemones Symbiotic with Pomacentr id Fishes , Trans . Am .

Ph i los . Soc . , 71 , Pt . 1 : 1 - 1 15 .

Johannes , R . E et ai . , 1 972 , The Metabo l ism of Some Co ralReef Commun i ties : A Team Study of Nutr ient and EnergyF lux at En iwetok , B ioscience, 22 541 -543 .

Josephson , Rober t K . , 1966 , Neur omuscu lar T r ansm i ss ion in a

Sea Anemone, J . Exp B io l . , 45 305-3 1 9 .

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64 DEVANEY

TABLE 1

Check l i s t of O ctocor al l ia of En ewetak A tol l

Phy lum COELENTERATA

Class ANTH OZOASubc lass OCTOCORALL IAOrder STOLON IFERAFam i ly CLAVULARIIDAEPachyclavu lar ia vio lacea (Quoy and Gaimard,

Ver seveldt , 1 972; H umes , 1 973 .

Fam i ly TUB IPORIDAETubipo ra mus ica L innaeus , 1 758 : Wel ls , 1 954 .

O rder TELESTACEAFam i ly TELESTIDAETeles to sp .

Order ALCYONACEAFam i ly ALCYON IIDAECladiella pachyclados (Klunz inger ,Lobophytum bo r bon icum (Von Mar enzeller

Ver seveldt , 1 977 .

Lobophytum cataloi T ix ier -Du r ivau lt , 1 957 :Ver seveldt , 1 977 .

Lobophytum den t icu latum T ix ier -Du r ivau lt , 1 956 :

Ver seveldt , 1977 .

Lobophyt um pauc iflor um (Eh renberg ,Odum and Odum ,

1 956 ; Ver seveldt , 1 972 .

H umes , 1 973 ; Ver seveldt , 1 977 .

Sar cophyton g laucum (Quoy and Gaimard,

Ver seveldt , 1 977 .

Sar cophyton tr ochel iOpho r um Von Marenzeller ,

1 886 : Ver seveldt , 1 972 ; 1 977 .

Sin u lar ia densa (Whi telegge, Ver seveldt ,

1 980 . Sin u lar ia bronger smai Ver seveldt , 1 972 :

Ver seveldt , 1 972 .

Sinu lar ia lep toc lados (Ehr enberg ,

Ver seveldt , 1 977 .

Sin u lar ia marenzeller i (Wr ight and Studer ,Ver seveldt , 1 972 .

‘New Enewetak record .

dr edged from depths of 1 00 m or deeper . One spec ies ,recor ded

,

as Subergo r g ia m o l l is , is ver y close taxonom i

cal l y to that determined as S . r et iculata from Enewetak ,but fu r ther wor k i s necessar y before thi s ques tion is

r eso lved (Bayer , per sonal commun icat ion) . Otherw ise, none

of the spec ies so far iden t i f ied i s common to the two

ato l ls . A t leas t one more shallow-water gor gonacean , as

yet un r ecor ded fr om Enewetak , was descr i bed from Ar noAto ll in the souther n Mar shal l Is lands (Bayer , The

gorgonacean Rumphella an t ipathes has been ut i l i zed fromspec imens co l lected at Enewetak for b iochem ical s tud ies(Cier es zko et al . , 1 968 ; Mar sh and Cier eszko ,

Dur i ng submer s ib le d ives off the seawar d s ide of threei s lets (B iken , Mut , and Enewetak) dur ing July 1 98 1 , the

author obser ved nephtheids and go rgonaceans as commonfaunal elemen ts at depths from appr ox imatel y 80 to

180 m . Sever al of these , as yet undeterm ined , wer e col

lected and wer e depos i ted at the B i shop Museum .

The clas s i ficat ion of octocorals used her ein fo l lows thatby Bayer although a r ev i sed class i fi cat ion has

recen t ly been pr oposed (Bayer ,

ACKNO“ &EDGMEN TS

J . Ver seveldt was mos t helpful in r ev iewing the d iscuss ion and checkl i s t on the alcyonaceans and al low ing the

inclus ion of the five new r ecor ds that he determ ined .

F . M . Bayer (USNM) pr ov ided updated names for the

mater ial co l lected by Go ldberg and was mos t helpful inperm i tt ing thei r inclus ion her e .

REF ERENCES

Bayer , F . M . , 1 949 , The A lcyonar ia of B ikin i and Ot her Ato l ls inthe Mar sha l l Gr oup , Par t I : the Gorgonacea, Pac . Sci . , 3

1 95-2 1 4 .

1 955 , Contr ibut ions to the Nomenclature, Systematics , an (

Fam i ly ALCYONIIDAE (Cont ’d)Sinu lar ia po lydactyla (Ehr enberg ,

Ver seveldt , 1 972 ; H umes , 1 973 .

Sinu lar ia r igida (Dana,Ver seveldt , 1977 .

Sin u lar ia sandens is (Ver seveldt ,Ver seveldt , 1 977 .

Fam i ly NEPH TH EIDAEDendronephthya (Roxas ia) m i rabi l is H ender son ,

1 909 : Ver seveldt , 1 977 .

Nephthea chabr ol i i Audou in , 1828 : Ver seveldt ,

1972 ; H umes , 1 973 .

Nephthea pacif ica Kuken thal ,Nephthea albida (H o lm ,

Stereonephthya un ico lor (Gray ,Fam i ly XEN IIDAEH eteroxen ia cohen i Ver seveldt , 1974 .

Order COENOTH ECAL IAFam i ly H EL IOPOR IDAEH eliopo ra coeru lea (Pal las , Wel ls 1954 .

Order GORGONACEASubo rder SCLERAXONIAFam i ly SUBERGORG IIDAESubergor gia nu tt ing i St iasny, 1 937.

Subergor gia r et icu lata (Ellis and So landerSuber gor gia suber osa (Pal las ,

Subo rder H OLAXONIAFam i ly ACANTH OGORG IIDAEAcan thogo r gia sp .

Fam i ly PARAMURICEIDAEAs t rogorgia sp .

As t rogorg ia? sp .

Fam i ly PLEXAURIDAERumphella an t ipathes (L innaeusFam i ly ELLISELL IDAEEll isella sp .

CHALLIA 65

Mar sh , M . E . , and L . S. Ciereszko , 1 973 , AlphaG lyceryl Ether sin Coe lenter ates , Proc . Ok lahoma Acad . Sc i . , 53 : 53 -56 .

Odum , E. P . , and H . T . Odum , 1956 , Zonation of Corals on Japtan Reef, En iwetok Ato l l , A to ll Res . Bu ll . No . 52 , pp . 1 -3 .

Schm itz , F . J. , D . J . Vanderah, and L . S. Ciereszko , 1974 ,

Mar ine Natur al Products : Nephenol and Epoxynephenol Acetate, Cembrene Der ivatives fr om a Soft Coral , Chem . Com

m un . , 1 974: 407-408 .

Vanderah, D . J. , N . Rut ledge, F . J . Schm itz , and L . S .

Ciereszko , 1 978 , Mar ine Natur al Products : Cembr ene-A and

Cembr ene-C fr om a Soft Cor al , Nephthea sp . , J . Org . Chem ,

43: 1 614- 1 61 6 .

Ver seveldt , J. , 1 972, Repor t on a Few Octocorals fr om En iwetokAto l l , Mar shal l Is lands , Zoo l . Meded . , 47: 457-465 .

1 977 , Octocorall ia from Var ious Local it ies in the PacificOcean , Zoo l . Ver h . (Leiden) N o . 1 50 ,

pp . 1 -42 .

1 980 , A Rev is ion of the Genus Sinu lar ia May (Octocoral l ia,

A lcyonacea), Zoo l . Verh . (Leiden) No . 1 79,pp . 1 - 1 28 .

Wel ls , J. , 1954 , Recent Corals of the Mar shal l Is lands , U . S .

Geo l . Sur vey Prof . Pap . 260 -I, pp . 385-486 .

Chapter 8

Enewetak Ato ll

the r efer ence co l lect ion w i th a set of labeled coral specimens r epr esen tat ive of each major r eef habi tat at

Enewetak and (b) pr epar e a’

reef cor al field iden t i ficationgu ide for gener al use by scientis ts wor k ing at Enewetak . I twas also hoped that the specimens and accompanying fieldobser vat ions would be used in indiv idual sys temat ic publ icat ions of the Enewetak scler act in ians .

To date, appr oximately 1 70 species in 54 genera havebeen ident i fied among the 2500 s tony cor al specimenswh ich wer e co l lected fr om 28 s i tes ar ound Enewetak Ato l l .These have been depos i ted in major m useums (B . P .

B ishop Museum , U n i ted States Nat ional Museum , B r i t ishMuseum [Natur al H is tor y], R i jksmuseum van Natuu r l ijke

H is tor ie, Queens land Museum), as wel l as in the Enewetakr efer ence co l lect ion .

A lthough the field gu ide is s t i l l in preparat ion , a prov is ional checkl i s t of scler act in ian cor als now bel ieved to

occur at Enewetak is pr esen ted in Table 1 . In man y famil ies , the species -Ievel class i fication general ly fol lows thatgiven in the r ecent publ icat ions about eas ter n Aus tr al ianscler act in ians by Ver on and P ichon (1 976 , 1 980 ,

by Ver on ,P ichon , and W i jsman -Bes t and by Ver on

and Wal lace Eco logi cal and zoogeog r aphic con

s iderat ion s w i l l awai t the completed gu ide .

Ston y corals co l lected at Enewetak dur ing the

wo r kshop have con tr i buted to several r ev is ionar y ‘

sys

temat ic s tud ies . For example, r ange extens ions were madeby Wallace (1 978) for spec ies of Acr opora; by D inesen( 1 980) for a species of -Lep toser is ; and by W i jsman-Bes t(1 980) fo r species in three fav i id gener a (Cyphas t rea, Leptas tr ea, Echinopo r a) . Lam ber ts 1 982) has also usedEnewetak specimens to rev ise the genus As t reopo ra. Six

of the n ine spec ies of As tr eopo r a (two of wh ich ar e new

species ) r ecogn i zed by Lamber ts (1 982) have been col

lected at Enewetak , wh i le two other s occur elsewher e in

the Mar shal l Is lands .

In addi t ion to these s ys tematic s tudies , ther e are

numerous publ icat ions on scleract in ian bio logy or geo logywh ich have resulted fr om field wor k and col lect ions madeat Enewetak . A select ion of these refer ences , wi th the genera exam ined , is presented in Table 2 . The author s of

these nons ys temat ic paper s ar e also c i ted in Table 1 underany species names given for these gener a.

68 DEVANEY AND LANG

TABLE 1

Check l i s t of Ston y Cor al s of Enewetak A tol l

Order SCLERACTINIASubo rder ASTROCOEN IINAFam i ly ASTROCOEN IIDAEStylocoen iella armata (Ehr enberg , H i ldemann et al . , 1975a.

‘1

’

Stylocoen iella guen ther i (Bassett -Sm i th ,Fam i ly TH AMNASTERIIDAE"l'Psammocora con t igua (Esper ,Psammocora digi tata Mi lne Edwards and H aime, 1851 .

Psammocora (Stephanar ia) togianens is Umbgr ove, 1940 : Gar th , 1964 ;Knutson et ai . , 1 972 ; Buddemeier et al . , 1974 . Synonymy after Veron and Pichon

‘Psammocor a exp lanu lata van der H or s t , 1922 .

‘Psammocora n ier s t r az i van der H or st. 1 921 .

°'

l'Psammocora p rofu ndacella Gard iner , 1 898 .

Fam i ly POCILLOPORIDAEPoc i llopo ra dam icor n is (L innaeus , Gar th , 1 964 ; Johannes et al . , 1972a; Clausen and Roth , 1975 ;

Co les and Jok iel , 1977; Stim son , 1978 ; R ichmond, 1982; Roth et al . , 1982 ; R ichmond and Jok iel , 1984 .

Poc i llopo ra setchell i H offmeister , 1929: Wel ls , 1954 . Synonym y v ia J . Veron (per sonal commun ication) .Poc i llopo ra eydoux i M i lne Edwards and H aime, 1860 : Gar th , 1 964 ; Pomer oy and Kuenzler , 1969 ;

Buddemeier et al . , 1 974 ; H i ldemann et al . , 1 975a, 1 975b.

Poc i llopo ra grandis Dana, 1 846 : H i ldemann et al . , 1975a, 1 975b. Synonym y after Veron and PichonPociIIOpo ra meandr i na Dana, 1 846 : Banner and Banner , 1 968 ; Muscatine and D

’

El ia, 1978 .

Poc i llopo ra e legan s Dana, 1846 : Wel ls , 1954 ; Odum and Odum , 1956 ; Gar th , 1964; H i ldemann et al . , 1975a,

1975b; Co les et al . , 1 976 ; D ’

El ia and Webb, 1977 ; Stim son , 1 978 . Synonymy v ia J . Veron (per sonal commun ication).Poc i llopo r a ver r ucosa (E l l i s and So lander , Wells , 1 954 ; Odum and Odum , 1956 ; Gar th , 1964 ; Pi lson , 1 974 ;

H i ldemann et al . , 1 975a, 1975b; Stim son , 1978 .

Poc i llopo ra danae Ver r i l l , 1 864 : Odum and Odum , 1 956 ; Knudsen , 1967 ; D’

Elia, 1 977 . Synonymy v ia J . Veron(per sonal commun ication ).

Ser iatopo ra hys tr ix Dana, 1 846 : Wel ls , 1954 ; Gar t h , 1964 ; Young , 1971 ; H i ldemann et al . , 1975a; Stimson , 1978 .

Ser iatopo ra angu lata Klunz inger , 1 879 : We lls , 1 954 . Synonymy after Veron and P ichonStylophora p is t i llata (Esper ,Stylophora mordax (Dana, Yabe et ai . , 1936 ; Wells , 1954 ; Odum and Odum , 1 956 . Synonym y after Veron

and PichonFam i ly ACROPORIDAE‘TAcr opo ra acu leus (Dana,

Acropo ra acum inata (Ver r i l l , Wel ls , 1 954 ; Gar th , 1 964 (as A . formosa or A . acum inata H i ldemann et ai . ,

1975a, 1975b.

Acropo ra asper a (Dana, Stimson , 1978 .

Acropo ra hebes (Dana, Wells , 1954 . Synonymy after Wal laceTAcropo ra aus tera (Dana, Wal lace, 1 978 .

‘TAcropo ra cereal is (Dana,

‘Acropo ra cuneata (Dana,

Acropo ra cythera (Dana, Wallace, 1978 .

Acropo ra ret icu lata (B rook , Vosburgh , 1 977 , 1978 , 1982 . Synonym y after Wal laceAcropora danai (M i lne Edwards and H aime,

Acr opo ra rotumana (Gardiner , Wells , 1954; H i ldemann ci al . , 1975a, 1975b . Synonym y after Veronand Wal lace

Acropo ra digi t ifera (Dana, St imson , 1978 .

Acr opo ra ech inata (Dana, Well s , 1954 ; Gar th , 1964 ; H i ldemann et al . , 1975a.

‘l'Acr opo ra flor ida (Dana, H umes and Stock , 1973 ; H i ldemann et al . , 1 975a; H umes , 1 981 ; H umes and Doj i r i , 1982 .

Acropo r a formosa (Dana, We lls , 1 954 ; Gar th , 1 964 (as “

A . formosa or A . Cheney , 1 975 ;H i ldemann et al . , 1 975a, 1 975b; Co les et al . , 1 976 ; Roth et al . , 1 982 .

Acr opo ra gemm ifer a B r ook , 1 892 : Odum and Odum , 1 955 ; in par t as Acr opo ra hum i lis (Dana, 1 846) in Wells ,1 954 ; in par t in : Odum and Odum , 1 956 ; Gar th , 1 964 ; H i ldemann et al . , 1975a, 1975b; Stimson , 1 978 .

Rev ised by Veron and Wal laceAcropo ra gr anulosa (M i lne Edwards and H aime,

Acr opo ra rayner i (Br ook , Wel ls , 1 954 . Synonym y after Wal lace

‘New Enewetak r ecord .

TNo t r ecorded from Mar shal l Is lands by th is name in Wells (T h i s table continued on next page.

SCLERACTIN IA (STONY CORALS) 69

TABLE 1 (con t’d)Fam i ly ACROPORIDAE (cont’d)TAcr opo ra hor r ida (Dana ,

Acr opo ra angu lata (Quelch , H i ldemann et al . , 1 975a. Synonymy after Wal laceAcr opo ra hum i l is (Dana, in par t in Wells , 1954 ; in part in : Odum and Odum , 1956 ; Gar th , 1964 ;

H i ldemann et al . , 1 975a, 1 975b; Stim son , 1 978 . Revi sed by Veron and Wal laceAcr opo r a hyacin thus (Dana, Wells , 1 954 ; Gar th , 1 964 ; Pomer oy and Kuenz ler , 1 969 ; Bruce , 1 969; Co les et al . , 1 976 ;

Sm i th and H ar r i son , 1 977 ; D’

El ia, 1 977; Stimson , 1 978 .

Acr opo ra corym bosa (Lamarck , in par t in Well s , 1 954 ; in part in : Odum and Odum , 1 956 ; Gar th , 1 964 ; St imson , 1 978 .

Synonym y after Ver on and Wal laceAcr opo r a recumben s (B rooks , Odum and Odum , 1 956 . Synonymy after Wal laceAcropo r a? sur culosa (Dana, Gar th , 1 964 ; H i ldemann et al . , 1 975a . Synonymy after Veron and

Wal laceAcr opo r a lor ipes (B rook , Young , 1 97 1 ; H i ldemann et al . , 1 975b.

Acr opo ra m icr ophthalma (Ver r i ll , Wal lace, 1 978 .

Acr opo ra m i l lepo r a (Ehr enberg ,Acropo ra cor ymbosa (Lamarck , in par t in Wells , 1 954 ; in par t in : Odum and Odum, 1 956 ; Gart h , 1 964 ;Stimson , 1978 . Synonym y v ia C. Wal lace (per sonal commun ication) .

Acr opor a nasu ta (Dana, Wells , 1 954 ; H i ldemann et al . , 1 975a, 1 975b; Stimson , 1978 .

Acr opo ra cymbicyathus (B r ook , Wells , 1 954 ; Odum and Odum , 1 955; Gar th , 1964 . Synonym y afterWal lace

Acropo ra pal ifera (Lamar ck , Wells , 1 954 .

Acr opo ra palmer ae Wel ls , 1 954 : Odum and Odum , 1 955 ; B ruce, 1 969 ; Co les et al . , 1 976 .

Acropo ra pan iculata Ver r i ll , 1 902 : Gar th , 1 964 .

”IAcropo ra robus ta (Dana,

Acr opo ra samoens is (Brook , in par t as Acr opo r a humu l is (Dana, 1 846) in Wells , 1 954 ; in par t in : Odumand Odum , 1 956 ; H i ldemann et al . , 1 975a, 1 975b ; St imson , 1 978 . Rev ised by Ver on and Wal lace

Acr opo ra secale (Studer ,Acr opo r a diver sa (B r ook , Wel ls , 1 954 . Synonymy after Ver on and Wallace

‘TAcr opora

“

sp . ca l led A . po lymo rp ha (B rook , 1 891 )Acr opo ra sp .

?

Acropo ra confer ta (Quelch , Wells , 1 954 ; Odum and Odum , 1956 ; Go r eau , 1959 ; H ildemann et al . ,

1975a. Equals A . hyacin thus (Dana, 1 846) or A . m i l lepo ra (Ehr enberg , 1 834) (C . Wal lace, per sonal commun ication ) .Acr opo ra del icatu la (B rook , Co les et al . , 1 976 .

Acropo ra gr andis (Br ook , H i ldemann et al . , 1 975a Pr obably not th is species (C. Wallace,

per sonal commun ication ) .Acr opo ra syr ingo ides (B rook , Stimson , 1 978 .

Acropo ra s t r iata (Verr i l l , Wel ls , 1 954 ; Stimson , 1 978 .

°

Acropo ra tenel la (B rook ,°

Acropo ra tenuis (Dana,

Acr opo r a teres (Ver r i l l , Wel ls , 1 954 .

Acr opo r a tubicinar ia (Dana, Odum and Odum , 1 956 .

Acropo r a va l ida (Dana,

‘

Acr opo r a var iabi l is (Klunz inger , Gar t h , 1 964 ; H i ldemann et al . , 1 975a. Synonym y after Veronand Wal lace

‘Acr opo ra vaughan i Wells , 1 954 .

TAs t reopora cucu l lata Lam ber ts , 1 980 : Lamber t s , 1 982 .

As t reopo ra gr aci l is Ber nard , 1 896 : Lam ber ts , 1 982 .

As t r eopo ra l is ter i Ber nard , 1 896 : Lamber ts , 1 982.

As tr eopo r a myr iophthalma (Lamar ck , Wells , 1 954 ; Odum and Odum , 1 956 ; Buddemeier et al . , 1 974 ;

H ighsm ith , l 980b, 1 98 1 b; Lamber ts , 1 982.

TAs t reopo r a scabr a Lam ber ts , 1 982 .

As t reopo ra sugges ta Wel ls , 1 954 : Lam ber t s , 1 982 .

TMon t ipor a aequ itubu r cu lata, Ber nard, 1 897 .

Mon t ipo r a compos ita Cross land , 1 952 : Odum and Odum , 1 956 .

Mon itpora?ver r i l l i Vaughan , 1 907 : Odum and Odum , 1 956 ; H ighsm ith , 1 980b.

Mon t ipo ra?ver r i li [s ic]Vaughan : H i ldemann et al . , 1 975a .

”

r Mon t ipo r a aus t r al ien s is Ber nard , 1 897 .

TMon t ipo ra ber ryi Be rn ard , 1 897 : H ighsm ith , 1 980b, 1 98 1a .

‘Mon t ipo ra cal icu lata (Dana ,

‘New Enewetak r ecord .

TNot recorded fr om Mar shall Is lands by th is name in Wel ls (T h is table cont inued on next page

70 DEVANEY AND LANG

TABLE 1 (con t’d)

Fam i ly ACROPORIDAE (cont ’d)‘Mon t ipor a con icu la Wel ls , 1954 .

‘Mon tipo r a danae Mi lne Edwards and H aime, 1 851 .

“

r Monn'

pora digi tata (Dana,

‘Mon t ipo ra ehrenbergi i Ver r i ll , 1 875 .

‘Mon t ipo ra flower i Wells , 1 954 .

‘TMon t ipo ra fo liosa (Pal las ,Mon tipor a [oveolata (Dana, Odum and Odum , 1956 .

Mon tipor a social is Bernard, 1 897 : Wel ls , 1954 ; H ighsm ith , 1 980b . Synonymy aft er Ver on and Wal lace0 “

I’

Mon t ipor a h isp ida (Dana,

Mon tipor a hoffmeis ter i Wells , 1954: H ighsm i th , l 98ob.

Mon tipor a inform is Bernard, 1 897 : H i ldemann ci ai . , 1 975a.

"I“

Mon tipor a lobu lata Ber nard , 1 897 .

‘Mon tipo ra mar shallens is Wells , 1954 .

‘TMon ti po ra sp . cf . M . m onas ter iata (For skal ,‘Mon tipor a myr iophthalma Bernard, 1 897 .

‘TMon tipor a po rosa Ber nar d , 1 897 .

Mon t ipor a sp .?

Mon tipo ra gr anu losa Ber nard, 1 897 : H ighsm ith , 1 980b.

‘Mon t ipo r a tu berculosa (Lamar ck ,Mon t ipor a venosa (Ehr enberg , Wells , 1 954 .

Mon tipor a ver r ucosa (Lamar ck, H i ldemann et al . , 1975a; Co les and Jokiel , 1977 .

Suborder FUNG IINAFam i ly AGARICIIDAE°

Gardineroser i s p lanu lata (Dana, As Pavona (Polyas tra) p lanu lata (Dana, 1 846) fr om Arno in Wells Synonymyafter Veron and Pichon

Lep toser is hawa i iens is Vaughan , 1907 : Dinesen , 1 980 .

Lep toseri s mycetoser o ides Wel ls , 1954 : Di nesen , 1980 .

‘Pachyser i s speciosa (Dana,

Pavona cactus (For skal ,Pavona p raetor ta (Dana, Wethey and Por ter , 1976a, 1 976b. Synonymy after Veron and PichonPavona clavus (Dana , H ighsm ith , 1981a, 1981b .

‘Pavona maldivens is (Gar diner , As P . (Pseudocolum nas tr aea) po ll icata Wells , 1954 fr om Bik in i and Ar no(Wells , Synonym y after Ver on and Pichon

‘TPavona pr ismatica B r i iggemann , 1 879 .

Pavona var ians Ver r i l l , 1 864 .

Pavona venosa (Ehr enberg ,Fam i ly SIDERASTREIDAE

‘Cosc inar aea co lumna (Dana,

Fam i ly FUNG IIDAE‘1'Cyclos er is erosa (D6der lein ,

Cycloser is hexagonal is Mi lne Edwards and H aime, 1 849 : Wells , 1954 .

Fungia (Ctenact is) ech inata (Pallas ,Fungia ech inata (Pallas , Young , 1 971 ; H i ldemann et al . , 1 975a, 1 975b.

‘TF ungia (Danafungia) va lida Ver r i l l , 1 864 .

Fungia (Fungia) fu ngi tes (L inneaus ,Fungia fungi tes (L innaeus , Pomeroy and Kuenzler , 1 967; Buddemeier et al . , 1974 ; H i ldemann et al . ,

1 975a, 1975b .

‘Fungia (Pleu ract is) paumotens is Stuchbur y , 1 833 .

Fungia (Pleu ract is) scu tar ia Lamar ck , 1 80 1 .

Fungia scu tar ia Lamar ck , 180 1 : Wells , 1954 ; Co les et al . , 1976 ; D ’

El ia, 1 977‘Fungia (Ver i llofungia) concinna Ver r i ll , 1864 .

'Fungia (Ver r i llofungia) r epanda Dana, 1 846 .

‘H alom it r a p i leus (L innaeus , As H . ph i lipp inens is Studer 190 1 fr om Jalu i t and Ar no (Wel ls , Synonym y.after VeronPichon

‘N ew Enewetak r ecord .

TNot recorded fr om Mar shal l Is lands by th is name in Wells (This table continued 0"next page.)

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72 DEVANEY AND LANG

TABLE 1 (con t'd)

Fam i ly FAVIIDAE (cont ’d)°

H ydnophora r igida (Dana,

I’

Lep tas t rea pruinosa Cr oss land, 1952 : W i jsman-Best , 1 980 .

Lep tas t rea purpurea (Dana, Pomeroy and Kuenz ler , 1969 ; W i jsman-Bes t , 1 980 .

Lep tas trea transver sa Klunz inger , 1 879 : W i jsman-Best , 1 980 .

Lep tor ia phr ygia (El l is and So lander , Yabe et ai . , 1936 ; Wells , 1954 .

°'

l'

Mon tas t raea ann u l igera (M i lne Edwards and H aime,

‘l'

Mon tas traea cur ta (Dana,

Ou lophyll ia cr ispa (Lamarck , H ighsm ith , 1 98 1a, 198 1b .

Ou lophyll ia aspera (Quelch , Buddemeier et al . , 1 974 . Synonymy af‘ter Veron et al .

Platygyra daedalea (El l is and So lander ,Platygyra r us t ica (Dana, Wel ls , 1 954 ; Odum and Odum , 1956 . Synonym y after Veron et ai .

l Platyg yra lamell ina (Eh ren berg ,Buddemeier et ai . , 1974 ; H ighsm ith , 1 980b, 1 98 1 b .

"I'Platygyra p in i Cheval ier , 1975 .

‘Platygyra s inens is (M i lne Edwards and H alme,

Ples ias tr ea ver s ipo ra (Lamarck , Yabe et al . , 1936 ; Wells , 1954 ; Odum and Odum , 1 956 .

Fam i ly RH IZANG IIDAE‘Cu licia r ubeo la (Quoy and Gaimard,Fam i ly OCUL IN IDAE

‘Acr hel ia hor rescen s (Dana,

Fam i ly MERULINIDAE‘Mer u l ina amp liata (Ellis and So lander ,

‘Scapophyll ia cy lindr ica M i lne Edwards and H aime, 1 848 .

Fam i ly MUSSIDAE‘Acan thas trea ech inata (Dana,

Lobophyll ia cor ym bosa (For skal , Odum and Odum , 1956 ; Young , 1971 .

‘Lobophyll ia hemp r ichi i (Ehr enberg ,

"I

“

Scolym ia sp . cf . S . vit iens is Br ii ggemann , 1 877.

‘1‘Symphyll ia agar icia Mi lne Edwards and H aime, 1 849 .

“I'Symphyll ia radians M i lne Edwards and H aime, 1 849 .

Symphyll ia sp .?

Symphyll ia nobi l is (Dana, Wel ls , 1 954 . Veron and Pichon (1 980) cons idered th is species to be a jun ior synonym of

Symphyll ia cf. recta (Dana,

Fam i ly PECTINIIDAE‘Echinophyll ia aspera (El l is and So lander ,

"f Pect in ic lactuca (Pal las ,

Suborder CARYOPHYLLIINAFam i ly CARYOPHYLL IIDAE°

Euphyll ia g labrescen s (Cham iss o and Eysenhardt ,°

Physogyra lichtens tein i (M i lne Edwards and H alme,

"I Plerogyra s inuosa (Dana,

Suborder DENDROPH YLL IINAFam i ly DENDROPHYLL IIDAE°

Endop samm ia sp .

”

l Tubas tr aea coccinea Lesson , 1 829.

‘Tu r binar ia crater (Pallas , B . R . Rosen and R . A . K inzle (per sonal commun ication) work ing wi th Enewetak Tu r binar ia on ly

recogn i ze th is species among a number of specimens exam ined .

Tu r binar ia danae Ber nard, 1 896 : Gar t h , 1 964 ; H i ldemann et al . , 1 975a, 1 975b . Cons idered to be a jun ior synonym of

Tu rbinar ia f rondens (Dana, 1 846) by Veron and P ichonTu r binar ia g lobu lar is Bernard, 1 896 : Odum and Odum , 1 956 . Cons idered to be a jun ior synonym of Tur binar ia

s tellu lata (Lamar ck , 1 8 1 6) by Veron and PichonTu r binari a mesen ter ina (Lamarck , Odum and Odum , 1 955 . Cons ider ed to be a val id species by Veron and Pichon

‘New Enewetak r ecord .TNot r ecorded fr om the Mar shal l Is lands by th is name in Wel ls

SCLERACTINIA (STONY CORALS)

TABLE 2

73

Selecti on s of N ons ys tema ti c Ston y Cor al Resea r ch at Enewetak A toll‘

T ype of s tudy

Skeletal organ iccompos ition

Asexual di sper sa lB ioeros ion

Productiv ity andtr oph ic s tructure

Radionucl ides inskeletons

Cor al gener a‘

B ioc hem i s tryAcropo ra, Fungia, H erpo l itha, Lobophyll ia,

Ser iatopora, Tu r binar ia

EcologyPor itesAs t reopo ra, Favia, Gon ias t r ea, Mon t ipo ra,

Ou lophyll ia, Pavona, P latygyr a,Por i tes

Acropo raAcr opo ra sp . , As treopo ra sp . , Favia sp . , Favites sp . ,

Fungia sp . , Mon tipo ra sp . , P les ias t rea sp . ,

Psammocor a sp . , Symphyll ia sp . , Tu r binar ia sp .

Acropo ra sp . , Poc i llopo ra, Po ri tes sp .

Acr opo r aCyphas t r ea

Acropo r a, Poc i llopo r a, Por i tes , Psammocora,

Ser iatopo r a, Tu r binar ia

As tr eopo ra, Favia, Gon ias t r ea, Mon t ipora,

Ou lophyll ia, Pavona, P latygyr a,Por ites

Acr opo ra

Acropo ra sp . , Poc i ll0po r a

Acropo r a sp . , Fungi a sp . , Poc i llopo r a

Acropo ra, As t reopo r a sp . , Cyphas tr ea,

Favia, Lepas treaj Lobophyll ia sp . ,

Mon t ipo ra sp . , Poc i llopo r a sp . , Por i tes ,Stylophor a sp . , Tu r binar ia

Acropo ra, As t reopo ra, Cyphas tr ea,

Echinopo r a, Favia, Favites ,

Gon ias tr ea,Lobophyll ia, Mon t ipo ra,

P latygyr a, Ples ias trea, Poc i llopor a,

Por ites , Stylophora, Tu r binar ia

Acr opo r aPavona

Geochem i s tryAs t r eopo r a, Favia, Fungia,

Gon iast rea, H erpo l itha, H ydnophor a,

Ou lophyllia, Parahalom i tra, P latygyra,

Poc i llopo ra, Por i tes , PsammocoraAs treopo ra sp . , Favia, Gon ias t r ea, Por ites

Acr opo r a sp . , Favia sp . , Mon t ipor a sp . ,

Platygyr a sp . , Poc i llopo r a sp . , Por ites sp . ,

Psammocor a sp .

Refer en ce

Young , 1 971

H ighsm ith , 1 980aH ighsm ith , l 980b, 1981a,

1 98 1 b, 1 98 1 c

Vosburgh , 1 977, 1 978 , 1 982

Johannes et al . ,1 972a

Banner and Banner , 1 968

B ruce, 1969

B ruce, 1 979

Gar th , 1964

H ighsm i th , 1 980b,1 980c, 1 98 1 b, 1 98 1 c

H umes , 1 98 1 ; H umes andDoj i r i , 1 982 ; H umes andStock , 1 973Knudsen , 1967

Johannes et al . , 1972bOdum and Odum , 1955

Odum and Odum , 1 956

Sm ith and H ar r i son ,1977

Wethey and Por ter , 1976a,

1 976b

Buddemeier et al . , 1 974

Knutson and Buddemeier ,1 973

H agger ty et al . , 1980

Buddemeier et al . , 1 982

‘Any species names g iven fo r each of these genera can be found in the checkl ist of corals (TableTReclass i f ied as a species of Ples ias t rea in Odum and Odum

(Th is table continued on next page. )

DEVANEY AND LANG

TABLE 2 ( con t'd )

Cor a l gener a‘

ImmunologyAcropor a, Cyphas t rea sp . , Favites ,

Fungia, Mon t ipo ra, Poc illopora,

Pori tes , Psammocora sp . , Ser iatopo ra,

Sty locoen iella, Tu r binar ia

Phys iologyAcr opor aAcropo raAcropo r a, Fungia, Poc i llopor a

Acropo ra sp . , Poc i llopo ra, Por ites

Acropor a, Fungia, Lep tas t rea,

Poci llopo r a, Pori tesAcropo ra sp .

As tr eopo ra, Favia, Fung ia,

Gon ias tr ea, H erpol i tha, H ydnophora,

Ou lophyll ia, Par ahalom i t ra, P latygyr a,

Poci ll0 pora, Por i tes , PsammocoraFavia, Gon ias t rea, Por i tesFaviaFavia, G on ias tr ea, Por i tes , PsammocoraAcr opo r a, Poc i llopor a

Acropor a sp . , Por ites sp .

Mon t ipora, Poc i llopo r a

Acropor a, Fungia, Lep tas t rea,

Por i tes , Poc i llopo r aSexual Reproducti on

Poc i llopor a

Poc i llopo r a

Acr opo ra, Poci llopo ra, Ser iatopo r a

Refer ence

H i ldemann et al . ,

1 975a, 1 975b

H ighsm ith , 1 979H agger t y et al . , 1980Knutson et al . , 1972

Roth et al . , 1982Co les and Strathmann , 1 973

Clausen and Roth , 1 975

Co les and Jok iel , 1 977Co les et al . , 1 976

R ichmond, 1 982R ichmond and Jok iel , 1984Stimson , 1 978

‘Any species names given for each of these genera can be found in the check l i st of corals (TableTRecIass i f ied as a species of P les ias t r ea in Odum and Odum

74

Type of s tudy

Immunorecogn i t ion

Growt h rate

ACKNOWLEDGMEN TSWe ar e gr ateful to par ticipants of the 1 976 reef co ral

wor kshop for perm i tt ing inclus ion of many of the new

r ecor ds of cor als , wh ich were based for the mos t par t onwor kshop and MPRL refer ence co l lection mater ial . We par

t icu lar ly w ish to thank the fo l low ing for thei r r ev iews of

th is checkl is t :Maya B . Bes t , R i jksmuseum van Natuur l ijke H i s tor ie (Fav i

idae, Mer u l in idae, Muss idae, Ocul in idae, Pect in i idae,

Caryophyll i idae)

Gérar d Faur e, Un iv . de la Réun ion (Thamnas ter ndae,

As t rocoen i idae)Pau l L . Jok iel , H awai i Ins t i tute of Mar ine B iology (Por i t i

dae; nons ys temat ic scleract in ian l i terature for

Enewetak)Rober t A . K in z ie III, Un iver s i ty of H awai i (Tu r binar ia)Aus t in E . Lam ber ts , G rand Rapids , M ich igan (As t r eopor a)James E . Mar agos , U . S. Army Corps of Eng ineer s , H awai i(Mon t ipor a)

M ichel P ichon , James Cook Un iver s i ty of Nor th Queensland (Mon t ipor a, Fungi ina, Tu r binar ia)

A (STONY CORALS) 75

u ra and Anomur a) of En iwetok Ato l l , Mar shal l Is lands , wi thSpecial Refer ence to the Obl igate Commensa ls of Branch ingCorals , M icr ones ica, 1 : 137- 1 44 .

Go reau , T . F . , 1 959 , The Phys io logy of Skeletal Formation in

Co r als . l . A Method for Measur ing the Rate of Calcium Depos i t ion by Corals U nder D iffer ent Condi tions,B io l . Bu l l . (WoodsH o le) , 1 1 6 : 59-75 .

H agger ty , J . A . , J . N . Weber (deceased) , J.

'

F . Cuffey , and P .Deines , 1 980 , Env i ronment-Related Morpho log ic and Geo

chem ica l Var iabi l i ty in the Modern Reef Corals Favia pa l lidaand Favia s tel l igera on Enewetak Ato l l , Pacific Geo logy, 1 4:95- 1 1 2 .

H ighsm ith , R . C . , 1 979 , Cor al Gr owth Rates and Env i r onmentalContr o l of Dens i ty Banding , J . Exp . Mar . B io l . Ecol , 37:1 07- 1 25 .

1 980a, Pass ive Co lon i za tion and Asexual Co lony Multipl icat ion in the Mass ive Coral Por i tes lutea M i lne Edwards and

H aime, J . Exp . Mar . B iol . Eco l . , 47: 55l 98ob, Bur r owing by the B ivalve Mo llusc l ithophaga cur ta inthe L iv ing Reef Co r al Mon t ipo ra ber ryi and a H ypothes is of

Rec iproca l Lar val Recru itment , Mar . B io l . , 56 : 1 55- 1 62 .

l 980c, Geog r aph ic Patter n s of Coral B ioeros ion : A Pr oduct iv i ty H ypothes is , J . Exp . Mar . B io l . Ecol , 46: 177- 196 .

1 98 1a, Coral B ioeros ion Damage Relative to Skeletal Dens i ty, Amer . Nat .

, 1 17: 1 93- 198 .

1 98 1 b, L ime-Bor ing A lgae in H er rnatyp ic Cor al Skeletons , J .

Exp . Mar . B io l . Ecol , 55: 267-28 1 .

1 98 1 c, Cor al B ioeros ion at Enewetak : Agents and Dynam ics ,In t . Rev . Gesam ten . H ydr obiol . , 66 : 335

-375 .

H i ldemann , W . H . , D . S. L inth icum , and D . C. Vann , 1 975a,

T ransplantation and lmmuno incompat ibi l i ty Reactions AmongReef-Bu i ld ing Corals , Immunogenet ics , 2: 269-284 .

D . S. L inth icum , and D . C . Vann , 1 975b, lmmuno incompat i

bi l i fy Reactions in Corals (Coe lenterata), Immuno log ic Phy logeny, W . H . H i ldemann and A . A . Benedict (Eds ) , P lenum ,

New York , pp . 1 05- 1 1 4 .

H umes , A . G . , 1 981 , A New Species of Tegas tes (Copepoda:H arpact ico ida) Associated wi th a Scleract in ian Co ral at

Enewetak Ato l l , Proc . B iol . Soc . Was h . , 94: 254-263 .

and M . Doj i r i , 1 982 , Xar i fi idae (Copepoda) Par as i ti c in IndoPaci fi c Scleract in ian Cor als , Beau/or tia, 32 : 139-228 .

and J . H . Stock , 1 973 , A Rev is ion of the Fami ly L ichomolgidae Kossman , 1 877, Cyclopo id Copepods Main ly Associatedwi th Mar ine Inver tebr ates . Sm ith . Con t ri . Zoo l . , 127: 1 -368 .

Johannes , R . E . , J . Maragos , and S. L . Co les , 1 972a, Oi l Damages Corals Exposed to A i r , Mar . Pollu t . Bu l l . , 3 : 29-30 .

et al . , 1972h, The Metabo l ism of Some Coral Reef Communi t ies : A Team Study of Nutr ient and Energy Flux at En iwetok,B ioscience, 22: 541 -543 .

Knudsen , J . W . , 1967, Tr apez ia and Tetr o lic (Decapoda, B r a

chyu r a, Xan thidae) as Obl igate Ectopa ras i tes of Poc i llopor id

and Acropor id Corals , Pac . Sci . , 21 : 51 -57.

Knutson , D . W . , and R . W . Buddemeier , 1 973 , Di st r i butions of

Radionucl ides in Reef Co ral s : Oppo r tun i ty for Data Retr ievaland Study of Effects , Radioact ive Con tainmen t of the Mar ineEnv i ronmen t , IAEA , V ienna, pp . 735-746 .

R . W . Buddemeier , and S. V . Sm i th , 1 972 , Coral Chronometer s : Seasonal Growth Bands in Reef Corals , Science, 177:

270-272 .

Lam ber t s , A . E. , 1980 , Two New Species of As treopo r a (Gni

dar ia, Anthozoa , Scleract in ia) fr om the M id-Pacifi c, Pac . Sci . ,34: 261 -267 .

1 982, The Reef Coral As treopo ra (An thozoa, Scleracti n ia,

As trocoen i idae) : A Revi s ion of the Taxonomy and Descr iption

76 DEVANEY AND LANG

of a New Species , Pac . Sci . , 36: 83-1 05 .

Muscatine, L . , and C F . D’

Elia, 1 978 , The Uptake, Retention ,

and Release of Ammon ium by Reef Corals , L imno l .

Oceanogr . , 23 : 725-734 .

Odum , E . P . , and H . T . Odum , 1 956 , Zonation of Cor als on Jap

tan Reef , En iwetok Ato l l , Ato l l . Res . Bu ll . No . 52 , pp . 1 -3 .

Odum , H . T . , and E . P . Odum ,1 955 , T r oph ic Structure and Pr o

duct iv i ty of a W indward Coral Reef Commun i ty on En iwetokAto l l , Eco l . Monogr .

, 25 : 291 -320 .

Pi lson , M . E. Q . , 1 974, Ar senate Uptake and Reduction byPoc i llopo ra ver r ucosa, L imnol . Oceanogr .

, 19: 339-341 .

Pomer oy , L . R . , and E. J . Kuenz ler , 1 969 , Phosphorus Tur noverby Cor al Reef An imals , in Sympos ium on Radioeco logy, D . J .Nelson and F . C. Evans (Eds ), U n iver s i ty of Mich igan Pr ess ,Ann Ar bo r , M ichigan , pp . 474-482 .

Pomer oy , L . R . , M . E . Q . Pi lson , and W . J . W iebe, 1 974 , T r acerStudies of the Exchange of Phosphorus Between Reef Waterand O r gan i sm s on

,the W indward Reef of En iwetok Ato l l , in

Pr oceedings of the Second In ter nat ional Co r al Reef Sympos ium , B r isbane, Aus tralia, 1 973 , 1 : 87-96 .

Randal l , R ., 1 971 , MPRL Ann . Rep . F Y 1 970— 71 , pp . 6-7 .

R ichmond, R . , 1982,Energet ic Con s ider at ions in the D isper sal of

Poc i ll0por a dam icor n is (L innaeus ) P lanu lae,in Pr oceedings of

the Four th In ter nat iona l Cor al Reef Sympos ium , Man i la, Ph i lipp ines , 1 98 1 , 2: 1 53 - 1 56 .

R ichmond , R . H . , and P . L . Jokicl , 1984 , Lunar Per iodici ty inLar va Release in the Reef Cor al Poc i llopo ra dam icor n is at

Enewetak and H awai i , Bu l l . Mar . Sc i . , 34: 280-287.

Roth , A . A . , C . C . Clausen , P . Y . Yab iku , V . E . Clausen , and W .

W . Cox ,1982 , Some Effects of L ight on Cor al Gr owth , Pac .

Sc i . , 36 : 65-8 1 .

Sm i th , S. V . , and J . T . H ar r ison , 1977 , Calcium Car bonate Production of the Mare incogn i tum , the Upper W indward ReefSlope, at Enewetak Ato l l , Sc ience, 1 97: 556

-559 .

Stimson , J. , 1978 , Mode and T im ing of Repr oduction in Some

Common H ermatyp ic Cor als of H awai i and Enewetak , Mar .

B io l . , 48 : 173- 1 84 .

Ver on ,J . E N . , and M . Pichon , 1 976 , Sc ler act in ia of Eastern

Aus tr al ia, Par t I : Fam i l ies Thamnas ter i idae, As t r ocoen i idae,

Poci llopor idae,Aus t . Ins t . Mar . Sci . Monogr . , 1 : 1 -86 .

and M . Pichon , 1 980 , Scleract in ia of Easter n Aus tral ia,

Par t III: Fam i l ies Agar ici idae, Sideras t reidae, Fungndae , Ocu

l in idae, Mer u lin idae, Mu s s idae, Pect in i idae, Car yophyll i idae,

Dendr ophyll i idae, Aus t . Ins t . Mar . Sc i . Monogr . , 4: 1 -422 .

and M . Pichon , 1 982 , Scler act in ia of Easter n Aus tr al ia,

Par t IV : Fam i ly Por t idae, Aus t . Ins t . Mar . Sc i . Monog r . , 5

1 -1 59 .

M . Pichon , and M . W i jsman -Best , 1 977 , Scler act in ia of

Easter n Austr al ia,Par t II : Fam i l ies Fav i idae, T r achyphyll i idae,

Aus t . Ins t . Mar . Sc i . Monog r . , 3 : 1-233 .

Ver on , J . E . N . , and C . C . Wal lace, 1 984, Scler act in ia of Easter nAustral ia, Par t V : Fam i ly Acr opor idae, Aus t . Ins t . Mar . Sci .

Monogr . , 6: 1-485 .

Vosburgh , F . , 1 977 , The Response to D rag of the Reef CoralAcr opor a ret icu lata, in Pr oceedings of the Thi r d In ter nat ionalCor al Reef Sympos ium , M iam i , F lor ida, 1 977, 1 : 477

-482 .

1 978 , Abstr act : The Str uctu r al Mechan ics and Eco logy ofthe Reef Cor al Acropora ret icu lata, J . B iomech . , 1 1 : 21 1 .

1 982 , Acr opo ra ret icu lata: Structur e,Mechan ics and Eco logy

of a Reef Cor al , Pr oc . R . Soc . Lond . B .,214: 481 -499.

Wal lace, C. C. , 1 978 ,The Coral Genus Acr opo r a (Scleract in ia:

As t rocoen i ina: Acropor idae) in the Centr al and SouthernGreat Bar r ier Reef Pr ov ince, Mem . Queens l . M us , 18:

273 -3 1 9 .

Weber , J. , and J . A . H agger ty , 1 976 , MPRL Ann . Rept . F Y

1 975— 76, 2 pp .

Wel ls , J . W . , 1 954 , Recent Corals of the Mar shal l Is lands , U . S.

Geo l . Sur vey Pr of . Pap . 260 - I,pp . 385-486 .

Wethey , D . S . , and J . W . Por ter , 1 976a, H abitat-Related Patter nsof Productiv i ty of the Fo l iaceous Reef Coral , Pavona p r aetor taDana, Coe len terate Eco logy and Behavior , G . O . Mack ie (Ed ),P lenum , New Yor k , pp . 59-66 .

and J . W . Por ter , 1976b, Sun and Shade D ifferences in Pr oduct iv i ty of Reef Corals , Nature, 262: 28 1 -282 .

W i jsman-Bes t , M . , 1 980 , Indo-Pacific Cor al Species Belong ing tothe Subfam i ly Mon tas t r einae Vaughan and Wel ls , 1 943

(Scleract in ia-Coelen terata) , Par t II : The Gener a Cyphas t rea,

Lep tas t r ea, Ech inopo r a and D ip loas t rea, Zoo l . Meded .

(Leiden) , 55: 235-263 .

Yabe , H . , T . Sug i yama, and M . Eguchi , 1936 , Recent ReefBu i lding Cor als fr om Japan and the South Sea Is lands Underthe Japanese Mandate, l . Sci . Rep . Tohoku Un iv . Second Ser .

Spec . Vol , 1 : 1 -66 .

Young , S. D . , 1 97 1 , Or gan ic Mater ial from Scler act in ian Cor alSkeletons — 1 . Var iation in Compos i tion Between SeveralSpecies , Comp . B iochem . Phys io l . B . Comp . B iochem .

, 40:

1 13 - 1 20 .

Chapter 9

TABLE 1

Check l i s t of Enewetak B r ach iopods

Class INART ICULATAOrder LINGULIDASuper fam i ly L INGULACEAFam i ly L INGUL IDAEL ingu la sp . , lar vae: Gi lmart in , 1 958 .

Class ART ICULATAOrder TEREBRATILIDASuper fam i ly TEREBRATELLACEAFam i ly MEGATH YRIDIDAEArgyrotheca argu ta Gr ant : Gr ant , 1983 .

(=A rgyrotheca sp . Cooper ,Fam i ly DALLIN IDAE

‘F r enu l ina sangu ino len ta Gmel in , 1 792 .

Order TH ECIDEIDASuper fam i ly TH ECIDEACEAFam i ly TH ECIDELL IN IDAEThecidell ina congr egata Cooper , 1 954 .

(= Thecidell ina max i l la H edley : Gr ant ,‘New Enewetak r ecord .

p lemen t mater ial for th is r epo r t . Each s tat ion where br achiopods wer e obtained has been ass igned a number in theUSNM r eg is ter of local i t ies , Depar tmen t of Paleobiology .Stat ion data are summar i zed in Table 2 . The Enewetakbr ach iopods cannot be seen under water because they aresmal l and l ive in dar k hab i tats ; ther efore,

co l lect ing had tobe somewhat r andom . P ieces of coral wer e broken ou t of

l ikel y look ing hab i tats and br ought to the sur face for examinat ion , much in the manner descr i bed by Jackson et al .

( 1 971 ) al though not on so vas t a s ca le. Mos t indiv idualswer e left attached to thei r coral s labs and were dr ied inthe sun ; those that became detached wer e preser ved inalcoho l for immed iate s tudy and were dr ied later .

ECOLOGYB rach iopods were found on Enewetak at depths r ang

ing fr om 5 m to 65 m, the l imi t of scuba equ ipmen t . At

B ik in i , wher e they wer e co l lected by dr edg ing , they are

repor ted as deep as 480 m (Cooper , The shallowes t occur r ence was a smal l F renu l ina sangu inolen ta on a

78

GR I N EM

GRANT

IO KM

|2"i v-VAN

ENEWETAK

Sou r/7Channe/

F ig . 1 Map of Enewetak s how i ng wher e br ach iopods wer e co l lected. Num ber s ar e las t digi ts of USNM local i ty number s32007-22.

coral fr ond at 5 m . The abundance and diver s i ty of brachiopods increase w i th depth and w i th incr eased dar knessin smal l caves and r ecesses in the r eef . On the seawar ds ide of B iken Is land , one p iece of cor al-cum -

sponge con

tained a dens i ty of Thec idellina congr egata that by ext r apolat ion amoun ts to 7000/m 2, and another has 40 ind iv iduals on a sur face 7 X 1 7 cm , amoun t ing to about3400/m

2(F igs . 6 through These dens i t ies compar e to

those found by Jackson et al . ( 1 971 ) at Jamaica and

Cur acao where dens i t ies of 5000 to 1 0 ,000/m2 were

encounter ed . These figur es may appear to exaggerate thedens i ty of br ach iopods on these reefs , because the com

mon impress ion i s that brach iopods ar e rare . Mos t d iver sdo not encounter br ach iopods , however , because they dono t sear ch for them del i beratel y , and br ach iopods ar e

essent ial l y inv is ible under water . In add i t ion , the di st r ibu

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80

forms that are tightly attached , T . congr egate and

A rgyrotheca argu ta, ar e clear ly adapted to avo id encr us tat ion by the ubiqu i tous sponges and algae; they gr ow awayfr om the subs tr ate and thus keep the commi ssur e elevatedfr om the encr us ter s . F renu l ina sangu ino len ta attaches by aped icle that ho lds the shel l wel l above the subs tr atum(actual l y below i t , on these under s ides ) and affor ds some

rotat ional mobi l i ty that can help to war d off encr us tat ion .

F r enu l ina sangu inolen ta is not as negat ive to l ight as isT . congregate. Several samples w i th F . sangu ino lenta alsocontain abundant H a limeda algae, but other pieces in the

co l lect ion have F . sangu inolen ta and T . congr egr eta in

assoc iat ion . The brach iopod found in the shal lowes t habi tatwas F . sangu ino len ta in 5 m off K idrenen (loc . 320 1 3a)where a s ing le tiny indiv idual was taken fr om a shadyplace in a n i che under an over hang . Apparently the somewhat warmer water at th is shal low depth was mor e a fac

tor than shelter fr om l ight in l imi ting the pr esence of

br ach iopods .

Two r epr esentatives of the Class lnar ticulata also probably occur at Enewetak . The lar va of L ingu la sp . is

r epor ted in p lankton hauls made in tr ansects acr oss the

souther n par t of the lagoon (G i lmar t in , Species of

L ingu la normal ly bur row in unconsol idated sed iment in the

in ter t idal'

and shal low subtidal zones . Thi s hab i tat was not

explored thoroughly , but the pr esence of the lar va ind icates that such a sear ch would produce specimens of a

L ingu la. Another inar t iculate genus that can be expectedto be found at Enewetak i s Cr en ia. An undeterm inedspecies in this genus was r epo r ted fr om B ik in i where thefauna is essent ial l y the same as at Enewetak (Cooper ,

Its smal l s i ze and tightly cemented habi t make i t

d i fficult to find (i t looks l ike a smal l , ver y flat l impet); thelarges t spec imen r epor ted by Cooper is 3 mm w ide, a pale

salmon color that blends in to the sur r ound ings , and a

shape that tends to con form to the contour s of the

subs trate.

ARTICULATAA rgyr otheca arguta i s char acter i zed by i t s small s i ze

and wh i te, clear , or tr ans lucen t shel l (F igs . 2 and 3 ) w i thweak to absent cos tae. I t had been r ecogn i zed fr omEnewetak as not belong ing to any descr ibed species(Cooper , 1 954) but was not descr i bed for fear that i t wasr epr esented on ly by juven i les . The pr esent Enewetak samp le is adequate, however , and contains small shel ls alongw i th the major i ty that seem to cease gr owth between 2and 3 mm in w idth (Table The nar row outl ine,

lack ofcos tae (other species are typ i cal ly s trongly cos tate) , and

lack of pigmen tat ion make i t un ique among species of

A rgyr o theca. I t was descr i bed recen tly (Gr ant , is

known on l y fr om Enewetak and B ik in i so far , and may

prove to be endem ic to the Mar shal l Is lands .

The two Paci fic spec ies descr ibed by B lochmann(1 910 , 1914) fr om Aus tralia and Tasman ia, A . aus tr a lisand A . may i, are m uch longer than A . argu ta and have thes tr ong cos tae that character i ze mos t species of the genus .

GRANT

F ig 2 A r gyr otheca ar guta fr om under s ide of coral in lagoonoff l ku r i n Is land at depth 15 m , local i ty 32014; j uven i le i n

smal l depr es s ion (4X ) .

F i g . 3 A r gyr otheca ar gu ta on same s lab fr om 32014 as in

F i g . 2; adu l t and j uven i le s hell s in close pr ox im i ty (4X ).

The co lor les s shel l of A . argu ta is another d is t ingu ishingfeature. A rgyr otheca argu ta d i ffer s from the other pediculate species of Enewetak , F r enu l ina sangu inolen ta,

in i t s smal lnes s , i ts flatter profi le, i ts wi de hinge, i ts t r i

angular r ather than round fo r amen , and, es pecially , i ts pelluc id shell mater ial .

A rgyrotheca argu ta attaches to the subst rate by meansof a shor t ped icle. It attaches somewhat loosel y , i .e. , the

pedicle is longer than i s normal in other member s of the

genus . A rgyr o theca arguta favor s cr ypt ic habi tats undercor al fr onds and in r ecesses , as would be expected fr om i t scolor less and tr ans lucent shel l . At Enewetak i t was found

H IOPODS 8 1

F i g . 5 F r enu l ina sangu inolen ta fr om n i ches at 24 m dep th in

the lagoon , shown -

attached to su bs t rate (4X ) . Shel l s ar e pi nk

w i th wh i te s tr i pes or so l id p ink .

chiopod the ver nacular name lamp shell (the ancient o i llamp) . The co lor is dis t inct ive, normal ly wh i te wi th p inks tr ipes or p ink mottl ing in i r regu lar r ows but occas ional ly aso l id p ink or r ed co lor wi thout s tr ipes . The dor sal valvebear s a shal low med ian depress ion result ing in a sulca teanter ior commis sur e.

The dor sal loop cons is ts of a lar ge hood of th in shell

mater ial attached to a shor t s l im median septum by meansof two br oad lateral bands . The ventr al valve has deltidialp lates outl in ing the anter ior par t of the foramen , which arecon junct in some spec imens but d is junct in mos t . Thespecies thus is var iable, but unm i s takable, in the Enewetakbr ach iopod fauna .

F renu l ina sangu ino len ta is the mos t widespr ead and

smal les t l iv ing spec ies of the genus (H atai , 1 940 ; Cooper ,Th is species l ike other br ach iopods at Enewetak

pr efer s cr ypt ic habi tats . It is not as abundant local l y as T .

cong regate but was found at more locali t ies than any

other br ach iopod species , being pr esen t at 14 of the 1 8

local i t ies sampled ( Table 2 Mos t specimens remainattached to thei r subs trates by the pedicle, so on ly thew idth measur emen t could be made w i thout danger of damage to the shel ls . Table 4 compar es the lengt h , w idth , andth ickness of 1 4 Enewetak specimen s . H owever , the smalles t specimen exam ined had a width of mm . The beakcur ves towar d the subs trate, so the do r sal valve is heldfac ing the coral fr ond or other “

over hang” that the shellscl ing to . Relat ive to the sea floor , the dor sal is thus upper

82

TABLE 4

Shel l Measur emen ts ofF r enu l ina sangu inolen ta in mm

Width Thi ckness Local i ty

and the ven tral lower as the shells hang fr om the cei lingsof thei r cr yptic reces ses . The red color looks gr een ishunder water , and the s tr ipes render the shells near ly inv is ible to the human eye in the dim and shimmer ing l ight .

Thecidell ina congregate was descr i bed or iginal l y fr omB ik in i Atol l where i t occur r ed as deep as 480 m (Cooper ,

It is the mos t abundan t br ach iopod at Enewetak ,although not the mos t w idespread . Thecidell ina congr egate

can be iden t i fied by i ts cemented habi t , w i th low cup

shaped ven tral valve and lid-l ike dor sal valve (F igs . 6

thr ough Its color ranges fr om whi te (mos tl y dead shells )to blu ish gray in the major i ty of liv ing shel ls . It is not

immediatel y di s tingu ishable fr om T . max i lla, known fromB ikin i (Cooper , al though close inspection r eveals

several di ffer ences . Thecidell ina congregate is smal ler(Table has the blue-

gr ay color , and the dor sal med ianseptum is wi der at the anter ior than at the pos ter ior ,wher e that of T . max i lla maintains near ly a un i form wi dthfr om back to fr ont .

Zumwalt (1 978) conducted a detai led anatomi ca l andfu nct ional s tudy of th is species , based upon mater ial heco l lected at Enewetak . H e found that bo th the diductorand the adductor muscles inser t in the hem ispondylium ,

wh ich i s a muscle platform that changes lengt h and shapew i th growth and gener al con figur at ion of the valve, to p ro

duce the mos t advantageous ang les for muscle act ion .

H ence the hem i spondylium var ies from one indiv idual tothe next .

As impl ied by the name , T . congr egate can occur inabundan t patches where extr apo lated dens i t ies r un to

several thousand per squar e meter . Judging fr om the sam

ples taken at Enewetak and r epo r ts of o ther species of

Thecidellina fr om elsewhere Jackson et al . ,

Thecidell ina sur v ives bes t in cr ypt ic habi tats . It feeds byopen ing the valves and expos ing the lophophor e to the

seawater , wh ich i s then pumped across the fi laments of

the lophophore. The lophophor e is cons ider ed a mod i fied

F i g . 6 Thecidel l i na cong r egate clus ter ed on under s i de of

cor al fr ond fr om sea s ide of B i ken Is land at USNM local i ty32022 (2X ). These ar e the specimen s whose measu r em en ts inthe speci es des cri pt ion indi ca te a width r ange fr om to

mm .

F ig . 7 Thecidel l ina congr egate fr om shal low cave at depthabou t 33 m on sea s ide of B i ken Is land. USNM local i ty 32022;com binati on of l i v i ng s hel ls and dead empty ven tr al valves

wi th nar r ow var i ati on i n s ize.

t r ocholophe, w i th a deep med ian inden tat ion along the

s ides of the dor sal med ian septum . The normal pos i t ion forfeed ing seems to be w i th the dor sal valve open to a pos it ion near ly per pend icular to the margin of the ventr al valve

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84 GRANT

H atai , K M . , 1 940 , Cenozo ic Br ach iopoda from Japan , Sci .

Rep . Tohuku Un iv , Second Ser . pp . 1 -413 .

Jackson , J . B . C . , T . F . Coreau, and W . D . H ar tman , 1971 ,

Recent Brach iopod— Corall ine Sponge Commun i ties and Thei rPaleonto log ical Signi fi cance, Science, 1 73: 623

-625 .

Rudw ick, M . J . S. , 196 1 ,‘Qu ick ’ and

‘Catch ’ Adductor Muscles

in B rachiopods , Nature 1 91 : 1 021 .

Su r lyk , F . , 1 972, Morpho logi cal Adaptations and Populati onStructures of the Dan ish Chalk Brach iopods , Kon ig . Donske

Vid . Se lek . , B io l . Skr ift . , 19: 57 .

Zumwalt , G . S., 1 978 , The Funct ional Morpho logy of the Tr op ica l

B rach iopod Th ecidell ina congr egate Cooper , 1 954, unpub

lished M .S. thes is , Un iver s i ty of Cal ifor n ia at Davi s .

Chapter 1 O

Reef-D welling B ryozoans of Enewetak A to ll

ROGER J . CU F FE Y‘and ROBERT S. COXT

‘Deper tmen t of Geos ciences (Deike B ldg ) , Penn sy lvan iaState U n iver s ity , Un iver s ity Par k, Penns y lvan ia 1 6802 ;TMuseum of Pa leon to logy, Un iver s ity of M ich igan ,

Ann A r bor , M ich igan 48 1 09

IN TRODUCT ION

Pu rpose

The goals of th i s r epor t ar e to l is t the br yozoans foundon the moder n reefs of Enewetak Ato l l , i l lus tr ate the mos tconspicuous species , ind i cate thei r biogeogr aph ic impl icat ions , and summar i ze thei r eco log y (par t icular ly ecozonal

di str ibut ions and cons tr uct ional r o les ) .

Si gn i f icance

The con tr ibut ion made by the phy lum B r yozoa to the

Enewetak r eefs , although known in general terms (Cuffey ,1970 , 1 973 , has not un t i l now been detai led at thespec ies level for the benefi t of those concer ned w i th var ious aspects of the Enewetak ecosys tem . Such in format ionw i l l also cons t i tute a preci se data base for compar at ivestud ies of foss i l r eefs contain ing these an imals . Mor eover ,the Enewetak species are of special br oader interes tbecause they represent the fi r s t moder n atol l-reef br yozoanfauna to be analyzed thoroughly . Thei r d is tr ibut ions thusw i ll shed l ight on br yozoan invo lvemen t in r eefs in gener al .

Col lect ion s

Over 40 loca l i t ies on Enewetak Ato l l were sampled byscuba d iv ing and reef walk ing in late 1 969 . Si tes wer eselected to pr ov ide balanced but comprehens ive coverageof the major env i ronmen ts , par t icular ly ecozones ,

developed on the atol l (Cuffey , 1 973 ,pp . 26 , 36

-38 ; Cu f

fey, 1 978 , pp . 68 , 72; H agger ty , Weber , Cuffey , and

Deines , 1980 , p . 97 ; Bas i le, Cuffey , and Kos ich , 1 984 ,

F ig . H owever , no samples wer e dr edged fr om below40 m ( 135 ft) .

L i ter atu r e

Mos t useful in iden ti fy ing the Enewetak br yozoans werethe monog raphs by Canu and Bass ler (1927 ,

H armer (1 9 1 5, 1 926 , 1 934 , Osbur n ( 1 950 , 1952 ,

Lev insen and Soule and SouleAdd i t ional paper s too numerous to l is t her e include occas ional relevant spec ies descr ipt ions as well .

F AUN AL COMPOSIT IONOver al l , 84 br yozoan species , 73 chei los tomes and 1 1

cyclos tomes in 6 1 genera have been ident i fied on

Enewetak (Table Of these species , s ix ar e new , beingpubl ished elsewhere (Cox and Cu ffey , submi tted) . In or derto r efer to these wi thout cr eating taxonom ic pr oblems ,they are denoted s imply as “

n . sp .

” fol lowed by the p roposed tr iv ial name in quotat ion mar ks and par entheses .

Among the remain ing species , sever al gener icr eass ignments wer e r equ i red (incorpo r ated in Table 1 ) dueto changes in concepts s ince publ icat ion of the ear l iermonog r aphs c i ted above.

Cer tain species ar e par t icular ly conspicuous among theEnewetak br yozoans and so compr ise a d is t inct ive as semblage (F igs . 1 and 2) char acter is t ic of th is ato l l and poss ibly o ther s . Such species appear consp icuous for di fferentr eason s . Some are r epr esented by several lar ge colon ies ,other s by many smal l co lon ies , s t i l l other s by a large totalsubs tr ate ar ea encr us ted , and other s yet by a h igh pr opo rt ion of the br yozoan fauna at indiv idual loca l i ties . Thesespecies ar e also d i fferen tly d is tr ibuted w i th in the atol l , as

d iscus sed below .

The Enewetak br yozoan species are character i zed byr elat ively few co lony growth forms (Cuffey , 1973 , pp .

29-3 1 and r efer ences ther ein) . Mos t numer ous by far (bothco lon ies and spec ies) are encr us t ing chei los tomes , th insheets or cr us ts r ang ing fr om many square m i l l imeter s inarea down to a few zooecia. Mos t of the Enewetak chei los tome fam i l ies

'

(Table 1 ) s tud ied by Cox ( 1 983 ; also Coxand Cuffey , subm i tted) grow as encr us t ing sheets . A few

sheets cons is t of more than one layer of zooecia, but noneencoun ter ed were th ickened into mult i lam inar nodularmasses (not even Cellepor er ie elbiros tr is , wh ich does formsuch masses in Baham ian br yozoan r eefs ; Cuffey , Gebelein ,

Fonda , B l iefn ick , Kos ich , and Sor oka, Mor e regular

CUFF EY AND COX

TABLE 1

Enewetak A tol l B r yozoan s

Systemati cs B iogeogr aphy‘ Ecology-

l“

Phy lum BRYOZOAClass GYMNOLAEMATAOr der CH EILOSTOMIDASuborder ANASCINAInfr aorder INOVICELLATA

Fam i ly AETEIDAEAetee tr uncate

Infr aorder MALACOSTEGA

Fam i ly CALLOPORIDAEA lder ina brevisp inaFam i ly H INCKSIN IDAEAn tropor e claus trecres se

A . lagu ncu la

:l :A . ovate

Cau lor amphus cf . br uneet Crenos ina cor onata

In fraorder COELOSTEGA

Fam i ly ONYCHOCELL IDAEi Sm i ttipora amer icanaFam i ly MICROPORIDAECeles chare levin sen i

Fam i ly TH ALAMOPORELL IDAEThelemopo relle dis tor ta

'

:l: T . expenseInfr aorder PSEUDOSTEGA

Fam i ly ASPIDOSTOMATIDAEMonopor ella fimbr iata

Infr aorder CELLULARINA

Fam i ly FARCIMINARIIDAED idymozoum marg inatumNel lie cf . tenu isFam i ly BUGUL IDAECau l ibugu la ci l iateFam i ly BEAN IIDAEBean ie cupu lar iens is

Fam i ly SCRUPOCELLARIIDAECeberea ell is i

Scr upocellar ie longi sp inose

Subo rder CRIBRIMORPH INAFam i ly CRIBRIL INIDAECr ibr i ler ie calamorphe

C . r adiateC . s imu lato rF igu lar ie n . sp .

Suborder ASCOPHORINAFam i ly ADEONIDAERep tadeonelle flegell ifere Ol ,Lg ;d,s

i R . joloens is Ol ,Lg ;d,s

‘Geogr aphy : P, Paci fic Ocean ; I , Indian Ocean ; C, Ca r ibbean and wes ter n At lantic; EEnewetak endem ic; F , prev ious ly known on ly as foss i ls .

TEcozone: Ol , Ow , oceanward reefs on leeward -or windward s ides of ato ll , respectively ; Lg ,lagoonal reefs . Depth : d , deep r eefs ; s , shallow reefs ; f, r eef flats .

i The several most conspicuous species on Enewetak (Figs . 1 and

(Th is table continued on next page. )

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88 CUFFEY AND COX

TABLE 1 (con t’d)Sys temati cs B iogeog r aphy

Fam i ly CLEIDOCH ASMAT IDAEi Cleidochasma po r cellanum

H ippopo rella sp in igera

Fam i ly MICROPORELL IDAEFenes tr u l ina melus i i

Fam i ly RETEPORIDAE (encrusting species )D r epanophore incisorD . tuber culatai Rhynchozoon cur tumR . so lidumFam i ly RETEPOR IDAE (fenestr ated species)Tr iphyllozoon cusp idatumFam i ly SAVIGNYELL IDAESevignyelle lefon t i iFam i ly TETRAPLARIIDAETetrap lar ia veleroee

Fam i ly TUBUCELLARIIDAETubuceller ie fus iform is

Class STENOLAEMATAOr der CYCLOSTOMIDASubo rder TUBULIPORINAFam i ly DIASTOPORIDAEStomatopo re cf . gr endipo raFam i ly TUBUL IPOR IDAEIdm idr onee flexuosa

P latonea ph i l ipp sae

Tubu lipo ra pu lcher r ima

Fam i ly ONCOUSOECIIDAEOncousoecie cf . ebr up teP roboscina cf . dichotomaFam i ly DIAPEROECIIDAED iaperoecia aff . r osea

Subo rder ARTICULINAFam i ly CRISIIDAECr is ia elongateC . cf. kerguelens is

Subo rder RECTANGUL INAFam i ly L ICHENOPORIDAED ispo rella cf . buskiL ichenopo ra novaz elendiae

Ecology'l‘

Ow ,Lg ;d,s

OI,Lg ;d

Ol ,Lg ;d,s

Ol ;Lg ;d,s

Ol ,Lg ;d,s

‘Geogr aphy : P , Pacific Ocean ; I , Indian Ocean ; C, Car ibbean and wester n At lantic; E,

Enewetak endem ic ; F , pr ev ious ly known on ly as foss i ls .

TEcozone : Ol , Ow , oceanward r eefs on leeward or windward s ides of ato l l , r espectively ; Lg ,

lagoonal reefs . Depth : d , deep r eefs ; s , shal low reefs ; f, r eef flats .

several mos t conspicuous species on Enewetak (Figs . 1 and

in outl ine, but always smal l and m uch r ar er , ar e th inencr us t ing d isks , Iichenopor id cyclos tomes . A lso along -the

subs tr ate ar e encr us t ing thr eads (aeteid chei los tomes , d iastopor id cyclos tomes ); encr us t ing br anches (some tubul ipor ine cyclos tomes ); and recumbent , though s l ightlyelevated , br anch ing tubes (other tubu l ipo r ine cyclos tomes ;Cuffey , Erect r igid latt ices are cer tain retepo r id

chei los tomes (Cuffey and McK inney, Er ect flexible

or jo in ted tufts include bo th extremel y del icate cr i s i id

cyclos tomes and somewhat mor e robus t chei los tomes (cellu lar ine, sav ignyell id, chei lopor in id, and tubucellar i id

chei los tomes ) . None of the Enewetak r eefal br yozoanswere obser ved to gr ow as er ect cyl indr i cal or flat tenedbranches .

All the Enewetak br yozoans men t ioned above ar e geo

log i ca l ly moder n . Deep dr i l l ing in to the late Cenozoi<

REEF -DWELLING BRYOZOANS

F i g . 1 The mos t abundan t Enewetak br yozoans (width of each fi eld in mm : specimen s num ber ed wi th in“

MAPEN I”

sui te in Paleobryozoo logi cal Resea r ch Col lect ion at Penns y lvan ia State U n i ver s i ty) . (a) Cr enos inacor onata, field mm wide. 27-A-1 ; (b) Cosc in iOps is n . sp . fi eld mm w ide, 16-G-12;

(c) Chor r'zopo r a ven t r i cose , field mm wide, 27-A-1a ; (d) The lemopor el le expense, field mm wide.

1 7-B-2; (e) R ep tadeonel le joloens is , field mm wide, 16-G -8 .

CUF FEY AND COX

F i g . 2 The o ther comm on Enewetak br yozoan s (s izes and num ber s as in F i g . 1 ) (a) A r echnopus le spa thu lete, fieldmm wide, 1 4-A-37; (b) H ippopodi ne feegeens is , field mm wide, 31 -C-4 ; (c) R hynchozoon cu r tum , fi eld mm wide,3-B~12; (d) Sm i t t ipore amer i cana, field mm wide, 21 -B-4 ; (e) Ca lyp totheca impar , fi eld mm wide, 21 -A-3:

( f) Trypos tega venus te, field mm wide, 15-A-2; (g ) Cleidochasma por cellanum , field mm wide, 21 -A-8 :

(h ) An t r opor e ovate,

“ field mm wide, 25-A-22; ( l ) Paresm i tt ine elenbanner i , field mm wide, 20-B-12.

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92 CUF FEY AND COX

aper tu r es , F ig . 2g)—

are common on deep and shal lowoceanwar d (leewar d and w indwar d) and lagoonal r eefs .

Comparably d is tr ibuted , except not on w indwar d oceanwar d r eefs , are An tropo re ovate (many t iny av icular ia,

F ig . 2h) and Per esm i tt ina elenbanner i (long cur ved av icu

lar ia, F ig . 2i ) .

B r yozoan roles in r eef cons tr uct ion are as impor tan t -tothe paleoecologis t as ecozonal d is tr ibut ions (Cuffey ,The Enewetak br yozoans ar e all cr yp tic or coelobi t ic , funct ion ing mos t ly as h idden encr us ter s under coral fr onds ,coral- rock ledges , detached r ubble, and occas ional shel ls . A

few ar e local ly cav i ty dweller s , but none wer e obser ved tobe cav i ty f i l ler s , fr ame bu i lder s , or sed imen t producer s .

The Enewetak br yozoans thus are more r es tr icted in con

s tr uct ional r o les than cer tain cav i ty- fi ll ing and r ock-bu i ld ingBaham ian relat ives (Stor r , 1 964 ; Cuffey , Gebelein , Fonda,

B liefn ick , Kos ich , and Sor oka, 1 977; Cuffey and Fonda,

ACKNOWLEDGMEN TSAs s is t ing in col lect ing were M . Zei gler , R . Por t ,

P . K ier , R . Gr an t , J . Pojeta, E . Kauf‘fman , T . Wal ler ,T . Phelan , A . Ch i lds , R . Mehlen , and C. Dawson ,

w i thlog is t ical suppo r t fr om En iwetok Mar ine B iolog i cal Labor ator y , Pennsylvan ia State Un iver s i ty , Nat ional Sc ience Foundation , and Atom ic Energy Commis s ion .

REFERENCES

Bas i le, L . L . , R . J . Cuffey , and D . F . Kos ich, 1 984 , Sclerosponges , Phar et r on ids , and Sphinctozoans (Rel ict Cr ypticH ard-Bodied Por ifer a) in the Moder n Reefs of Enewetak Ato l l ,J . Paleon t .

, 58 : 636-650 .

B rown , D . A . , 1 964 , Foss i l Br yozoa from ‘

Dr i ll H o les on En iwetokAto l l , U . S. Geo l . Sur v . Prof . Pap .

, 260 -EE,pp . 1 1 1 3 - 1 1 1 6 .

Canu , F . , and R . S. Bas s ler , 1927, B ryozoai r es des l les H awai i ,Bu l l . Soc . Sci . Seine-et -Oise, 7 1 -66 .

1 929 , Br yozoa of the Phi l ippine Region , U . S. Nat . M us .

Bu ll . , 1 -685 .

Cox, R . S. , 1983 , Encrus t ing Chei los tome B r yozoans of Enewetak(En iwetok) A to l l— Iden t if ication Manual and D i s tr ibuti onalData

, unpubl ished r epo r t for M id-Paci fic Mar ine Labor atoryand B i shop Museum (supplemented fr om MS . thes is , Pennsylvan ia State Un iver s i ty , Un iver s i ty Park).and R . J . Cuffey , Di s tr ibution , Eco logy , and New Species of

Encrusting Chei lostome Br yozoans on Enewetak Ato l l ,M icrones ica, (subm i tted) .

Cuffey , R . J. , 1 970 , En iwetok B ryozoans , En iwetok Mar ine B io l .Lab . Ann . Rept . , 1 969— 1 970, 3 pp .

1973 , B ryozoan Di s tr i bution in the Moder n Reefs of

En iwetok Ato ll and the Bermuda P latform , Pac . Geo l . , 6 :25-50 .

1 977 , B ryozoan Contr ibutions to Reefs and B ioher rns

through Geo log ic T ime, Amer . As soc . Pet r o l . Geo l . Stud .Geo l . , 4 : 1 8 1 - 1 94 .

1 978 , A Note on B r yozoans in the Moder n Reefs of

En iwetok Ato l l and the Aus tral ian Great Bar r ier Reef, Pec.

Geo l . , 1 3 : 65-76 .

1 985 , Reef-Dwel l ing Cyclostome Br yozoans on EnewetakAto l l— Compar ison s and Contras ts for Foss i l Reefs , Geo l .Soc . Amer . Abs t r . Prog .

, 1 7: 283 284 .

blages in Modern Coral Reefs off Andr os and Eleuthera, Bahamas , Proceedings of the Thi r d In ter nat ional Coral Reef Sympos ium , Miam i , F lor ida, May 1 977, 1 : 81

-86 .

and S. S. Fonda, 1 979 , Br yozoan Species Assemblages inthe Moder n B r yozoan -R ich Reefs at Jou lter s Cays(Bahamas ) — General Character and Paleoenv i ronmen tal Imp licat ions , Geo l . Soc . Amer . Abs t r . P r og r .

, 1 1 : 407-408 .

C . D . Gebe lein , S. S. Fonda, D . M . B liefn ick , D . F .Kos ich , and L . G . Sor oka, 1977 , Modern T idal-ChannelBr yozoan Reefs at Jou lter s Cays (Bahamas ) , Proceedings ofthe Th i r d In tem et ionel Cor a l Reef Sympos ium ,

Miam i , F lor ida,

May 1 977, 2: 339-345 .

and F . K . McKinney, 1 982, Retepor id Chei los tome Brydzoans from the Modern Reefs of Enewetak Ato ll and Thei rImpl ications for Paleozoic Fenes tr ate B ryozoan Paleoeco logy ,Pec . Geo l. , 1 6 : 7- 13 .

Fonda, S. S. , and R . J . Cuffey , 1 976 , B r yozoan Assemblages inthe Modern Bermuda Reefs — Comparative Data for Analys isof Cenozo ic B ioher rn s , Geo l . Soc . Amer . Abs tr . P rogr . , 8 :

1 75 .

H agger t y , J . A . , J . N . Weber , R . J . Cuffey , and P . Deines , 1 980 ,

Env i r onment-Related Morpho logi c and Geochem ical Var iabi l i tyin the Modern Reef Corals Favia pa l lida and Favia s tell igera on

Enewetak Ato l l , Pec . Geo l . , 1 4 : 95— 1 1 3 .

H armer , S. F . , 1 91 5 , The Po lyzoa of the Siboga

Expedi tion , Pt . 1 : Entoprocta, Ctenostomata, Cyclos tomata,

Siboga-Expedit ie, 28a : 1 - 1 80 .

1 926 , The Po lyzoa of the Siboga Expedi tion , Pt . 2 : Chei los tomata Anasca , Siboga

—Expedit ie, 28b : 1 8 1-50 1 .

1934 , The Polyzoa of the Siboga Expedi tion , Pt . 3 : Chei los tomata Ascophor a I , Fam i ly Retepor idae, Siboga-Expedit ie,

28C : 502-640 .

1957 , The Po lyzoa of the Siboga Expedition , Pt . 4 : Chei lostomata Ascopho ra II , Siboga-Expedit ie, 28d : 641 - 1 1 47.

Lev insen , G . M . R . , 1909, Morpho log ical and Sys tematic Studies on the Chei los tomatous B ryozoa, Nat . For fatteres For lag ,

Copenhagen , pp . 1 -43 1 .

Osbur n , R . C. , 1 950 , B r yozoa of the Pacifi c Coas t of Amer ica ,

Pt . 1 , Chei los tomata— Anasca, A l lan H ancock Pacif . Exped. ,

14 : 1 -269 .

1 952 , B r yozoa of the Pacific Coast of Amer ica , Pt . 2 :Chei los tomata— Ascophora, A l lan H ancock Pacif . Exped . , 1 4

27 1 -61 1 .

1 953 , Bryozoa of‘

the Paci fic Coast of Amer i ca , Pt . 3 :Cyclostomata, Ctenos tomata, Entoprocta, and Addenda, A l lanH ancock Pacif . Exped . , 14 : 613-84 1 .

Soule, D . F . , and J . D . Soule, 1 973 , Morpho logy and Speciationof H awai ian and Eastern Pacific Sm it t in idae (B ryozoa,

Ectopr octa) , Amer . Mus . Nat . H is t . Bu ll . 1 52 , pp . 365-440 .

Storr , J . F . , 1 964 , Eco logy and Oceanogr aphy of the Coral -ReefT ract , Abaco Is land , Bahamas , Geo l . Soc . Amer . Spec . Pep .

79, pp . 1 -78 .

Chapter 1 1

Enewetak Atoll

TABLE 1

Checkl i s t of Enewetak Sipuncu la and Ech iu r a

Phy lum SIPUNCULAFam i ly SIPUNCUL IDAE

‘Siphonosome (Damos iphon) cumanense (Kefer s tein ,

‘Siphonosome (Siphonosome) r otumenum (Shipley, 1 898) as‘Siphonosome (Siphonosome) en iwetoki Fisher , 1950 ;Fisher , 1 950 ; Stephen and Edmonds , 1972 .

‘Sipuncu lus indicus Peter s , 1 850 .

Fam i ly ASPIDOSIPHONIDAEAsp idos iphon mue l ler i Dies ing , 1 851 : H ighsm ith , 1 981 .

Cloeos iphon asperg i l lus (Quatrefages , H ighsm i th ,198 1 .

L ithecros iphon cr is tatus (Slu iter ,L i thecros iphon gu rjanovee Mur ina, 1967 : H ighsm i th ,1 98 1 .

L ithecr os iphon odhner i Fischer , 1 922 : H ighsm ith , 1981 .

L ithecros iphon un iscu tetus (Ikeda, H ighsm ith ,1 98 1 .

Par asp idos iphon gigas (Sluiter , H ighsm ith , 1981 .

‘Paresp idos iphon johns ton i Edmonds , 1980 .

Fam i ly PH ASCOLOSOMATIDAE‘Phescolosome (Phascolosoma) n igr escens Kefer s tein ,

1 865 .

Phylum ECH IU RAOrder ECH IU ROINEAFam i ly BONELLIIDAE

‘Achaetobonell ie macu lete Fisher , 1953 .

‘Fam i ly ECH IURIDAEAnelas sor hynchus ? sp .

‘New Enewetak r ecord .

Is lands w i thout fu r ther local i ty . Geog raph ic range of s ipuncu lans i s shown in Table 2 .

ECH IURA

Echiu ran s are pr ev ious ly unr ecor ded from Enewetak orthe other Mar shal l Is lands . H owever , the bi fur cate-tippedpr oboscis (somet imes up to a meter in lengt h) of a bonell idechiu r an has been obser ved in the shallower water s of

Enewetak Lagoon extending fr om beneath attached cor alreef outcrops . Two specimens which appear to be of the

94 DEVANEY

TABLE 2 final determ inat ion is pos s ible (S. J . Edmonds , per sonalcomm un i ca t ion ) .

Iden t i f ied Enewetak Si puncu lan sRecor ded an d Thei r Geog r aph i c Range‘

REF ERENCESSiphonosome (D .) cumanense Pan -tr opica lSiphonosome (S .) r o tumenum Mar shal l Is lands (Enewetak)Sipuncu lus inducu s Eas t Afr ican coast to Car o line

Is landsAsp idos iphon m ue l ler i At lantic bo r eal ; W . Indian

Ocean ; JapanCloeos iphon aspergi l lus Indo-West Paci ficL ithecr os iphon cri s tatus Indones ia, tr opica l Pacific

and Car ibbean , Japan , Austr al iaPhes co losoma (P .) n igr escen s Pan -tr opical

‘Based on Stephen and Edmonds , 1972; Edmonds , 1 980 .

same form wer e co l lected at Kawajalein Atol l in 1 982 byScott Johnson . S. J . Edmonds has iden t i fied these as

Achaetobonell ia macu lata F i sher , pr ev ious ly r ecor ded fr omOnotaa in the G i lber t Is lands . A lso a complete specimenmember of the fami ly Echiu r idae was collected

'

in the in tert idal ar ea of a shal low channel beneath r ocks . The latter istentat ively determ ined as a mem ber of the genusAnelas sorhynchus . Add i t ional mater ial is r equ i r ed befor e a

Cut ler , E . B . , and N . J . Cut ler , 1 98 1 , A Recons ider at ion of

Sipuncula named by I . Ikeda and H . Sato ,Publ . Seto . Mar .

B io . Lab . , 26: 51 -93 .

1 982 , Rev is ion of the genus Siphonosome,Pr oc . B io l . Soc .

Wash . , 95: 748-762 .

Edmonds , S. J. , 1 980 , A Rev i s ion of the Sys tematics of Aus

t r alian Rec . 8 . Aus t . M u s,1 8: 1 -74 .

Fischer , W . , 1 928 , Uber zwei neue Siphonosome Ar ten der

Wur tembu rgischen natur al ien Samm lung zu Stuttgar t , Zoo l .Anz . ,

76 : 1 38- 1 43 .

Fisher , W . K . , 1 950 , The New Subgenera and a New Species of

Siphonosome (Sipunculoidea) , Ann . Mag . N at . H is t , Ser . 1 2,

3 : 805-808 .

H ighsm ith , R . C . , 1 98 1 , Cor al B ioeros ion at Enewetak : Agentsand Dynam ics , In t . Revue Gesamen . H ydrobiol 66 : 335-375 .

Ikeda, l . , 1 924 , Fur ther Notes of the Gephyr ea of Japan withDescr ipt ions of Some New Species fr om the Mar shal l , Car ol ine, and Palau Is lands , Jap . J . Zoo l . , 1 : 23-44 .

Stephen ,A . C. , and S. J . Edmonds , 1972, The Phy le Sipuncu la

and Echiu ra, T rustees Br it . Mus . (Nat . H is t ), London .

Suchanek , T . H . , and P . L . Co l in , 1 986 , Rates and Effects of

B iotur bation by Inver tebr ates and Fishes at Enewetak and

B ik in i Ato l ls , Bu l l . Mar . Sc i . , 33 : 25-34 .

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96 DEVA

Resear ch Labor ator y (MPRL ) (Mar t in , As a r esult ,one new spec ies of trematode (fluke) was descr ibed fr omspec imens of a butter fly fish (Mar t in and H ammer ich ,

Four new spec ies of tur bel lar ian po l yclad flatwormswer e repor ted by H yman (1 959) fr om Enewetak as a

r esult of col lect ions made by D . I . Reish in the summer of1 956 from r eef flat rocks and algal thal l i . An undeterm inedr habdocoel was co l lected fr om a shell and coconut con taining fresh water found on the gr ound dur ing the summer of1963 (Magu i r e,

N EMERTEA (R ibbon Worm s )

Th ree species of ben th ic nemer t ine worms found at

B ik in i and Enewetak in 1946 included on lyone

— Baseod i scus delineatus , 30 cm long , 4 to 5 mmw ide— from Enewetak (Coe, Th i s spec imen was

co l lected in shal low water on the lagoon s ide of

Bogon , (Boken ) Is let . Subsequen tly , i t was noted that nemer teans made up 2596 of the meiobenthos fr om in ter s t i t ialsands exam ined at Enewetak . Two undeterm ined nemer

tean s , co l lected on the r eef flat beachr ock near the quar r yon Enewetak i s let , have been depos i ted in the MPRL r eference co l lect ion .

N EMATODA (Roundworm s )

The di s tr ibut ion and biological r elat ion s of a s tomachnematode, P r otosp i r ura mur icola, found in r ats and house

Chapter 1 3

Two ectocommensal po lychaetes have been recor dedas symbion ts fr om ech inoderm s at Enewetak . H ar tman(1 954) repor ted Ges t r olep idie cleviger e assoc iated w i th thesea cucumber s St ichopus hor rens at B ik in i , Enewetak , andRongelap and w i th H o lothur ia gyr ifer at Enewetak . Two

add i t ional holo thu ro ids (Act inopyge maur itiana and

H o lothur ia atr a) wer e recor ded as hos ts for th is po lyno idby Rei sh Devaney ( 1 967) noted H o lo lep idella

n ig rop unctata as a symbion t w i th the br i ttle-

s tarOphiocoma aneg lyp t ica at Enewetak . Straughan (1969)made the fi r s t iden t i ficat ions of sp i r orbid tube worm s fromEnewetak . Two spec ies wer e r ecogn i zed . One, er roneous lydescr ibed as a new spec ies , is the s in is tr al l y co i led speciesVinear ia koehler i [as Sp i r or bis (P i leo lar ie) po lyopercu letusStraughan ,

the other is a dextr al species ,Neodex iosp i ra bras i liens is [as Sp i r or bis (Ci r ceis) be llu lusBus h]. Young ( 1 969) r ecor ded a nud ibranch that preyedon the above two spec ies and on a serpul id .

Lev i ten ( 1976 ,1 978) r ecor ded po l ychaetes fr om the

fam i l ies Eun ic idae and Nereididae that ser ved as pr ey forseveral con id gas tr opods on the seawar d r eef bench at

Enewetak Is let and elsewher e in the Indo-Wes t Pac i fic .Kohn in his con t inu ing eco log ica l s tud ies on

Conus , r epor ted r epr esen tat ives of eight po lychaete fam il ies that are pr ey for sever al of these gas tropods . Some of

the iden t i fied po lychaetes are appar ent ly the fi r s t r ecor dsof these spec ies fr om Enewetak where they occur r edmain ly in the sandy lagoon among algae (H al imeda

An under s tand ing of the habi tat pr efer ences and d ivers i ty of r eef flat po lychaetes along the seawar d s ide of

Enewetak Is let was gr eatl y enhanced thr ough a s tudy conducted in 1 976 and 1 978 under labo rator y suppo r t byBa i ley-B r ock et ai . As a r esult of th is wor k , i t wasdeterm ined that 44 po lychaete taxa included 1 8 ident i fiedspec ies that wer e new Enewetak r ecords , w i th representat ives of two add i t ional fam i l ies (Paraon idae and Scal ibr egm idae) . Over 20 of the 3 1 syll ids had been pr ev ious lyunr ecor ded , but 12 have yet to be ass i gned spec i ficnames .

Po l ychaetes r epr esen t ing 26 fami l ies wer e abundan tin faunal componen ts as soc iated w i th selected Enewetakcor als and co r al r ubble (H ighsm i th , The po l ychaetes wer e the mos t common inver tebr ates found in thecoral heads , w i th over 80% of all ind iv iduals repr esen ted

98 DEVANEY AND BAILEY-BROOK

TABLE 1

Check l i s t of Polychaetes of Enewetak A tol l‘

ANNEL IDAClass POLYCH AETAFam i ly POLYNOIDAE

Ges t rolep idie clevigere Schmarda, 1 86 1 : H ar tman , 1954 .

Ges t r olep idie clever igere [s ic]Schmarda: Reish, 1968 .

H armothoe imbr iceta (L innaeus , H ar tman , 1 954 ; Reish, 1 968 .

H ololep idella n igr Op unctete (H or st , Devaney , 1967 .

Iphione mur icate Sav igny , 1 8 1 8 : Reish, 1 968 .

Perelep idonotus empu ll ifer us (Gr ube,

Per elep idonotus ampull ifera H artman ,1 954 ; Beish , 1968 .

Fam i ly PALMYRIDAEPalmyra eu r ifera Sav igny , 1 8 18 : H ar tman , 1954 ; Reish, 1968 .

Fam i ly CH RYSOPETAL IDAEBhawen ia goodei Webs ter , 1 884 : Reish, 1968 .

Bhawen ia cryp tocephdla Gravier , 1901 : H ar tman , 1 954 .

ChrySOpetelum ehler s i Grav ier , 1902 : H ar tman , 1 954 ; Reish, 1 968 .

Peleeonotus debi lis (G rube,

Peleeonotus debi lis [s ic]: Bai ley-Brock et al . , 1980 .

Fam i ly AMPH INOMIDAEEu rythoe comp lanete (Pallas , H ar tman , 1954 ; Reish , 1968 ; Bai ley-B rock et al . , 1980 .

Eu rythoe pacif ica Kinberg , 1 857 : Pomer oy and Kuenzler , 1967 .

Pseudeu rythoe ocu l ifera‘(Augener , H ar tman , 1 954 ; Reish, 1 968 ; Ba i ley-Brock et al . , 1980 .

Fam i ly EUPH ROSIN IDAEEuphr os ine myr tosa Savigny , 1 8 18 : H ar tman , 1954 ; Reish, 1 968 .

Fam i ly PHYLLODOCIDAEAneit ides madeir ens is (Eangerhans , H ar tman , 1954 ; Rei sh, 1968 .

Eu la lia vi r idis (L innaeus , Bai ley-Brock et al . , 1980 .

Eu lalia tenex Grube, 1878 : H ar tman , 1954 ; Reish, 1 968 .

Eum ide sangu inee (Oer sted, Reish, 1 968 .

Genety l lis gr aci lis (Kinberg , H ar tman , 1 954 ; Reish, 1 968 .

Phy l lodoce marquesen s is Monr o , 1 939 : H ar tman , 1954 ; Reish, 1968 .

Phy l lodoce p r uvo t i Fauvel , 1930 : H ar tman , 1954 ; Rei sh, 1968 .

Fam i ly H ESION IDAEH es ione genetta Grube, 1 866 : H ar tman , 1954 ; Rei sh , 1 968 .

Leocretes ch inens is Kinberg , 1 866 : H ar tman , 1 954 ; Reish, 1 968 .

Poder ke augu s tifr ons (Gr u be , Reish , 1968 .

Irma? augus t ifr ons H ar tman , 1 954 .

Fam i ly PILARGIDAESynelm is albin i (Langer hans , 1 88 1 ) in Pettibone, 1 966 .

Ancis trosyllis r igida Fauvel , 191 9 : H ar tman , 1 954 ; Rei sh, 1 968 .

Fam i ly SYLL IDAESubfam i ly SYLL INAELangerhans ia cor nuta (Rathke,

Ehler s ia cor nuta: Ba i ley-Br ock et al . , 1 980 .

Langer hans ia sp . A : Bai ley-Br ock et al . , 1 980 .

Langerhans ia sp . B : Bai ley-B r ock et al . , 1980 .

H ap losyll is eber rans (Fauvel , H artman , 1 954 ; Reish , 1 968 .

H ap losyll is Spong icola (Grube, H ar tman , 1 954 ; Reish, 1 968 ; Bai ley-B rock et al . , 1 980 .

Op is thosyll is aus t ralis Augener , 1 91 3 : Ba i ley-B r ock et al . , 1 980 .

Op is tosyll is [s ic]aus tral is Reish, 1 968 .

Op is thosyll is brunnea Langer hans , 1 879: Bai ley-Brock et al . , 1 980 .

Op is tosyll is [s ic] brunnea Reish, 1 968 .

Op is thosyll is corall icole (H ar tmann—Schr é der , Bai ley-Br ock et al . , 1980 .

Op i thosyll is long ic i r r eta Monro , 1939.

Op is tosyll is [s ic] long ici r r ate: Reish , 1 968 .

Syl lis gr aci lis Grube, 1 840 : H ar tman , 1 954 ; Reish , 1 968 .

‘The author s and dates for species have been checked us ing H ar tman (1 959) and Day Refer ences fo l lowing the co lon are

Enewetak records .

(Th is table continued on next page. )

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100 DEVANEY AND BAILEY-BROOK

TABLE 1 (con t’d)

Fam i ly NEREID IDAE (cont ’d)P letynereis dumen

‘

l i i (Audouin and M i lne Edwards , Kohn, 1 98 1 .

Pletynereis po lyscalma Chamber l in , 191 9 : Reish, 1 968 .

P letynereis pu lchel la Gr av ier , 1 901 : Reish, 1 968 ; Bai ley-B rock et al . , 1980 .

Pseudonereis enomela Gr av ier , 1 90 1 : H ar tman , 1 954; Reish, 1968 .

Pseudonereis var iegata (Grube, Bai ley-Brock ci al . , 1 980 .

Pseudonereis gellepagens is Kinberg , 1 866 : H ar tman , 1954 ; Reish, 1 968 .

Fam i ly NEPH TYIDAEM icronephtys sphaeroc i r r eta (Wesenberg-Lund , Reish, 1 968 .

Fam i ly SPH AERODORIDAESpheer odor idium capense (Day,Spheerodorop s is capen se Day : Bai ley-B r ock et al . , 1980 .

Fam i ly GLYCERIDAEG lycera tes se late Grube , 1 863 : H ar tman , 1 954 ; Reish , 1 968 ; Bai ley-B r ock et al . , 1 980 ; Kohn , 198 1 .

G lycera sp . : H ar tman , 1 954 .

Fam i ly EUN ICIDAEEun ice af ra Peter s , 1 854 : H ar tman , 1954 ; Reish, 1968 ; Lev iten , 1976 , 1978 ; Bai ley-Br ock et al . , 1980 ; H ighsm ith , 1 98 1 .

Eun ice an tennata (Sav igny , H ar tman , 1 954 ; Reish 1968 ; H ighsm i th , 1 98 1 .

Eun ice aus t ral is Quatr efages , 1 865 : Reish, 1 968 ; H ighsm i th , 198 1 .

Eun ice (N icidion) g raci l is Cr oss land, 1 904 : Reish, 1968 .

Eun ice vittata (Del le Chiaje, Kohn , 1 98 1 ; H ighsm i th , 1 98 1 .

Lys idice n inetta Audouin and Mi lne Edwards , 1 833 .

Lys idice co l ler is Grube, 1 868 : H ar tman , 1 954 ; Reish , 1 968 ; Lev iten , 1976 , 1 978 ; Bai leyBrock et ai . , 1980 ; H ighsmith , 1Nematonereis un icom is (G rube , H ar tman , 1 954 ; Reish, 1 968 ; Bai ley-Br ock et al . , 1 980 ; H ighsm ith , 1 981 .

Pelo la s ici l iens is (Grube , H ar tman , 1 954 ; Rei sh, 1 968 ; Lev iten , 1976 , 1 978 ; Bai ley-Brock et al . , 1980 ; Kohn , 1 98 1 .

Eun ice s ici l iens is Grube: H ighsm i th , 1 98 1 .

Peremarphysa or ien ta l is W i l ley , 1 905 : H ar tman , 1954 ; Reish , 1968 .

Fam i ly LUMBRINERIDAELumbr iner is latrei ll i Audou in and M i lne Edwards , 1 834 : Bai ley-B r ock et al . , 1980 .

Lumbr inen‘

s sphaerocephala (Schmarda, H ar tman , 1954 ; Reish, 1968 .

Fam i ly ARABELLIDAEA r abe lla ir ico lor (Montagu , H ar tman , 1 954 ; Reish , 1 968 .

A rabe l la mutans (Chamber l in , H ar tman , 1 954 ; Reish, 1968 ; Kohn , 1 98 1 .

D r i lonereis sp . cf . D . major Cross land, 1 924 : MPRL Co l lection .

Fam i ly LYSARETIDAEOenone fu lgide Sav igny , 1 8 1 8 : Bai ley-Br ock et al . , 1 980 ; H ighsm ith , 1 98 1 .

Ag lau r ides fu lgida (Sav igny) : H ar tman , 1 954 ; Reish, 1 968 .

Fam i ly DORVILLEIDAEDor vi llea p seudor ubrovi tteta Berkeley , 1 927 : H ighsm ith , 198 1Do r vi llea s im i lis Cross land , 1 924 : H ar tman , 1 954 ; Rei sh, 1968 ; Bai ley-B rock et al . , 1 980 .

Pap i ll iodor vi llee gardiner i (Cross land, Reish, 1968 .

Do r vi llea gardiner i (Cross land) : H ar tman , 1954 ; H ighsm ith , 198 1 .

Fam i ly ORBINIIDAENainer is sp . : H ar tman , 1954 ; Reish , 1 968 .

Scolop los sp . : Bai ley-Brock et al . , 1980 .

Fam i ly PARAONIDAELevinsen ia gr aci lis (Tauber , H ar t ley, 1 98 1 .

Par aon is gr aci lis : Bai ley-B rock et al . , 1 980 .

Fam i ly SPION IDAELeon ice ci r rata (Sar s , H ar tman , 1 954 .

Malacoceros (Malacocer os) indicus (Fauvel , Rei sh , 1968 .

TM icr osp io m icr ocer e (Dor sey , Ba i ley-B rock et al . , 1980 .

Po lydora armata La nger hans , 1 880: Woodwick, 1968 ; Ba i ley-B rock et al . , 1 980 .

Po lydora cf. P . armata Langer hans : H ar tman , 1954 .

Po lydora tr iden ticu lata Woodw ick , 1964 : Woodw ick , 1964 ; Reish, 1 968 .

‘The author s and dates for species have been checked us ing H ar tman ( 1 959) and Day Refer ences fo l lowi ng the co lon a

Enewetak records .

TNeeds r e-

conf i r rnat ion , poss ibly refer r able to M icr osp io gr anu late B lake and Kudenov , 1 978 (L inda Ward, per sonal commun ication) .(Th is table con tinued on next page

POLYCHAETES

TABLE 1 (con t'd)Fam i ly SPION IDAE (cont’d)

M in usp io c i r r ifer a (W i ren , Foster , 1 971 .

P r ionOSp io ci r r ifer a W i ren , 1 883 : Reish, 1 968 .

Pseudopo lydora an tennata (Claparede, Woodw ick , 1964 ; Reish, 1 968 ; Bai ley-Br ock et ai . , 1 980 .

Pseudopo lydora corall icole Woodw ick , 1964 : Woodwi ck, 1964 ; Reish , 1968 .

Pseudopo lydor a p igmen tate Woodw ick , 1 964 : Woodwi ck , 1964 ; Rei sh , 1 968 .

Cerez ielle r eishi (Woodwick , B lake, 1 979 .

PseudOpolydora reishi : Woodw ick , 1 964 ; Reish , 1 968 ; Bai ley-Brock et al . , 1 980 .

Sco lelep is (Sco lelep is) bonn ier i (Mes n i l , Reish , 1 968 .

Sp io sp . : H ar tman , 1 954 .

Tr ipo lydora sp inosa Woodwi ck , 1964 : Woodw ick , 1 964 ; Reish , 1 968 ; B lake and Woodw ick, 1 98 1 .

Fam i ly CH AETOPTERIDAEChaetopterus sp . : Rei sh , 1 968 .

Mesocheetop ter u s segi ttar ius (Claparede, Bai ley-Br ock et al . , 1 980 .

Mesocheetop ter us m inutas Pott s , 1 91 4 : H ar tman , 1 954 ; Reish, 1 968 ; Kohn , 1 98 1 .

Phyllochaetop ter u s ramosus W i l ley , 1 905 : Reish , 1968 .

Phyllochaetop ter us racemosus : Ba i ley-B rock et al . , 1980 .

Phyllochaetop ter us soc ial is Claparede, 1 870 : Reish, 1 968 .

Phyllochaetop ter us p ictus Cr os s land , 1 903 : H ar tman , 1 954 .

Phyllochaetop ter us sp . cf . P . ar abicus Grube, 1870 : Kohn , 1 98 1 .

Fam i ly SCALIBREGMIDAEH yboscolex long iseta Schmarda, 1 86 1 : Ba i ley-B r ock et al . , 1 980 .

Fam i ly MAGELONIDAEMagelona sp . : Beish , 1 968 .

Fam i ly CIRRATUL IDAECir r atu lus sp . : H ar tman , 1 954 ; Reish , 1 968 .

Ci r r iform ie punctate (Grube, Kohn , 1 981 .

Ci r r iform ie sem ic incta (Ehler s , H artman , 1 954 ; Beish , 1 968 ; Bai ley-Br ock et al . 1 980 .

Dodecacer ia loddi H ar tman , 1954 : H ar tman , 1954 ; Reish, 1 968 ; Bai ley-Br ock et al . , 1980 .

Fam i ly CTENODR IL IDAECtenodr i lus ser ratus (Schm idt , Reish , 1 968 .

Fam i ly OPH ELIIDAEA rmandia in termedia Fauvel , 1 901 .

A rmandia Ianceo late W i l ley , 1905 : Rei sh , 1 968 .

Polyophthalm us p ictus (Dujardin , H ar tman , 1 954 ; Reish , 1 968 ; Ba i ley-Br ock et al . , 1 980 .

Fam i ly CAPITELL IDAECep itella cap iteta (Fabr icius , Rei sh, 1 968 ; Ba i ley-B rock et al . , 1 980 .

Dasybranchus ceducus (Grube, H artman , 1954 ; Reish , 1 968 ; Kohn , 1 98 1 .

Dasybranchus lumbr ico ides Grube , 1 878 : H ar tman , 1954 ; Reish, 1 968 .

Dasybr anchus sp . : Bai ley-B rock et al . , 1 980 .

Decamas tus sp . : Ba i ley-B rock et al . , 1980 .

H eter omas t ides bif idus Augener , 1 91 4 : Reish , 1 968 ; Ba i ley-B r ock et al . , 1 980 .

Leiochr ides sp . : Bai ley-Brock et ai . , 1 980 .

Notomas tu s later iceus Sar s , 1 85 1 : Ba i ley-B r ock et al . , 1 980 .

Pu ll ielle sp . : Ba i ley-B r ock et al . , 1 980 .

cap i tell id sp . : Bai ley-B rock et al . , 1 980 .

Fam i ly ARENICOL IDAEB r enchiomaldane vincen t i i Langer hans , 1 88 1 : H ar tman , 1 954 ; Rei sh, 1 968 .

Fam i ly MALDANIDAEAx io thella aus t r a lis Augener , 1 914 : Reish , 1 968 .

Euclymene sp .

maldan id sp . : Ba i ley-B rock et al . , 1 980 .

Fam i ly TEREBELL IDAEEupo lym n ia t r igonos toma (Schmarda,

Eupo lymn ia t r igos toma [s ic] (Schmarda) : Rei sh, 1 968 .

Leaena m inu te H ar tman , 1 954 : H ar tman , 1 954 : H ar tman , 1954 ; Bai ley-B r ock et al . , 1 980 .

Terebel la ehrenberg i Grube , 1870 : Bei sh , 1 968 .

Terebe l la ehrenbergi Grube : H ar tman , 1 954 .

(This table continued on next page

102 DEVANEY AND BAILEY-BROOK

TABLE 1 (con t ’d)

Fam i ly TEREBELL IDAE (cont ’d)Thelepu s sp . : Beish , 1968 .

ter ebe l l id sp . : Bai ley-B r ock et al . , 1980 .

Fam i ly SABELL IDAEEuchone en iwe tokens is Reish , 1968 : Rei sh, 1 968 .

Notau lax phaeotaen ia (Schmarda,

H yp s icom u s phaeotaen ia [s ic]: H ar tman , 1954 ; Beish , 1968 .

Megalomma t r iocu latum Reish, 1 968 : Beish , 1968 .

Sabel la notata Grube ,1 878 : H ar tman , 1 954 ; Bei sh , 1 968 .

sabe ll id sp . : Bai ley-B r ock et al . , 1 980 .

Fam i ly SERPUL IDAEH ydr o ides albicep s (Grube,

Eupomatus albicep s Grube : H ar tman , 1954 ; Reish , 1968 .

Pomatos tegus s tel latus (Abi ldgaard, Bai ley-B rock unpubl .Fi logr ana imp lexa Berkeley , 1835 .

Salmacina dys ter i (H ux ley , Young , 1969 .

Salmac ina incrus tons Clarapede, 1 870 : Reish, 1968 .

Salmacina sp . : H ar tman , 1954 .

Serpula har tmanae Bei sh , 1 968 : Reish, 1 968 .

Serpula sp . : H artman , 1 954 .

Sp i robranchus giganteus (Pal las , H ar tman , 1 954 ; Rei sh, 1 968 .

Verm i l iop s is g landiger us Grav ier , 1 906 : Reish , 1968 ; Bai ley-Brock et al . , 1980 .

Verm i l iop s is sp . : H ar tman , 1 954 .

serpul id sp . : Bai ley-Br ock et al . , 1 980 .

Fam i ly SPIRORBIDAENeodexiosp ir a br as i liens is (Grube, Kn ight-Jones , 1984 .

Sp iror bis bellu lus Bush [s ic]: Young , 1969 .

Vinear ia koehler i (Caul ler y and Mesn i l , Kn ight-Jones , 1 984 .

Sp iror bis (P i leolar ia) n . sp . : Young , 1969 .

sp i ror bids , dextral : H ar tman , Reish , 1 968 .

spi r or bids , s in i str al : H ar tman , 1954 ; Rei sh, 1 968 .

sp i r or bid sp . : Bai ley-B rock et al . , 1980 .

by th is group . H ighsmi th ind icated that mos t of the

po lychaetes found in assoc iat ion w i th the corals were“

nes tler s”r ather than bioeroder s (bo rer s ) . Eun icids and syl

l ids were mos t numer ous among the po lychaete fam i l ies ,w i th the former hav ing a far gr eater biomass . Po lychaetes ,together w i th Sipuncu lan s , wer e found to be the mos t common inver tebrates associated w i th r ubble in H ighsmi th ’

s

s tudy . Six iden t i fied eun icids , two dor v i lleids , includ ing one

new Enewetak recor d and one lysaret id, wer e among thespecies enumer ated , together w i th undeterm ined member sfrom other fam i l ies . The fam i l y Aphr od i t idae was r ecor dedfor the fi r s t t ime from Enewetak in that paper , but no gener a or species was lis ted . A great deal of wor k on the

annel ids of Enewetak remains to be done. Th is is especially tr ue of the soft bottom in the lagoon where th isfauna has yet to be inves t igated to any extent .

One hundr ed th i r ty-two po lychaete species have beeniden t i fied fr om Enewetak Ato l l . Of them , 52 speciesar e also found in the h igh H awai ian Is lands (Bai ley-B rockand H ar tman ,

1 984) and 43 (3396) ar e known fr om the

Solomon Is lands (Gibbs , Twenty-seven Enewetakspec ies occur on the shal low cor al r eefs of Sumatra and

Thai land , and 20 are from Eas ter ls land , r epr esen t ing 23and 1596, respect ively (Kohn and L loyd , 1973a,

b) . Col

lect ions fr om H er on ls land r eefs off Aus tral ia (Reichelt1 979) compr ised mos t ly er rant po lychaetes (Eun icidae an<

Syll idae) and shar ed on l y 1 096 of the spec ies w i t f

Enewetak . All these co l lect ions were fr om s im i lai

habi tats — har d subs trata of shal low coral r eefs —

except fO iReichelt

’

s , wh ich was r es tr icted to the infauna of sand ancr ubble fr om under neath “

smal l cor al boulder s . Thesenumber s sugges t faunal s im i lar i t ies of Enewetal

po l ychaetes w i th both the So lomon and H awai ian ls landsThe h igh num ber for H awai i pr obably r eflects more in ten's ive sampl ing and r esear ch ther e than elsewhere in the

tropical Paci fic . It is expected that the s im i lar i ty betweerEnewetak and wes t Pac i fic locat ions w i ll incr ease as the

po lychaetes from those ar eas become better known .

Among the Enewetak po l ychaetes , two s yll id species(H ap losyll is spong icola and Exogone ver uger a) and two eun

~

icids (Nematoner eis un icom is and Pa lo la s ici l iens is ) havebeen found at the Ind ian and Paci fic Ocean locationsmen t ioned above. Other species wi th br oad Pac i ficd i s tr ibut ions ar e the amphinom id Eu rythoe comp lanata,

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M o llus ca of Enewetak Ato ll

E . ALISON KAY and SCOTT JOH NSON

Un iver s ity of H awai i at Manoa,

H ono lu lu, H awai i 96822

IN TRODUCTIONThe mo l lusks of the Mar shall Is lands were fi r s t s tudied

by Demond for a summar y of the eco logy and

zoogeog raphy of cen tr al Paci fic mo l lusks . H er summar ywas based on co l lect ions made between 1 944 and 1 957 in

the Mar shal l , Caro l ine,G i lber t , and Mar iana Is lands .

A lthough Demond’

s paper does not spec i ficall y ci te spec iesfrom Enewetak , her wor k was der ived in par t from s tudyof col lect ion s made in 1 946 and 1 947 at Enewetak by thelate J . P . E . Mor r ison of the Nat ional Museum of Natur alH is tor y dur ing Operat ion Cross r oads and fr om co l lect ionsmade in 1 952 at Enewetak by H . S. Ladd , M . Rus sel l ,

‘Ear l ier r ecords of the mo l lusks of the Mar shal l Is lands include

those of Es cho ltz H offman H atai and

Di etr ich and Mor r is None mentions Enewetak .

Chapter 1 4

“

(Abs tr acts fr om the annual EMEL (later known as MPML )r epor ts ar e cited on ly i f we could not tr ace subsequent publ icat ion .

TABLE 1

Anno tated Checkl i s t of Mol lusks fr om Enewetak Atol l‘

Phy lum MOLLUSCAClass GASTROPODASubclass PROSOBRANCH IAOrder ARCH AEOGASTROPODAFam i ly H AL IOT IDAEH al iot is clathrata Reeve, 1 846 ; Talmadge, 1963 : 1 32 ; as H . ruber clathrata Reeve.

H a liot is gemma Reeve, 1 846 ; Talmadge, 1 963 : 1 36 , pl . 1 4 , F ig . 8 ; as H . jacnen s is Reeve.

H . ovina Gmel in , 1 791 ; [Cer nohor sky, 1 972 : 33 , pl . 2, F ig .

H . p lanata Sower by , 1 833 ; [T almadge, 1 963 : 1 35 , pl . 1 4 , F ig .

H . van'

a L innaeus , 1 758 ; [Cernohor sky, 1 972 : 34 , pl . 2 , F ig .

‘T hi s check l i st r epr esents all of the mo l lusks pr esent ly known fr om Enewetak , both l iv ing and foss i l . Specimens of species r ecorded as

l iv ing at Enewetak are depos ited in the National Museum of Natural H isto r y , Wash ingt on , D C , and/or Ber n ice P . B ishop Museum , H ono

lulu , H awai i . The fo l lowing conventions ar e used in the check l i st : ( 1 ) species found l iv ing at Enewetak are l isted wi th a ci tation to a r eference fi gur e in squar e brackets ; where species have been pr ev ious ly r ecorded , the reference is also cited ; (2) fam i l ies are gi ven in app r oxi

mate phy logenetic sequence, and wi th in each fam i ly the genera and species are l isted alphabetical ly .(This table cont inued on next page. )

and R . C . Townsend of the U . S. Geo log ical Su r vey .T hese Enewetak col lect ions ar e depos i ted in the Nat ionalMuseum of Natu r al H is to r y (USNM ), Wash ington , D . C .

Another reference col lect ion of Enewetak mo llusks was

begun in 1 955 when appr ox imately 1 80 spec ies co l lectedby A . D . H inckley wer e determ ined by Demond (H iatt ,1 956 ,

1957) and depos i ted at the Enewetak Mar ine B iolog ical Labo rator y (EMEL ) . Subsequently , cephalopods collected and iden t i fied by S. J . Towns ley (H iatt , 1 957) andother mo l lusks co l lected by EMEL v i s i tor s and by theauthor s wer e added to the EMEL co l lect ions . A checkl is tof the Mo l lusca of Enewetak A to l l is prov ided in Table 1 .

Recor ds of the mo l lusks of Enewetak ar e found in a

w ide r ange of publ icat ions T Monogr aph ic tr eatments of

fam i l ies and gener a wh ich r ecor d shel ls from Enewetak ar ethose on As s im inea (Abbott , Str ombidae (Abbott ,1 960 , 1 96 1 ; Jung and Abbo tt , Vas idae (Abbo tt ,

KAY AND JOH NSON

TABLE 1 (con t’d)Fam i ly SCISSURELL IDAESc issurel la concinna A . Adams , 1 862 ; [H abe , 1 951 : 5 , pl . 1 1 , Figs . 10 ,

S . coronata Watson , 1 886 ; Ladd , 1966 : 26— 27 , pl . 2, Figs . 7 , 8 ; H o locene.

S . decl inans Watson , 1 886 ; Ladd, 1 966 : 27 , p l . 2, F ig . 6 ; H o locene.

S . equator ia H edley , 1 899 ; Ladd, 1 966 : 27 , pl . 2, Figs . 9, 1 0 ; H o locene.

Fam i ly FISSURELLIDAED iodora [oveolata Gar r ett , 1 872 ; F ig . 1A .

D . gr an ifera (Pease, Ladd, 1966 : 3 1 — 32 , pl . 3 , Figs . 3 , 4 ; H o locene; [Kay, 1979 : Figs 1 1 E— F].

TD . mar shallens is Ladd , 1 966 ; Ladd, 1 966 : 3 1 , pl . 3 , Figs . 1 , 2 ; late Miocene.

D . r uppell i (Sower by , (Kay, 1 979 : 42, Figs . 1 IC— D].Emargi nu la bicancellata Montr ouz ier , 1 860 ; Ladd, 1966 : 28 , pl . 2 , Figs . 1 1 , 1 2 ; H o locene.

E . concinna A . Adams , 1 852; Figs . 1 8 , 1C.

l’

E . cf . E . clypeus A . Adams , 1 851 ; Ladd , 1 966 : 28 , p l . 2, Figs . 15 , 16 ; H o locene.

E . cf . E . di lecta A . Adam s , 1 852 ; Ladd, 1 966 : 28 , pl . 2, Figs . 1 3 , 1 4 ; as Emarginu la sp . cf . peas ei Th iele, 1915 ;

H o locene.

E . n igr omacu lata Th iele, 1 91 5 ; F ig . 1D .

E . souverbiana Pi lsbry, 1 890 ; Ladd , 1966 : 28 , pl . 2 ; Figs . 1 7 , 18 ; H o locene.

“

l’

Emarginu la sp . B ; Ladd , 1 966 : 29 , pl . 2, Figs . 21 , 22; H olocene.

Punct u rella sp .

‘

l H em i toma bikin iens is Ladd , 1 966 ; Ladd , 1 966 : 29 , pl . 2, Figs 24 , 25 ; H o locene.

TH . (Mon tfor t ia) sp . A ; Ladd, 1 966 : 29 , p l . 2, Figs . 26 , 27; H o locene.

R imu la mar iei Cr osse, 1 866 ; Ladd, 1 966 : 30 , pl . 2, Figs . 32, 33 ; as R imu la sp . ; M iocene? ; H o locene.

TScu tus sp . A . ; Ladd , 1966 : 3 1 , pl . 2 , Figs . 34 , 35; late M iocene.

l'Scu tus sp . B . ; Ladd , 1966 : 3 1 , pl . 2 , Figs . 36 , 37 ; late M iocene .

Fam i ly PATELLIDAEPatel la flexuosa (Quoy and Gaimard, Ladd , 1966 : 32 , pl . 3 , F ig . 7; as P . s tellaeform is Reeve, 1 842;

H o locene.

Fam i ly ACMAEIDAEPatello ida sp .

Fam i ly TROCH IDAEAngar ia delph inus (L innaeus , [Cernohor sky, 1972 : 42 , text F ig .

As tele engebiens is Ladd , 1 966 ; Ladd , 1 966 : 37 , pl . 4 , Figs . 1 1 — 13 ; ear ly M iocene; F ig . I F .

Call ios toma sp ; Zmar z ly, 1 984 : 1 5 ; F ig . 1 E .

Clancu lus at ropurpureus (Gould , Zmar zly, 1 984 : 1 1 5 ; [Cernohor sky, 1978 : 32 , pl . 8 , F ig .

C . clangu lo ides (Wood , [Cer nohor sky, 1 972 : 40 , pl . 8 , F ig .

C . margar i tar iu s Ph i l ippi , 1 847 ; [Cer nohor sky, 1 972 : 40 , p l . 8 , F ig .

Euchelu s angu latus Pease, 1 867 ; Ladd, 1 966 : 34 , pl . 3 , Figs . 1 7— 19; H o locene.

E . at ratus (Gmel in , [Cer nohor sky, 1972 : 40 , pl . 8 ,F ig .

E . foveolatu s (A . Adams , [Mar shal l , 1 979 : 524 , Figs . 2A— E].E . ins tr ictu s (Gould , Ladd, 1 966 : 33 , pl . 3 . Figs . 1 1 - 1 3 ; Miocene?; P leistocene.

E . pauper cu lus (L ischke, [Cer nohor sky, 1 978 : 33 , pl . 8 , F ig .

TE . cf . E . quadr icar inatus (D i llwyn , Ladd , 1 966 : 33 , pl . 3 , Figs . 9 , 1 0 ; late? M iocene; H o locene.

”

( Euchelus sp . A . ; Ladd , 1 966 : 34 , pl . 3 , F igs 20— 22 ; H olocene.

TG ibbu la engebiens is Ladd , 1966 ; Ladd , 1966 : 36 , p l . 4, Figs . 6 , 7 ; ear ly Miocene.

G . marmorea (Pease, Ladd, 1966 : 36 , pl . 4 , Figs . 8— 1 0 ; as Fossar ina hoffmeis ter i Ladd , 1 966 ; M iocene.

H ybochelu s cance llatus or ien ta lis P i lsbry, 1904 ; Ladd , 1966 : 34— 35 , pl . 3 , Figs . 23 , 24 ; H o locene.

l sanda op icina (Gould, Ladd, 1 966 : 39— 40 , pl . 5, Figs . 1 — 4 ; H o locene .

Mon i lea belcher i (Ph i l ippi , [Cernohor sky, 1 972 : 41 , pl . 8 , F ig .

M . dan ieli Cr osse 1862 ; [Cer nohor sky, 1 972 : 41 , pl . 8 , F ig . 1 2; as M . phi l ipp iana Dunker ,TM . mar shallens is Ladd, 1966 ; Ladd, 1966 : 40 , pl . 5 , Figs . 9— 1 2; ear ly M iocene.

M . vem icosa Gould, 186 1 ; Ladd , 1 966 : 40— 41 , pl . 5, Figs . 1 3 , 14 ; as M . l ifuana Fischer , 1 878 ; H o locene;[Cer nohor sky, 1 978 : pl . 9 , F ig .

Tectus fenes tratus (Gmel in , [Cer nohor sky, 1 972 : 39, pl . 6 , F ig .

T . hi r asei Pi lsbry , 1 904 ; [Pi lsbry, 1 904 : 32 , pl . 5 , Figs . 52, 52a].T . pyr am is (Bo r n , [Cer nohor sky, 1 972 : 39 , pl . 8 , F ig .

l Thalo t ia berauens is (Beets , Ladd, 1 966 : 34— 35, pl . 3 , Figs . 27 , 28 ; late M iocene.

T . g i lber t i (F ischer , [Cer nohor sky, 1 978 : 34 , p l . 8 , F ig. 1 0 ; as Can thar idus].Tosat rochu s atten uatus (Jonas , [Ladd , 1 966 : 35 , pl . 3 , F ig . 29 ; as Thalot ia aff . elongatus (Wood ,Trochus h is tr io Reeve, 1 848 ; Ladd, 1 966 : 37 , pl . 4, Figs . 1 6 — 1 8 ; H o locene; [Cernohor sky, 1978 : 32, pl . 8 , F ig

T . n i lot icus L innaeus , 1 767 ; in tr oduced by the Japanese about 1935 ; [Cer nohor sky, 1972 : 38 , pl . 8 , F ig .

‘l‘Species not now l iv ing at Enewetak ; ho r i zons r ecorded by Ladd (1 966 , 1 972, 1977 , (This table continued on next page.)

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108 KAY AND JOHNSON

TABLE 1 (con t'd)

Fam i ly NERIT IDAE (cont ’d)TP is u l ina adam s iana G . and H . Nev i l l , 1 869 ; Ladd , 1 977 : 1 4 , pl . 1 , Figs . 1 , 2 ; H o locene.

Puper ita cf . P . ben son i (Recluz , F ig . 1 P , l Q .

TSmaraqdia co lei Ladd , 1966 ; Ladd , 1966 : 58 — 59 , pl . 1 1 , Figs . 5— 7; Miocene.

TS . joqjacar tens is (Mar t in , Ladd , 1 966 : 58 , pl . 10 , Figs . 28— 3 1 ; M iocene.

TSmaraqdia sp . A . ; Ladd, 1966 : 59 , pl . 1 1 , Figs . 8 , 9 ; late M iocene.

[Ner ita p icea Recluz , 1 841 , r epor ted by Stokes (1966) and Kay (1979) has not been confi rmed].Fam i ly NERITOPSIDAENer i top s is radu la (L innaeus , Ladd , 1 966 : 55 pl . 1 0 , Figs . 1 2— 14 ; M iocene.

Fam i ly PH ENACOLEPADIDAEPhenaco lepas granocos tata (Pease, [Kay, 1 979 : 68 , F ig . 21A].P . scobinata (Gould , [Kay, 1979 : 68 , F i g .

Order MESOGASTROPODAFam i ly L ITTOR IN IDAEL i ttor ina coccinea (Gmel in , Rosewater , 1 970 : 439 , p l . 325 , Figs . 1 1 , 1 2 .

L . p in tado (Wood , Rosewater , 1 970 : 447 , p l . 325 , Figs . 1 5, 16 .

L . scabra L innaeus , 1 758 ; Rosewater , 1 970 : 436 , pl . 325 , Figs . 8— 10 .

L . undu lata Gray , 1 839 ; Rosewater , 1 970 : 298 , pl . 325 , F igs . 8— 1 0 .

Nod i l i ttor ina m i llegr ana (Ph i l ippi , Rosewater , 1970 : 491 , pl . 378 , Figs . 1 3 , 1 4 .

Peas iella tan t i l la (Gou ld , [Kay, 1979 : 74, F ig . 24A].‘l‘Tectar ius rehder i Ladd, 1966 ; Ladd , 1966 : 59 , pl . 1 1 , Figs . 1 1 — 1 3 ; M iocene.

Fam i ly RISSOIDAEA lvan ia crys talline Gar r ett , 1 873 ; F ig . I L .

A . iso lata (Laseron , [Kay, 1 979 : 77, F ig . 27C].B r ooku la sp . , F ig . 1M .

TCinqu la par ryens is Ladd, 1 966 ; Ladd, 1966 : 6 1 , pl . 1 1 , Figs . 23 , 24 ; M iocene.

C . cf . C . r oseoc incta (Suter , Ladd, 1966 : 61 , p l . 1 1 , Figs . 25, 26 ; M iocene; H o locene.

TMerel ina p is inna (Melv i l le and Standen , Ladd, 1966 : 63 , pl . 1 2, F igs . 4 , 5; Miocene.

M . wanawana Kay, 1979 ; [Kay, 1979 : 82, Figs . 28 A , B].Parashiela beets i Ladd , 1966 ; Ladd, 1 966 : 64 , pl . 12 , Figs . 8 , 9 ; H o locene.

TPus i ll ina je/fcoat i (Ladd , Ladd , 1 966 : 62 , pl . 1 1 , Figs . 27— 3 1 ; M iocene; H o locene; Ponder , 1985: 30 .

P . marmorata (H edley , [Kay, 1979 : 79, Figs . 27 E, F ; as Vitr icithna].TPu t i lla morana Ladd , 1 966 ; Ladd, 1966 : 1 6 , pl . 1 1 , Figs . 1 6 , 1 7; Miocene .

TR isso ina abbott i Ladd , 1 966 ; Ladd , 1 966 : 70 , pl . 1 3 , Figs . 1 9- 21 ; M iocene.

R . abrardi (Ladd , Ladd , 1 966 : 65 , pl . 1 2, Figs . 1 5— 1 8 ; as Zebina; ea r ly M iocene; H o locene; Ponder ,1 985 ; 80 .

TR . ai l inana Ladd , 1 966 ; Ladd , 1 966 : 70-71 , p l . 1 3 , F ig . 26 ; M iocene.

TR . alex is i Ladd , 1966 ; Ladd , 1 966 : 70 , pl . 1 3 , F igs . 1 3 — 16 ; M iocene.

R . ambiqua (Gould , La dd , 1966 : 71 , pl . 1 4 , Figs . 23 , 24 ; H o locene.

TR . ambiqua par r yens is Ladd , 1 966 ; Ladd, 1 966 : 72 , pl . 1 4 , Figs . 25 , 26 ; ear ly M iocene.

R . andaman ica Weinkau ff , 1 885 ; Ladd , 1966 : 67, pl . 1 2 , Figs . 22— 25 ; as R . m ijana Ladd, 1 966 ; ear ly Miocene;H o locene; [Cer nohor sky, 1 978 : 44 , pl . 1 1 , F ig .

R . balteata Pease, 1 869 ; Ladd , 1966 : 69, pl . 1 3 , Figs. 5— 8 ; ear ly Miocene; H o locene.

TR . bik in ien s is Ladd , 1 966 ; Ladd , 1 966 : 69, p l . 1 3 , F igs . 9 , 1 0 , 1 7 , 1 8 ; M iocene ; Ponder , 1 985 : 84 .

R . clathrata (A . Adam s , [Cernohor sky, 1 978 : 47 , pl . 12, F ig .

TR . concinna A . Adam s , 1 851 ; Ladd , 1 966 : 72 , p l . 1 3 , F ig . 33 ; M iocene.

R . ekkanana Ladd, 1 966 ; Ladd, 1 966 : 71 , p l . 1 3 , Figs . 3 1 , 32; H o locene.

TR . emnanana Ladd, 1 966 ; Ladd, 1 966 : 67 , pl . 12, Figs . 3 1 , 32 ; Miocene.

R . fimbr iata (Souver bie, [Cernohor sky, 1 978 : 45 , p l . 1 1 , F ig .

R . cf . R . flex uosa Gould , 1 86 1 ; Ladd , 1 966 : 66 , pl . 1 2 , Figs . 21 , 22 ; H o locene .

TR . her r ingi Ladd , 1 966 ; Ladd , 1 966 : 74 , F ig . 1 4 , Figs . 13,14 ; late M iocene.

R . i llus tr is Sower by , 1 894 ; Ladd 1966 : 66 , pl . 1 4 , Figs . 5- 8 ; as R . epham i lla Watson , 1886 ; ear ly M iocene;H o locene.

R . im br icata Gould , 1 832 ; [Kay, 1979 : 86 , F ig . 29L ; as Zebina].TR . cf . R . indr ai Beets , 1 94 1 ; Ladd, 1 966 : 66- 67 , p l .

'

12 , Figs . 23 , 24 ; ear ly Miocene.

R . lam ber t i Sou ver bie, 1870 ; [Cer nohor sky, 1 978 : 46 , pl . 1 2 :TR . lomaloana Ladd , 1 966 ; Ladd , 1 966 : 72 , pl . 13 , Figs . 27 , 28 ; Miocene.

TR . mar shallens is Ladd , 1 966 ; Ladd , 1 966 : 73 , pl . 1 3 , Figs . 38 — 40 ; pl . 14 , Figs . 1 — 4 ; M iocene.

R . meji lana Ladd , 1966 ; Ladd , 1 966 : 67,pl . 1 2, Figs . 25 , 26 ; ear ly M iocene .

‘l‘Spec ies not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd ( 1 966 , 1 972, 1 977, (Th is table conti nued on next page. )

MOLLUSCA 109

TABLE 1 (con t'd)

Fam i ly R ISSO IDAE (con t'd)R . p licata A . Adams , 1 851 ; Ladd , 1 966 : 74— 75 , pl . 1 4 , Figs . 21 , 22 ; ear ly M iocene.

R . ret icu lata (Sower by, [Cer nohor sky, 1 978 : 46 , pl . 1 2 :TR . cf . R . s up r acos tata Gar r ett , 1 873 ; Ladd, 1 966 : 68 , p l . 1 2 , Figs . 35 , 36 ; ear ly Miocene.

R . tenu is t r iata Pease, 1 868 ; Ladd , 1 966 : 68 , p l . 1 2 , Figs . 33 , 34 ; ear ly Miocene; H o locene.

R . tor nat i l is Gould , 1 86 1 ; Ladd , 1 966 : 69 , pl . 1 3 , Figs . 1 1 , 1 2; as R . transenna Watson , 1 886 ; ear ly Miocene;H o locene; [Cer nohor sky, 1 978 : 48 , p l . 12 :

R . tur r icu la Pease, 1 861 ; Ladd, 1 966 : 72 , pl . 13 , Figs . 36 , 37; Miocene; H o locene.

Schwar tz iella t r it icea (Pease, [Kay, 1979: 86 , F ig . 29l].

TStos icia gardnerae (Ladd , Ladd, 1 966 : 60 , pl . 1 1 , Figs . 1 4 , 1 5 ; as l r avadia; Miocene; Ponder , 1 985 : 94 .

TZebina cf. 2 . cooper i (O l iver , Ladd, 1 966 : 65 , p l . 12, Figs . 1 1 , 12; as Zebina; H o locene.

Z . ki lleblebana Ladd, 1 966 ; Ladd , 1 966 : 65 , p l . 12, Figs . 13 , 14 ; ear ly Miocene.

Z . sem ip l icata (Pease,Ladd , 1 966 : 64 , pl . 1 2, F ig . 10 ; as Z . metalt i lana Ladd, 1 966 ; ear ly Miocene; H o locene;

[Kay, 1 979 : 86 , F ig . 29K].2 . s tn

’

atu la (Gould, [Kay, 1979: 86 , F ig . 29L].Z . tri den tata M ichaud, 1830 ; [Kay, 1 979 : 86 , F ig .

Fam i ly ASSIMINEIDAETAs s im inea n it ida en iwetokens is Ladd , 1 966 ; Ladd, 1966 : 75 , pl . 1 0 , F ig . 8 ; Miocene.

A . n i t ida mar shallens is Abbo tt , 1958 ; Abbott , 1958 : 256 , pl . 15, F ig . 8 ; ho lotype fr om Japtan Is land, Enewetak .Fam i ly BARLEEIDAETP rotobar leeia myer s i (Ladd, Ladd, 1966 : 62 , p l . 1 1 , Figs . 32, 33 ; as Amph ithalamus (Cer atos traca?) myer s i ;late M iocene; Ponder , 1 983 : 243 ; [Ponder , 1 983b : 243 , F igs . 1 1E— I]

Fam i ly CAECIDAETCaecum berberense Ladd , 1 972 ; Ladd, 1 972: 22, pl . 5 , Figs . 1 1 , 1 2 ; ear ly Miocene; P l iocene.

C . cf . C . fo l in i Kisch , 1 959 ; F ig . 28 .

C . c]. C . g label lum A . Adams , 1 868 ; [Kay, 1 979 : 1 10 , F ig . 4_

2G].TC . par ryen s is Ladd, 1 972; Ladd, 1 972: 23 , pl . 5 , Figs . 1 — 7; ear ly Miocene.

C . sep imen tum de Fo l in , 1 867; [Kay 1979 : 109, F ig .

TO ver tebrale H edley , 1 899 ; Ladd , 1 972 : 22 , pl . 5, F igs . 8— 10 ; P leis tocene; H o locene.

TElephan tenellum sp . A . ; Ladd , 1972 : 23 , pl . 5 , F ig . 15 ; pos t-M iocene.

‘l’

Far tu lum cf . F . ampu tatum (H edley , Ladd, 1 972 : 24 , pl . 5, Figs . 1 6 , 1 7 ; ear ly M iocene.

TFar tu lum sp . A . ; Ladd, 1972 : 24 , pl . 5 , F ig . 1 8 ; pos t-M iocene.

M icr anellum schlanger i Ladd, 1 972 ; Ladd, 1972 : 23 , pl . 5 , Figs . 13 , 1 4 ; Miocene; H o locene.

Fam i ly VITRINELL IDAELeucor hyncha caledon ica Cr osse, 1 867 ; Ladd , 1 966 : 76 , p l . 1 4 , Figs . 30 , 3 1 ; H o locene.

L . crossei T r yon , 1 888 ; Ladd , 1966 : 78 , pl . 1 4 , Figs . 32 , 33 ; H o locene.

TL . s tephenson i i Ladd, 1 966 ; Ladd, 1966 : 76 , pl . 1 4 , Figs . 34, 35 ; ear ly M iocene.

L iot ina cf . L . bo tan ica (H edley , Ladd, 1 966 : 46 , pl . 7, Figs . 9— 1 1 ; H o locene.

L . locu losa (Gould , Ladd , 1 966 : 46 , pl . 7 , Figs . 1 2— 14 ; H o locene.

L . peron i i (K iener , [Cer nohor sky, 1 972 : pl . 1 0 , F ig .

L iotina sp . A . ; Ladd , 1966 : 47 , pl . 7 , Figs . 15— 1 7; ear ly Miocene .

Lophocochl ias m inu tiss imus (Pi lsbry, Ladd, 1 966 : 77, pl . 1 5, Figs . ear ly Miocene.

TL . paucican'

natus Ladd, 1 966 ; Ladd, 1 966 : 77, p l . 1 5 , Figs . 6— 8 ; ear ly Miocene.

TLyd iphn is enewetokens is Ladd , 1 966 ; Ladd , 1 966 : 79 , pl . 16 , Figs . 6— 8 ; ear ly M iocene; Pl iocene.

Mar elepetopoma sp . ; F ig . 1 q .

M icr ol iot ia dau tz enbergi (Bavay , F ig . I R .

TMunditella par ryen s is Ladd, 1 966 ; Ladd , 1 966 : 78 , pl . 1 5, Figs . 1 2- 1 4 ; late Miocene; H olocene.

M . gualum (H edley , Ladd , 1 966 : 78 , pl . 1 5 , Figs . 9— 1 1 ; P leistocene.

Sanson ia andaman ica (Preston , [l redale, 1 91 7 : 372 , F ig . F ig . I N .

S. cor ayi (Ladd, Ladd_, 1 966 : 63 , pl . 12, F ig . 2; as A lvan ia (Taramellia) ; H o locene.

TS . kenneyi (Ladd, Ladd, 1 966 : 63 , pl . 12, F ig . 3 ; as A lvan ia (Taramell ia) ; Miocene.

So lar ior bis tr ican'

nata (Melv i lle and Standen , Ladd , 1966 : 79 ,pl . 1 6 ,Figs . 1 — 3 ; ear ly Miocene.

Teinos toma engebiense Ladd, 1966 ; Ladd, 1 966 : 78 , pl . 1 5 ,Figs . 1 5— 1 7 ; H o locene.

T . mar shallen se Ladd , 1 966 ; Ladd , 1 966 : 78 , pl . 1 5 ,F igs . 1 8 — 20 ; ear ly M iocene.

T . par vu lum H edley , 1899; [H edley , 1 899 : 553 , F ig .

Teinos toma sp . ; F ig . l l , N .

Fami ly ORB ITESTELL IDAEOr bites tella sp . ; F ig . 1G , I H .

“

I'Species not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd ( 1 966 , 1972 , 1 977 , (Thi s table cont inued on next page. )

1 10 KAY AND JOHNSON

TABLE 1 (con t’d)

Fam i ly OMALOGYRIDAEOmalogyra japon ica (H abe, [Kay, 1979 : 92 , F igs . 32A— C].Fam i ly CINGULOPSIDAERufodardanu la sp . ; F ig . 1K .

Fam i ly TURRITELLIDAE‘l'Tu r r i tella cf . T . c ingu l ifera Sower by , 1 825; Ladd, 1 972 : 1 4 , pl . 1 , F ig . 1 ; ear ly Eocene to ear ly M iocene.

Tur r i te l la sp . A . ; F ig . 2C.

Tur r ite l la sp . B . ; F ig . 20 .

Verm icu lar ia sp . A . ; Ladd 1 972 , pl . 2 , Figs . 5— 7 ; ear ly Miocene; F ig . 2K .

Fam i ly TRUNCATELL IDAETruncatel la guer in i i A . and J . V i lla, 1 84 1 ; [Cernohor sky, 1 972 : 58 , pl . 1 2, F ig .

Fam i ly VERMET IDAEDendr opoma gr egar ia H adfield and Kay, 1 972; [Kay, 1979 : 1 03 , F ig .

D . mer ocl is ta H adfield and Kay, 1 972; [Kay, 1 979 : 104 , F ig . 39A].D . p latypus (Morch , [Kay, 1 979 : 106 , F ig .

D . p sarocephala H adfi eld and Kay, 1 972; [Kay, 1979 : 1 06 , F ig . 39D].D . r hyssoconcha H adfield and Kay, 1 972 ; [Kay, 1 979 : 1 06 , F ig . 39C].

Petaloconchus keenae H adfield and Kay, 1972 ; [Kay , 1979 : 1 08 , F ig . 40A — B].TP . lamellos us Ladd , 1 972; Ladd , 1972 : 20 , pl . 3 , F ig . 13 ; late M iocene.

‘l’

P . mer kana Ladd , 1972 ; 20 , pl . 3 , Figs . 4— 12; ear ly M iocene; H o locene.

P . tokyoens is (P i lsbry, [Pi lsbry , 1 895 : 61 , pl . 1 , Figs . 9

TSerp u lor bis cf . 8 . javan us Mar t in , 1 879 ; Ladd , 1 972 : 20— 21 , pl . 3 , Figs . 1 4, 51 ; late M iocene.

S. var iabi lis H adfield and Kay, 1 972 ; [Kay, 1 979 : 1 08 , F igs . 40 E, F].Vermetus ali i H adfield and Kay, 1 972 ; [Kay, 1 979 : 1 08 , Figs . 40 C, D].Fam i ly SIL IQUARIDAEStephopoma cf . 8 . r oseum Quoy and Gaimard, 1833 .

Fam i ly PLANAXIDAEPlanax is su lcatus (Bor n , [Cer nohor sky, 1972 : 58 , pl . 1 2, F ig .

P . zonatus A . Adam s , 1 851 ; F ig . 2A .

Fam i ly CER ITH I IDAEB i tt ium impenden s (H edley , Ladd, 1 972 : 30 , p l . 7 , F ig . 1 5 ; H olocene.

B . sergen tum (Jousseaume, Ladd , 1972 : 30— 3 1 , pl . 7 , Figs . 16— 20 ; ear ly Miocene; H o locene.

B . zebr am (K iener , Ladd , 1972 : 3 1 , pl . 7, Figs . 21 , 22; as B . ian thinum ; H o locene; Ladd , 1 972 :3 1 , pl . 4 , F ig . 1 3 ; as B . en iwetoken s is Ladd , 1972 ; ear ly M iocene; Ladd , 1972 : 32, pl . 4 , Figs . 1 4— 1 6 ;

as B . toddae; late M iocene.

Cer i th ium alveo lus H ombr on and Jaqu inot , 1 84 1 ; [Cernohor sky, 1 972 : 68 , pl . 15 , F ig .

C . atr omargi natum Dau tzenberg and Bouge, 1 933 ; [Kay, 1 979 : F ig . 45 F].C . claviforme Schepman , 1 90 1 ; F ig . 2L .

C . co lumna Sower by , 1 834 ; Ladd, 1 972 : 39 , pl . 1 0 , F ig . 2 ; H o locene .

C . echinatum (Lamarck , Ladd, 1 972 : 37 , pl . 9 , F ig . 6 ; H o locene; as C . m u tatum Sower by , 1 834 ; H oubr ick ,

1 974 ; [Kay , 1 979 : 1 22, F ig . 45L ; as C . mu tatum Sower by ,C . egenum Gould , 1 849 ; Ladd, 1 972 : 39 , pl . 9 , Figs . 9 , 1 0 ; H o locene.

TC . en iwetokens is (Ladd, Ladd, 1 972 : 37; pl . 9, Figs . 1 , 2 ; Miocene; as Rhinoclavis .

TC . floranen s is (Ladd, Ladd, 1 972 : 37, pl . 9 , Figs . 4 , 5 ; M iocene; as Rhinoc lavis .

cf . in ter s tr iatum Sower by , 1 855 ; [Kay, 1 979 : 122, F ig . 45R].

T mar shallens is (Ladd , Ladd, 1972 : 34 — 35 , pl . 8 , Figs . 1 2, 1 3 ; M iocene; as Rhinoc lavis .

mun itum Sower by , 1 855 ; Ladd, 1 972 : 39 , pl . 10 , F ig . 2 ; H o locene; as Cer ith ium aff . C . co lumna.

nes iot icum (P i lsbry and Vanatta, [Kay, 1979 : 1 23 , F ig .

nodu los um Brugu ier e, 1 792; H oubr ick , 1971 : 560-565 ; 1 974 : 20 ; [Cer nohor sky, 1 972 , pl . 1 3 , F ig

cf . C . p lanum Anton , 1 839 ; [Cer nohor sky, 1 972 : pl . 1 4 , F ig .

r os tr atum Sower by , 1855 ; Ladd, 1972 : 32 , pl . 4 , F ig . 1 7; as Co lina ros trata; P leis tocene; H o locene.

salebr os um Sower by , 1 855 ; Ladd , 1972 : 37 , pl . 9, F ig . 1 2 ; P l iocene; H o locene.

tenu ifilos um Sower by , 1866 ; Ladd , 1972 : 38 , pl . 9, F ig . 1 5 ; as C . tene l lum Sower by , 1855; P leis tocene; H o locene;[Cer nohor sky, 1 972 : 70 , pl . 1 4 , F ig .

C . t rai l i i (Sower by , [Cernohor sky, 1972 : 71 , pl . 1 6 , F ig .

Clypeomor us bat i llar iaeform is H abe and Kosuge, 1 966 ; Ladd, 1 972 : 37, pl . 9, Figs . 7 , 8 as Cer i th ium alveo lusH ombron and Jaqu inot , 1841 ; H o locene; H oubr ick , 1 974 ; as Cer ith ium sejunctum l r edale; [H oubr ick ,1 985 : 5 1 — 52, F ig .

o

o

o

o

o

o

o

o

o

TSpecies not now l iv ing at Enewetak ; hor i zons recorded by Ladd (1 966 , 1 972, 1977, (This table cont inued on next Page

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KAY AND JOH NSON

TABLE 1 (con t ’d)

Fam i ly CERITH IOPSIDAE (cont ’d)J . uveanum (Melv i ll and Standen , [Kay, 1979 : 1 28 , F ig . 46H].

TSei la waluens is Ladd , 1 972 ; Ladd , 1972 : 44 , pl . 1 1 , F i g . 1 3 ; late M iocene.

Fam i ly EUL IMIDAEBalc is acicu la (Gould , Lamber son , 1978 ; F ig . 38 .

B . bryan i (P i lsbry, [Kay, 1 979 : 1 61 , F ig . S6I].B . so lida (Sower by , [Kay, 1 979 : 1 62 , F ig . 56K].B . thaanum i (P i lsbry, [Kay, 1979 : 1 64 , F ig . 56M].

TBalcis sp . A . Ladd, 1 982 ; Ladd, 1 982 : 55, pl . 41 , Figs . 14 , 1 5 ; as Eu lima; ear ly M iocene.

H em i leisot r acum sp . , F ig . 3C .

Le ios t raca metcalfei (A . Adams , [Kay, 1 979 : 1 65 , F ig . 56Q].M ucr onalia n it idu la Pease, 1 86 1 ; [Kay, 1 979 : 1 65, F ig . 56T].Pyr am idello ides suta (P i lsbry, [Kay, 1 979 : 84 , F ig . 29G].Stylapex sp . , F ig . 3A .

Styl ifer l inckiae Saras in and Saras in , 1 887 ; [Kay, 1 979 : 167 , F ig . S7E].Thyca cry s tal lina (Gould , [Kay, 1979 : 167, F ig . 56U].Fam i ly STROMB IDAELam bis ch i r ag r a (L innaeus , Abbott , 196 1 : 1 72 , pl . 1 2 1 , Figs . 1 0 — 1 2 ; Berg , 1 974 .

L . cr ocata [Abbo tt , 1 961 : 1 57, pl . 1 21 , F ig .

L . lamb is (L innaeus , Abbo tt , 196 1 : 1 54 , pl . 1 2 , F ig . 4 ; Berg , 1 974 .

L . scorp ius (L innaeus , Berg , 1 974 ; [Abbott , 1 961 : 1 64 , pl . 1 21 , F ig .

L . t r u ncata (H umphrey , Abbott , 196 1 : 157, pl . 1 21 : 9 ; Be rg , 1 974 .

Str om bus den tatus L innaeus , 1 758 ; Abbo tt , 1960 : 85 , pl . 1 4 : 23 ; Berg , 1 974 .

S. er yth r inus Di l lwyn , 1 81 7; Abbo tt , 1960 : 8 1 , pl . 20 , Figs . 1 — 5 .

8 . f r ag i l is (R6ding , [Abbott , 1 960 : 82 ,pl . 14 , F ig .

S . g ibber u lus L innaeus , 1 758 ; Abbott , 1960 : 143 , pl . 1 4 , F ig . 20 ; Berg , 1 974 ; Ladd, 1972 : 62,

pl . 1 9, F ig . 9, pl . 20 , Figs . 1 — 3 ; H o locene .

S . haemas toma Sower by , 1 842; Abbo tt , 1 960 : 1 1 7, pl . 1 7, Figs . 1 1 , 1 2; Ber g , 1 974 .

S. len t iginosus L innaeus , 1 758 ; Abbot t , 1960 , pl . 17 , Figs . 1 1 , 1 2 .

S. luhuanus L innaeus , 1 758 ; Abbo tt , 1960 : 1 36 , pl . 14 , F ig . 1 5 ; Berg , 1 974 ; G i llary, 1 974 ; G i llary and G i llary , 1 979 .

S . maculatu s Sower by , 1842; Abbo tt , 1 960 : 77 , p l . 20 , Figs . 24 , 25 .

T3 . m icklei Ladd , 1972; Ladd , 1 972 : 60 — 61 , pl . 1 8 , Figs . 5— 8 ; late M iocene .

S. m icrou r ceus (Ki ra, [Abbo tt , 1 960 : 71 , pl . 20 : 24 ,

S. mutabi lis Swainson , 1821 ; Abbo tt , 1 960 : 74 , pl . 20 : 1 5 , 1 6 ; Berg , 1 974 ; Ladd , 1 972: 58— 59, pl . 1 5 ,

Figs . 1 0 — 1 5 ; H o locene .

S . s inuatus L ightfoot , 1 786 ; [Abbott , 1 960 : 60 , pl . 1 7 , F ig .

S . taur u s Reeve, 1 857 ; Cer nohor sky, 1 972 : 73 , Cl . 1 7 , F ig . 5 .

S . var iabi l is Swain son , 1 820 ; [Abbott , 1 960 : 1 03 , pl . 1 4 , Figs . 2 1 ,

S . wi lson i Abbott , 1 967 ; [Abbott , 1967 : 455 , pl . 328 , Figs . 1

TSt r ombus sp . C. ; Ladd , 1972 : 6 1 , pl . 1 8 , Figs . 10 , 1 1 ; late M iocene.

Terebe l lum terebel lum (L innaeus , Jung and Abbott , 1 967 : 453 , pl . 321 .

Fam i ly VAN IKORIDAEVan ikoro cancel lata (Lamar ck , Ladd , 1 972 : 52, pl . 1 3 ,

Figs . 6— 8 ; ear ly Miocene, H o locene.

V . quer in iana (Recluz , Ladd , 1 972 : 52 , pl . 13 , Figs . 9, 1 0 ; ear ly M iocene; H o locene.

V . hel ico idea (le Gu i l lou , [Cer nohor sky, 1 972 : 86 , pl . 22, F ig .

TV . cf . V . kanakar um P i lsbry , 1 92 1 ; Ladd , 1 972 : 52,pl . 1 3 , F ig . 1 1 ; H o locene.

Fam i ly H IPPON ICIDAETH ippon ix fo liaceus Quoy and Gaimard, 1 835 ; Ladd , 1972 : 53 ,

pl . 1 3 ,Figs . 1 5— 21 ; M iocene .

TH . revellei Ladd , 1 972 ; Ladd , 1972 : 54, pl . 14 ,Figs . 1 — 1 8 ; ear ly M iocene; P l iocene.

Sabia con ica (Schumacher , Ladd , 1 972 : 54 ,pl . 13 ,Figs . 22— 27; ear ly M iocene; H o locene; [Kay, 1 979 :

1 79 , Figs . 62F — H ].Fam i ly CAPUL IDAEChei lea eques tr i s (L innaeus , Ladd , 1 972 : 52 ,

pl . 1 3 , F ig . 1 2 ; Miocene.

Cr ep idu la acu leata (Gmel in , Found on pi l ings only ; [Kay, 1 979 : 18 1 , Figs . 63G , H ].Cap u lus dan iel i (Crosse, [Cer nohor sky, 1 972; 88 ,

pl . 22 , F ig .

TCapu lus sp . A . ; Ladd, 1972 : 55 ,pl . 14 , F ig . 2 1 ; late M iocene.

TCap u lus sp . B . ; Ladd , 1 972 : 55 , pl . 1 4 , F ig . 22 ; late M iocene.

Fam i ly XENOPHORIDAEXenophor a cerea (Reeve, Ponder , 1 983a : 23 — 25 , F ig . 23 .

TSpecies not now l iv ing at Enewetak ; ho r i zons r ecorded by Ladd (1 966 , 1 972, 1977, (Th is table cont inu ed on next pag

MOLLUSCA 1 13

TABLE 1 (con t'd)

Fam i ly LAMELLARIIDAELame llar ia cf . L . berghi Deshayes , 1 863 ; F ig . SD .

Fam i ly ERATOIDAEP roterato gemma (Bavay , [Ca te, 1977: 353 , F ig .

P . s talagm ia Cate, 1 975 ; [Cate, 1 977: 347, F ig .

Fam i ly TRIVIIDAETr ivia hordacea Kiener , 1845; [Kay, 1979 : 187, F ig .

T . or yza (Lamarck , Ladd, 1 977 : 17, pl . 1 , Figs . 10— 1 2; Miocene and younger beds .

Fam i ly CYPRAE IDAECypr aea annulus L innaeus , 1 758 ; Ingram , 1947; [Burgess , 1970 : 341 , pl . 41A].

ar abica L innaeus , 1 758 ; Ladd , 1 977 : 19 ,pl . 2, Figs . 1 , 2; H o locene; [Burgess , 1 970 : 21 5 , pl . I SA].

argu s L innaeus , 1 758 ; [Burgess , 1970 : 253 , pl . 28 B— C].

asel lus L innaeus , 1758 ; [Burgess , 1970 : 1 1 8 , pl . 8H ].auran tium Gmel in , 1791 ; [Burgess , 1970 : 1 1 8 , pl . 8H ].beck i i Gaskoin , 1836 ; [Burgess , 1970 : 277 , pl . 3OL].bis tr inotata (Sch i lder and Sch i lder , [Burgess , 1970 : 271 , pl . 30C].caputserpen t is L innaeus , 1 758 ; Ingram , 1 947; [Burges s , 1970 : 1 77, pl . 1 2F].car neo la L innaeus , 1 758 ; Ingram , 1 947; [Burgess , 1970 : 20 1 , pl . 3ZO].catho l ico r um (Sch i lder and Sch i lder , [Burgess , 1 970 : pl . 13A].cau r ica L innaeus , 1 758 ; [Burgess , 1970 : 292 , pl . 3ZO].chi ldren i Gr ay , 1 825 ; Ladd, 1 977: 22, pl . 3 , Figs . 1 1 — 1 3 ; H o locene; [Burgess ,1970 : 262, pl . 29G].ch inens is Gmel in , 1 791 ; [Burges s , 1 970 : 79, pl . SH ].cicer cu la L innaeus , 1 758 ; [Burgess , 1970 : 266 , pl . 3OA].clandes tina L innaeus , 1 767; [Bur gess , 1 970 : 93 , pl . 7C].cr ibrar ia L innaeus , 1 758 ; [Burgess , 1970 : 1 89, pl . 136K].depr es sa Gr ay , 1 824 ; Ingram , 1947; as Cypraea in termedia; [Burgess , 1 970 : 21 7, p l . 1 8E].

di llwyn i (Sch i lder , [Burgess , 1 970 : 273 , pl . SOH ].

eg lan t ina Duclos , 1 833 ; [Burgess , 1 970 : 21 8, pl . 1 9A].erosa L innaeus , 1 758 ; [Burgess , 1 970 : 1 75, pl . 1 2 A— D].

f im br iata Gmel in , 1 791 ; [Burgess , 1 970 : 139 , pl . 9I].g lobu lus L innaeus , 1758 ; [Burges s , 1970 : 272, pl . 30D].goodall i Sower by , 1832; [Burgess , 1 970 : 1 03 , pl . 7aC].helvo la L innaeus , 1 758 ; Ladd, 1977: 23 , pl . 5, Figs . 1 — 3 ; late Miocene; [Burgess , 1 970 : 154 , pl . 10G].h i rundo L innaeus , 1758 ; [Bur gess , 1 970 : 286 , pl . SI F].humphreys i i Gr ay , 1 825 ; [Cernohor sky, 1967: 92, p l . 1 7:

isabe l la L innaeus , 1 758 ; Ingr am , 1 947; [Burgess , 1 970 : 41 , plcf. C . kamai (Beets , Ladd, 1 977 : 25, pl . 5 , Figs . 1 3 — 1 5 ; ear ly Miocene.

labro l ineata Gaskoin , 1 849; [Burgess , 1 970 : 1 43 , pl . 9aF].leviathan Sch i lder and Schi lder , 1 938 ; [Burgess , 1970 : 202 , pl . 1 5 , F ig . E, E1].

l imacina Lamar ck , 1 8 1 0 ; [Cernohor sky, 1 967, pl . 13 , F ig .

lynx L innaeus , 1 758 ; [Burgess , 1970 : 24 1 , pl . 24C].macu l ifera (Sch i lder , (Burgess , 1 970 : 21 7, p l .

mappa L innaeus , 1 758 ; [Burges s , 1 970 : 248 , pl . 26 A , C— D].

margar i ta (Sch i lder and Sch i lder , [Cernohor sky, 1 967 : 72 , p l . 1 1 :

mar iae (Sch i lder , [Burgess , 1 970 : 274 , pl . 3OI].mar tin i Schepman , 1 907 ; [Burgess , 1970 : 60 , p l . 4D].

maur i t iana L innaeus , 1 758 ; [Burgess , 1 970 : 247, pl .m icrodon Gr ay , 1 828 ; [Burgess , 1 970 : 130 , pl . 9D

— E].m inor iden s Melv i lle, 1 90 1 ; [Burges s , 1970 : 132 , pl . 9F].moneta L innaeus , 1 758 ; Ingr am , 1 947 ; Menge, 1 973 ; Renaud , 1 977 ; [Burgess , 1 970 : 343 ,

pl . 4 l B — H].nucleus L innaeus , 1 758 ; [Burgess , 1 970 : 261 , pl . 29H].onyx L innaeus , 1 758 ; [Burges s , 1 970 : pl . 6A].poror io L innaeus 1 758 ; Ingr am , 1 947 ; [Burgess , 1 970 : 1 52 ,

pl .punctata L innaeus , 1 771 ; [Burgess , 1970 : 1 04 , pl . 7aE].schi lderor um (Ir edale, [Burgess , 1 970 : 1 90 , pl . 14A].scur r a Gmel in , 1 791 ; [Burgess , 1 970 : 225 , pl . 1 9O].s taphylaea L innaeus , 1 758 ; [Bur gess , 1 970 : 259 , p l . 29E].s to lida L innaeus , 1 758 ; [Burgess , 1 970 : 290 ,

pl . 3 1E].

0

0

0

0

0

0

0

0

0

0

6

TSpecies not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd (1 966 , 1 972, 1 977, (Th is table cont inued on next page . )

1 1 4 KAY AND JOH NSON

TABLE 1 (con t ’d)Fam i ly CYPRAEIDAE (con t ’d)

talpa L innaeus , 1 758 ; [Burgess , 1970 : 199 , pl . 14F].teres Gmel in , 1 791 ; [Bur gess , 1970 : 109 , pl .tes tudinar ia L innaeus , 1 758 ; [Burgess , 1970 : 249 , pl . 27A].t igr is L innaeus , 1 758 ; [Burgess , 1 970 : 229 , pl . 22A].u r sellus Gmel in , 1791 ; [Burges s , 1970 : 287, pl .ven tr icu lus Lamar ck , 1 8 1 0 ; Ingr am , 1 947; [Burges s , 1 970 : 194 ,

pl 1 4O].vitel lus L innaeus , 1 758 ; [Burgess , 1970 : 206 , pl . 17A].

Fam i ly OVULIDAECalpum us lacteus (Lamarck , [Cernohor sky, 1 967 : 1 07, p l . 21 , F ig .

C . ver r ucosus (L innaeus , [Cernohor sky, 1967 : 107 , pl . 21 , F ig .

Pedicular ia pacifica Pease, 1 865 ; Ladd , 1977: 25, p l . 6 , Figs . 5— 7 ; H olocene.

Phenacovo lva br unneiterma Cate, 1 969 ; [Cer nohor sky, 1 972 : 91 , pl . 23 :P r imovu la margar i ta (Sower by , [Cernohor sky, 1 972 : 90 , pl . 23 , F ig .

Fam i ly ATLANTIDAETA t lan ta peron i i Leseu r , 1 8 1 7; Ladd, 1977 : 26 ; H o locene; [Ladd, 1 977 : 26 , pl . 6 , Figs . 1 4

Fam i ly NAT ICIDAETAmpu ll ina berauens is (Beets , Ladd, 1977 : 27 , pl . 7, Figs . 5— 7; ear ly M iocene.

Eunat icina papi l la (Gmel in , [Cernohor sky, 1972 : 1 02, pl . 27 ,F ig .

Natica bougai Sower by , 1 908 ; [Cernohor sky, 1 972 : 98 , pl . 25 , F ig .

N . qualter iana Recluz , 1844 ; Ladd , 1 977 : 30 , pl . 9 , F igs 3 — 1 0 ; as Nat icar ius marochiens is (Gmel in ,

H o locene; [Cernohor sky, 1 972 : 95 , p l . 24 , F ig .

N . onca (Rod ing , [Cernohor sky, 1 972 : 98 , pl . 25 , F ig .

N . vio lacea Sower by , 1 825; [Cernohor sky, 1972: 96 , pl . 25 , F ig .

Never i ta cf . N . pet iver iana Recluz , 1 855 ; F ig . 3H , 3 l .

Never i ta sp . ; F ig . 3F , 3G .

Po lin ices albumen (L innaeus , [Cernohor sky, 1972 : 99, p l . 26 , F ig .

P . au ran t ius (Rod ing , [Cernohor sky, 1 972 : 99, pl . 25 , F ig .

P . flem ingiana (Recluz , [Cer nohor sky, 1972 : 98 , pl . 25, F ig .

P . melanos tomus (Gmel in , [Cernohor sky,1972 : 1 01 , pl . 26 , F ig .

P . tum idus (Swainson , Ladd , 1 977 : 28 , pl . 8 , F ig . 3 ; as P . mam i l la (L innaeus ,late M iocene.

TSinum sp . A . ; Ladd, 1977: 30 , pl . 9 , Figs . 1 , 2 ; H o locene.

Fam i ly CASSIDIDAECasmar ia en

'

naceu s (L innaeus , [Abbott , 1 968 : 190 , pl . 1 4 , Figs . 8 , 1 0

C . ponderosa (Gmel in , [Abbott , 1 968 : 1 95 , pl . 1 4 , Figs . 1 ,

Cass is cor nuta (L innaeus , [Abbott , 1 968 : 47 , pl . 3 , Figs . 1

Phal ium bis u lcatum (Schuber t and Wagner , [Abbott , 1968 : 1 26 , pl . 8 , F ig .

Fam i ly CYMAT IIDAECharon ia tr i ton is (L innaeus , [Kay, 1 979: 2 1 5 , F ig . 77F].Cymat ium aquat i le (Reeve, [Kay, 1979: 220 , F ig . 77A].C . clandes tinum (Lamar ck , [Kay, 1 979 : 220 , F ig . 79A].

flaveola (R6ding , (H inton, 1 978 : 30 , F ig .

gemmatum (Reeve, [Kay, 1 979 : 222, F ig . 79C].hepat icum (L innaeus , [Cer nohor sky, 1 967 : 48 ,

p l . 4 , F ig

in termedium (Pease, [Kay, 1 979 : 220 , F ig . 77C].lotor ium (L innaeus , [Cer nohor sky, 1 967 : 48 , pl . 4 , F ig .

m ur icinum (Roding , [Kay, 1979 : 21 7, F ig . 77E].n icobar icum (Rii ding , H oubr ick , 1 974 ; [Kay, 1 979 : 21 6 , F ig . 77D].

p i leare (L innaeus , [Kay, 1 979 : 221 , F ig .

pyr um (L innaeus , [Kay, 1 979 : 2 1 8 , F ig .

r ubecu lum (L innaeus , [Cernohor sky, 1967: 52, pl . 4 , F ig .

vespaceum (Lamarck, [Kay, 1979: 223 , F ig . 79E].D is tor s io anus (L innaeus , [Kay, 1 979 : 223 , F ig . 79K].D . pus i l la Pease, 1 861 ; Ladd , 1 977 : 35 , pl . 1 1 , Figs . 1 2 , 1 3 ; H o locene.

Gyr ineum r os eum (Reeve, [Cer nohor sky, 1 972 : 1 1 7, pl . 32, F ig .

[T he occur rence of Cymat ium cf . C . caudatum (Gmel in , 1 79 1 ) ci ted by Ekdaleet al . ( 1 979) has no t been con fi rm ed].

O

O

O

O

Q

O

O

0

0

0

0

0

0

0

0

0

0

0

TSpecies no t now l iv ing at Enewetak ; ho r i zons r ecorded by Ladd ( 1 966 , 1972, 1977 , (This table

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1 16 KAY AND JOH NSON

TABLE 1 (con t ’d)

Fam i ly TRIPHORIDAE (cont ’d)N . cf . N . leucomys (H er v ier , F ig . 2l .

N . t r it icea (Pease, [Kay, 1979: 1 50 , F ig . SI E].NanOphora sp . ; F ig . 2H .

Tet raphora her vien'

(Kosuge, [Cernohor sky, 1 978 : 1 72, pl . 60 , F ig .

Tr iphora gr anu lata (Adam s and Reeve, [Cernohor sky, 1 978 : 1 69, pl . 59, F igT . taen iolata (H erv ier , [Cernohor sky, 1978 : 1 72, pl . 61 , F ig . 1 ; as Inella (Cau totr iphora)monovi tta Laser on ,

Vir io lo abbotti (Baker and Spicer , [Kay, 1979: 1 39, F ig . 50E].V . bayan i Jousseaume, 1 884 ; [Kay, 1979 : 139, F ig . 50J].V. cancel lata (H inds , Ladd , 1972: 50 , pl . 1 3 , F ig . 1 ; late M iocene; [Kay, 1979 : 1 40 , F ig . 50D].V . cf . V. cor r ugata (H inds , [Cer nohor sky, 1 978 : 173 , pl . 6 1 , F ig . 6 ; as Vir io loin terfi lata

V. incisa (Pease, Ladd , 1 972 : 50 , pl . 15, F ig . 22; H olocene.

V. in tergr anosa (H er v ier , [Cer nohor sky, 1978 : 123 , pl . 61 , F ig .

Fam i ly EPITON IIDAEEp iton ium alatum (Sower by , [Kay, 1979 : 153 , F ig . 54C].

au r i tum (Sower by , F ig . SE.

i r r egu lar is (Sower by , [Cernohor sky, 1 972 : 197, pl . 56 , F ig .

paumotens is (Pease, [Kay, 1 979 : 1 56 , F ig . 54A].

perp lexum (Deshayes , Ladd, 1972 : 51 , p l . 13 , F ig . 3 ; H o locene; [Kay, 1 979 : 156 , F ig .

symmet r ica (Pease,

revo lutum (H edley , [Kay, 1979 : 1 56 , F ig . 54F].cf . E . ves talis (H inds , [Cernohor sky, 1978 : 168 , pl . 59, F ig .

umbi l icatum (Pease, [Kay, 1979 : 157, F ig . S4K].Opalic bicar inata (Sower by , [Cernohor sky, 1 978 : 168 , pl . 59 , F ig . 9; as Nodiscala].mmmmmmmm

Fam i ly JANTH IN IDAEJan th ina g lobosa Swainson , 1822; [Kay, 1 979 : 158 , F ig . SSA, D].J. jan th ina (L innaeus , [Kay, 1 979: 158 , F ig .

Order NEOGASTROPODAFam i ly MUR ICIDAEAspella producto (Pease, [Kay, 1 979 : 235, Figs . 83C, D].A tt i l iosa caledon icu s (Jou sseaume, [Vokes and D ’

Att i lio, 1982 : 70 , F ig

Chicoreu s br u nneu s (L ink , [Cer nohor sky, 1967 : 120 , pl . 25, F ig .

C . m icrophy l lus (Lamarck , [Radwin and D’

Att i lio , 1 976 : 39, pl . 4 , F ig .

C . ramosus (L innaeus , [Cernohor sky, 1967: 1 22 , pl . 25 , F ig .

Favar t ia guamens is Emer son and D’

Att i lio , 1979 ; [Emer son and D’

Att i l io, 1 979 ,4 , Figs . 1 1 ,

H omalocan tha anatom ica (Perr y , [Radwi n and D’

Att i lio, 1976 , pl . 9, F ig .

March ia bip innata (Reeve, [Radwi n and D’

Att i lio , 1976 : pl . 9 , F ig .

M . laqueata (Sower by , [Radwi n and D’

Att i lio , 1 976 : 58 , pl . 9, F ig .

M . mar t inetana (R6ding , [Radwin and D’

Att i lio , 1 976 : pl . 27, F ig .

M . tr i p tera (80m , [Radw in and D’

Att i lio , 1 976 : pl . 9 , F ig .

Naguet ia tr igon u la (Lamar ck , [Radw in and D’

Att i lio, 1 976 , pl . 15, F ig .

N . tr iquetra (8 0m , [Radw in and D’

At t i lio, 1976 , pl . 15, F ig .

Phyllocoma convo luta (Broder ip , [Cernohor sky, 1967 , pl . 24, F ig .

Phyllonotus lacin iatus (Sower by , [Radwi n and D’

Att i lio, 1976 , pl . 6 , F ig .

TPterynotus sp . ; Ladd, 1977 : 39, p l . 1 4 , F ig . 3 ; H olocene.

Fam i ly TH AIDIDAECr on ia fiscella (Gmel in , Ladd, 1 977 : 40 , pl . 1 4 , F ig . 8 ; late Miocene; [Cernohor sky, 1972 : 1 28 , pl . 36 , F ig 8 ;

as Mor u la margar i t icola (B roder ip ,

D r upa ar achno ides (Lamarck , Ber nstein , 1 974 ; Lev iten , 1974 ; Kohn and Lev iten , 1976 ; Kohn , 1980a;

[Emer son and Cer nohor sky, 1 973 : pl . 2, F ig . 6 ; as D rupa r icinus r icinus].clathrata (Lamar ck, Ber nstein , 1 974 ; Emer son and Cernohor sky, 1973 : 33 , pl . 2 , Figs . 16— 1 8 .

elegans (Broder ip and Sower by , Ber n stein , 1974 ; Emer son and Cer nohor sky, 1973 : 25, pl . 2, F ig . 12 .

gr os su lar ia Rod ing ,1 798 ; Emer son and Cer nohor sky, 1973 : 37 , pl . 2 , Figs . 23 ,

24 ; Kohn , 1 980a.

morum Rad ing ,1 798 ; Emer son and Cer nohor sky, 1973 : 1 7 , pl . 2, Figs . 1 — 3 ; Ber nstei n , 1974 ; Kohn and Lev iten ,

1976 ; Kohn , 1 980a; Lev iten and Kohn , 1980 .

r icina (L innaeus , Emer son and Cer nohor sky, 1973 : 22,pl . 2 , Figs . Ber nstein , 1974 , Lev iten , 1974 ;

Kohn and Lev i ten , 1 976 ; Kohn 1 980a; Levi ten and Kohn , 1 980 .

D

0

0

0

0

TSpecies not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd ( 1 966 , 1 972, 1 977 , (This table cont inued on next pa

MOLLUSCA 1 1 7

TABLE 1 (con t'd)

Fam i ly TH AIDIDAE (cont ’d)D . r ubus idaeu s Roding , 1 798 ; Emer son and Cernohor sky, 1 973 : 29, pl . 2, Figs . 1 3 — 1 5 .

D r upe lla elata (B lainv i l le, Kohn , l 98oa; as D r upella com us (Rod ing , [Kay, 1979 : 244 , F ig . 87A].D . fenes t r ata (B lainv i l le, [Cer nohor sky, 1 978 : 68 , pl . 1 9 , F ig .

D . [r agum (B lainv i l le, [Moyer et al . , 1 982 : F ig .

D . r ugosa (80 m , Ladd , 1 982 : 44 , pl . 1 1 , Figs . 1 2 , 1 3 ; as D . concat enate (Lamarck , H o locene ;[Cer nohor sky, 1 972 : 1 26 , pl . 35 , F ig .

D r upe lla sp . , [Kay, 1979 : 244 , F ig . 878 ; as D r upella ochros toma (B lainv i lle,

Macu lo tr i ton ex im ius (Reeve, [Cer nohor sky, 1 978 : 71 , pl . 20 , F ig .

M . sculpt i le (Reeve, [Cer nohor sky, 1972 : 1 30 , p l . 36 , F ig .

M . ser r iale (Deshayes , Lev i ten , 1974 ; as M . br acteatu s (H inds , Kohn and Lev iten , 1 976 ;

Kohn , 1 980a; [Cer nohor sky, 1 972 : 1 29 , pl . 36 , F ig .

Mor u la anaxeres (Kiener , [Ce r nohor sky, 1 972 : 1 26 , pl . 35 , F ig .

M . fun icu lata (Reeve, Ladd , 1 977 : 40 ; as D rupa (D r upella) cf . D . mon i l ifer a (Pease, H o locene ; [Kay,

1 979 ; 247 , F ig . 83l].M . gr anulate (Duclos , Ber n s tein , 1 974 ; Levi ten , 1 974 ; Kohn and Levi ten , 1976 ; Ladd , 1977 : 40 ,

pl . 1 4F ig . 7; H o locene; Kohn , 1 980a; as Moru la fi scel la; Lev iten and Kohn , 1 980 ; [Cer nohor sky, 1 972 : 1 27 ,

pl . 36 , F ig .

M . nod icos tata (Pease, [Cernohor sky, 1 972 : 1 27, pl . 36 , F ig .

M . sp inosa (H . and A . Adams , [Cernohor sky, 1 978 : 1 28 , pl . 36 , F ig .

M . squamos'a (Pease, Figs . 3J, 3K .

M . uva (R'

od ing ,Ber ns tein , 1 974 ; Lev iten , 1 974 ; Kohn and Lev iten , 1 976 ; Kohn , 1 980a ; Lev iten and Kohn

1 980 ; [Kay, 1 979 : 249, F ig . 87F].Mu r icodr upa fun icu lus (Wood , Kohn and Lev i ten , 1 976 ; [Kay, 1 979 : 238 , F ig . S3I].Nas sa ser ta (Brugu ier e, [Cer nohor sky, 1 967 : 1 35 , pl . 29 , F ig .

Thais acu leata (Deshayes , [Cernohor sky, 1 967 : 1 30 , pl . 28 , F ig . 1 72 ; as Thais h ippocas tanum (L innaeus ,T . arm iger a (L ink , Menge, 1 973 ; Ber ns tein , 1 974 ; Lev iten and Kohn , 1 980 ; [Kay, 1 979 :

251 , F ig . 88G].T . in termedia (Kiener , Ber n stein , 1 974 ; [Kay, 1 979 : 251 , F ig . 88H ].T . tuberosa (R6ding , Lev iten , 1 974 ; Kohn and Lev i ten, 1 976 ; Lev iten and Kohn , 1 980 ; Kohn , l 98oa;

[Cer nohor sky, 1 967 ; 130 , pl . 28 , F ig .

Vex i l la lineata A . Adam s , 1 853 ; [Kay, 1 979 : 252, F ig . 88E].V . vex i l lum (Gmel in , [Kay, 1979 : 252, F ig . 8BD].

[T he record of Purpura aper ta (B lainv i l le, 1 832) ci ted by Kay ( 1 979) has not been confi rmed].Fam i ly CORALL IOPH ILIDAECor all iophi la cos tu lar is (Lamar ck , [Cer nohor sky, 1978 : 73 , pl . 21 , F ig .

C . d'or bignyana (Pet i t , [Kay, 1 979 : 255 , F ig . 90A].

C . erosa (Roding , [Cer nohor sky, 1 972 : 1 3 1 , pl . 37, F ig .

TC . macnei l i Ladd , 1977 ; Ladd, 1 977 : 4 1 — 42 ,pl . 14 ,F igs . 1 2, 1 3 ; late Miocene.

C . v io lacea (K iener , [Kay,1 979 : 256 , F ig . 90C].

TCor all iophi la (Corall ioph i la) sp . ; Ladd , 1 977 : 41 , pl . 1 4 , F ig . 1 1 ; H o locene.

TCorall ioph i la (F usom ur ex) sp . A . ; Ladd , 1 977 : 42 , pl . 1 5 , F ig . 3 ; late Miocene.

TCor all ioph i la (F usom u rex) sp . 8 ; Ladd , 1 977 : 42, pl . 1 5 , F i g . 4 ; ear ly M iocene.

Lat iax is inflata (Dunker , Ladd , 1 977 : 43 , pl . 1 5 , F ig . 5; late M iocene .

Quoyu la m onodon ta (B lainv i l le , Ladd , 1 977 : 43 — 44 , pl . 1 5 , F ig . 7 ; H o locene; [Kay, 1979 : 258 , F ig . 906 ;

as Quoyu la madrepo rar um Sower by].Rapa rapa (L innaeus , [Cer nohor sky, 1 972 : 130 , pl . 37 , F ig .

Fam i ly BUCCIN IDAECaduci/er t runcata (H inds , [Cer nohor sky, 1972 : 1 41 , pl . 30 , F ig .

Can tharus ex im iu s Reeve, 1 846 ; [Cer nohor sky, 1 978 : 76 , pl . 22, F ig .

C . far inos us (Gould , [Kay, 1 979 : 261 , F ig . 9ZO].C . pu lcher (Reeve, [Cer nohor sky, 1 972 : 1 42, pl . 38 , F ig .

C . undosus (L innaeus , Lev i ten , 1 974 ; Kohn , 1 980a; [Cer nohor sky, 1972 : 1 41 , pl . 38 , F ig .

C . wagnen’

(Anton , [Cer nohor sky, 1 972 : 1 42 , pl . 39 , F ig .

Cl ivipo ll ia f r agar ia (Wood , [Kay, 1 979 : 263 , F ig .

Eng ina egr eg ia (Reeve , [Cer nohor sky,1 972 : 1 43 , pl . 39 , F ig.

E . l ineata (Reeve, [Cernohor sky, 1 972 : 1 43 , pl . 39 , F ig .

E . mendicar ia (L innaeus , [Cer nohor sky, 1 972 : 144 ,pl . 38 , F ig .

TSpecies not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd (1 966 , 1 972 , 1 977 , (Th is table con t inued on next page. )

1 18 KAY AND JOH NSON

TABLE 1 (con t ’d)

Fam i ly BUCCIN IDAE (cont ’d)E . zonal is (Lamar ck , [Sal isbur y , 1 983a:P rodot ia ignea (Gmel in , [Kay, 1 979: 264 , F ig.

P . ios toma (Gr ay , [Kay, 1 979 : 264 , F ig . 92J].

Fam i ly COLUMBELL IDAEAesop u s spicu lum (Duclos , [Cer nohor sky, 1 972 : 1 38 , pl . 41 , F ig .

Anach is m iser (Sower by, [Kay, 1979 : 266 , F ig . 94F].

Co lumbel la cf. C . molecu l ina Duclos , 1 835 ; F ig . 3N .

TC . pardal ina Lamarck , 1 822 ; Ladd, 1 977 : 44 , pl . 15, F ig . 1 1 ; late M iocene.

Eup l ica am i r an t ium (Sm i th , F ig . 3M .

deshayes i i (Cr osse, [Cernohor sky, 1 972: 134 , pl . 40 , F ig

cf . E . l iocyma Pi lsbry, 1 904 ; F ig . 3Q .

cf . E . m inuta Gould , 1 860 ; F ig . 3R .

obes u la H erv ier , 1 899 ; F ig . 3W .

tu r tu r ina (Lamarck , Ladd , 1 977 : 44 , pl . 15, Figs 1 2 , 13 ; Quater nar y .var ian s (Sowerby , Ber ns tein , 1974 ; Ladd, 1977 : 45 , pl . 15 , F ig . 14; H o locene.

TLavesopus en iwetokens is Ladd, 1 977; Ladd, 1977 : 45, p l . 1 6 , F ig . 1 ; H o locene.

M i t rella fu s i/orm is (Pease, [Kay, 1979 : 268 , F ig . 94H — I].M . cf . M . albina (Kiener , F ig . 3P .

M . ligu la (Duclos , [Cer nohor sky, 1 972 : 135 , pl . 41 , F ig .

TM . sagi tta (Gasko in , Ladd , 1977 : 47, pl . 16 , F ig . 1 1 ; ear ly Miocene to H o locene.

Pyrene flava (Brugu iere, [Cer nohor sky, 1972 : 132, pl . 40 ,F ig .

cf . P . l ivescens (Reeve, F ig . 30 .

obtusa (Sower by, Ladd, 1977 : 47, pl . 1 6 , F ig . 1 0 ; H o locene; F ig . 3L .

oce l lata (L ink , [Cer nohor sky, 1 972 : 1 32, p l . 40 , F ig .

punctata (B rugu ier e, F ig . 3V .

Sp lendidu la (Sower by , [H abe, 1 964 : 89, pl . 28 , F ig .

tes tudinar ia (L ink , [Cernohor sky, 1972 : 133 , pl . 40 , F ig .

Sem inella cf . 8 . dau tzenberg i (H er v ier , F ig . 3T .

8 . divar icata (P i lsbry, F ig . 35 .

TS. sm i thi (Angas , Ladd, 1 977 : 47, pl . 1 6 , F ig . 9 as Anachis ; late M iocene.

S. s ubphi lodicia (H erv ier , F ig . 3U .

Zafr a trog lodytes (Souver bie, [Cern ohor sky, 1972 : 1 38 , pl . 41 , F ig .

Zafr ona pus i l la (Pease, Ladd, 1 977 : 45 , pl . 1 6 , F ig . 2; as Zafr ona l ifuana (H er v ier , H o locene.

Fam i ly COLUBRARIIDAECo lubrar ia an t iquata (H inds , [Cer nohor sky, 1 972 : 1 21 , pl . 37, F ig .

C . cf . C . cum ingi (Dohr n , [H i rase, 1 95 1 : pl . 1 05 , F ig .

C . mur icata (L ightfoot , [Kay, 1979 : 271 , F ig . 92L].C . n i t idu la (Sower by , [Cer nohor sky, 1 972 : 1 21 , pl . 37, F ig .

C . tor tuosa (Reeve, [Kay, 1979: 272, F ig . 92M].Fam i ly MELONGEN IDAETPugi l ina swar tz i Ladd , 1 977; Ladd, 1977 : 51 , pl . 1 7 ,

Figs . 6 , 8 ; ear ly M iocene.

Fam i ly NASSARIIDAENassar ius comp tus (A . Adams , [Cer nohor sky, 1 978 : 87,

pl . 27, F ig .

N . concinnus (Powys , [Cer nohor sky, 1 978 : 88 ,pl . 27, F ig .

TN . en iwe tokens is Ladd , 1 977 ; Ladd , 1 977 : 52 , pl . 1 7,Figs . 1 4 , 15 ; late M iocene.

N . gaudiosu s (H inds , Kohn , 1 980a; [Cer nohor sky, 1978 : 86 , pl . 27, F ig .

N . g lans (L innaeus , [Cer nohor sky, 1 972 : 1 51 , pl . 45, F ig .

N . gr an ijer us (Kiener , [Cer nohor sky, 1 972 : 146 ,pl . 42 , F ig .

TN . mar shallens is Ladd , 1 977; Ladd , 1 977 : 55, pl . 1 8 ,Figs . 6 , 7; M iocene.

N . pap i llos u s (L innaeus , [Kay, 1 979 : 275 , F ig . 95A].N . pauper us (Gou ld , [Cer nohor sky, 1 972 : 1 52, pl . 44 , F ig .

TN . t iar u la (Kiener , Ladd , 1 977 : 52 , pl . 1 7 , F ig . 9 ; pl . 21 , F ig . 9 ; H o locene .

Fam i ly FASCIOLAR I IDAEDo l icholat i r us lancea (Gmel in , [Cer nohor sky, 1 972 : 1 61 , pl . 46 , F ig .

Fascio lar ia f i lamen tosa (Rod ing , [Cer nohor sky, 1972 : 1 53 ,pl . 45, F ig .

F us in us undatu s (Gmel in , [Cer nohor sky, 1 972 : 1 62,pl . 48 , F ig .

Lat i rus crat icu latus (L innaeus , [Cer nohor sky, 1972 : 156 ,pl . 46 , F ig .

L . macu latus (Reeve, [Cer nohor sky, 1 978 : 92 ,pl . 29, F ig .

mmmmmm

TSpec ies not now l iv ing at Enewetak ; hor izons recorded by Ladd ( 1 966 , 1 972 , 1977 , (This table con t inued on next pa

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120 KAY AND JOHNSON

TABLE 1 (con t’d)

Fam i ly MITR IDAE (cont ’d)M . cucumer ina Lamarck , 1 8 1 1 ; Bern s tein , 1 974 ; Kohn and Lev i ten , 1 976 ; Kohn , 1 980a; [Cer nohor sky, 1976 :

400 , pl . 256 , Figs . 13

M . eden tu la Swainson , 1 823 ; [Cer nohor sky, 1 976 : pl . 257, Figs . 35,

M . erem itar um (R6ding , 1798 ; [Cer nohor sky, 1 976 : 324 , pl . 255, Figs . 7,

M . fas t igium Reeve, 1 845 ; Ber ns tein , 1 974 ; as M itra brunnea Pease, 1 868 ; [Cernohor sky, 1976 : 494 , pl . 258 ,

Figs . 28 ,

M . fer r ugi nea Lamar ck , 18 1 1 ; [Cer nohor sky, 1 976 : 397, pl . 256 , Figs . 9,

M . f r aga Quoy and Gaimard, 1833 ; Ladd, 1 977 : 60 , p l . 1 9 , F ig . 9 ; H o locene; [Cernohor sky, 1 976 : 404 , pl . 256 ,

Figs . 1 8 ,

M . fu lvescen s B roder ip , 1 836 ; [Cer nohor sky, 1 976 : 43 1 , p l . 257, Figs . 10 ,

M . imper ialis R6d ing ,1 798 ; Cer nohor sky, 1976 : 321 , pl . 255 , Figs . 1 5 , 16 .

M . incomp ta (So lander in L ight foot , [Cernohor sky, 1 976 : 322, pl . 253 , F ig .

M . li tterato Lamarck , 1 8 1 1 ; Bern stein , 1974 ; Levi ten , 1 974 ; Kohn and Lev i ten , 1 976 ; as St r igatella; Kohn , 1 980a;

as Str igatella ; Lev i ten and Kohn , 1980 ; [Cern ohor sky, 1 976 : 482, p l . 258 , Figs . 9

luctuosa A . Adam s , 1 853 ; [Cernohor sky, 1 976 : 443 , pl . 257, Figs . 23 ,

m it r a (L innaeus , [Cer nohor sky, 1976 : 307 , pl . 253 , F ig .

n ivea (B roder ip , [Cer nohor sky, 1976 : 335 , pl .popo l is (L innaeus , [Cer nohosky, 1 976 : 308 , pl . 253 , F ig .

pauper cula (L innaeus , Lev iten and Kohn , 1 980 ; [Cernohor sky, 1 976 : 477, pl . 258 , Figs . 1

pecu l iar is Reeve, 1 845 ; [Cer nohor sky, 1 976 : 424 , pl .pet r osa Sower by , 1 874 ; [Cer nohor sky, 1976 : pl . 292 , F ig . 13 ; albino form of M . us tu latapyram is (Wood, [Cer nohor sky, 1 976 : 424 , pl .r ubr i t incta Reeve, 1844 ; [Cer nohor sky, 1 976 : 399 , pl . 256 , Figs . 1 1 ,

scu tu lata (Gmel in , [Cer nohor sky, 1 976 : 488 , pl . 258 , Figs . 1 7

s t ict ica (L ink, [Cer nohor sky, 1 976 : 309 , pl . 253 , Figs . 3 ,

tabanu la Lamar ck , 1 8 1 1 ; [Cernohor sky, 1 976 : 408 , pl . 256 , F ig .

t icaon ica Reeve, 1 844 ; [Cer nohor sky, 1 976 : 429, p l . 257 , Figs . 4 ,

turgida Reeve, 1845 ; Ladd, 1 977 : 60 , pl . 1 9 , Figs . 1 0 , 1 1 ; late Miocene.

typha Reeve, 1845 ; [Cer nohor sky, 1976 : 499, p l .us tu lata Reeve, 1 844 ; [Cer nohor sky, 1 976 : 337 ,

pl .vu ltuoso Reeve, 1845; [Cernohor sky, 1976 : 442, p l . 391 , F ig .

Neocanci lla c lathrus (Gmel in , [Cernohor sky, 1 967 : 154, pl . 35 , F ig .

N . pap i lio (L ink , [Cer nohor sky, 1967 : 154, pl . 32, F ig .

Pter ygia crenu lata (Gmel in , [Cernohor sky, 1 967 : 1 6 1 , pl . 38 , F ig .

P . dacty lus (L innaeus , [Cer nohor sky, 1 967 : 1 62 , pl . 38 , F ig .

P . fenes t rata (Lamarck , [Cernohor sky, 1 967 : 162 , pl . 38 , F ig .

P . n ucea (Gmel in , [Cer nohor sky, 1 967 : 1 62 , pl . 38 , F ig .

Scabr icola cas ta (Gmel in , [Cernohor sky, 1 967 : 1 60 , p l . 37, F ig .

S. coeru lea (Reeve, [Cernohor sky, 1972 : 1 73 , pl . 51 , F ig .

S. desetangs i i (K iener , [Cer nohor sky, 1 967 : 16 1 , pl . 37 , F ig . 251 ; as Swain son ia suffecta (Dau tzenberg and

Bouge,

S. var iegata (Gmel in , [Cer nohor sky, 1 978 : 10 1 , pl . 32 , F ig .

Subcanc i lla in ter l ir ata (Reeve, [Cernoho r sky, 1978 : 1 05 , pl . 34 , F ig .

S. ver r ucosa (Reeve, [Cer nohor sky, 1 967 : 158 , p l . 36 , F ig . 249 ; as Canci l la].

3

3

3

3

3

3

3

3

3

3

3

3

3

3

3

3

3

Fam i ly COSTELLARIIDAEVex i llum acup ictum (Reeve, [Cer nohor sky, 1967 : 1 64 ,

pl . 38 , F ig .

V . amabi lis (Reeve, [Cernohor sky, 1967 : 90 , pl . 44 , F ig .

TV . app rox imat um (Pease, Ladd , 1977 : 66 , pl . 20 , F ig . 1 9 ; ear ly M iocene.

V . bem hardina (Rod ing , [Cernohor sky, 1 967 : 1 82, pl . 44 , F ig .

V . cadaverosum (Reeve, [Cer nohor sky, 1 967 : 166 , pl . 39 , F ig .

V . cancellar io ides (Anton , Ber nstein , 1 974 ; Lev iten , 1 974 ; Kohn and Lev iten , 1976 ; Lev iten and Kohn ,

1 980 ; as Pus ia nodosa (Swainson , Kohn , 1 980a; as Pus ia cancellar io ides (Anton,

[Kay, 1 979 : 325 , F ig .

catenatum (B roder ip , [Cer nohor sky, 1 978 : 1 1 6 , pl . 40 , F ig .

cem ohor skyi Ladd , 1977 ; Ladd, 1977 : 63 , pl . 20 , Figs . 6 , 7 ; late M iocene.

co r bicu lum (Sower by , [Cernohor sky, 1 978 : 1 12, pl . 37 , F ig .

cor onatum (Helbl ing , [Cer nohor sky, 1 967 : 1 66 ,pl . 39 , F ig .

cos tatum (Gmel in , [Cer nohor sky, 1967 : 167 , pl . 39 , F ig .

TSpec ies no t now liv ing at Enewetak ; ho r i zons r ecorded by Ladd ( 1 966 , 1 972 , 1 977 , (This table con t inued on next Pa

MOLLUSCA 12 1

TABLE 1 (con t'd)

Fam i ly COSTELLARIIDAE (cont ’d)cr ocatum (Lamarck , [Cer nohor sky, 1 967 : 1 67 , pl . 39 , F ig .

deshayes i (Reeve, Ladd , 1 977 : 63 , pl . 20 , F ig . 8 ; late M iocene.

echinatum (A . Adams , [Cernohor sky, 1 967 : 1 68 , pl . 40 , F ig .

em i l iae (Gar r ett , [Kay, 1 979 : 327, pl . 1 09F].

exasperatum (Gmel in , [Cer nohor sky, 1 967: 1 68 , pl . 40 , F ig .

exqu is itum (Gar r ett , Ladd , 1 977 : 66 , pl . 21 , F ig . 1 ; H o locene.

fes tum (Reeve,Ladd , 1 977 : 64 , pl . 20 , F ig . 1 0 ; M iocene.

fil is tr iatum (Sower by , [Cernohor sky, 1978 : 1 1 0 , pl . 36 , F ig .

for t ip l icatum (Pease, [Cer nohor sky, 1 972 : 1 76 , pl . 52 , F ig .

cf . V . gembacan um (Mar tin , Ladd , 1 977 : 65 , pl . 20 , F ig . 1 4 ; Miocene.

goubin i (H er v ier , [Cernohor sky, 198 1 : 1 0 1 , pl . 3 , F ig .

gr anosum (Gmel in , [Cer nohor sky, 1 967 : 1 70 , pl . 40 , F ig .

gr uner i (Reeve, [Cer noho r sky 1967 : 1 71 , pl . 41 , F ig .

hum i lis (H er v ier , [Cer nohor sky, 1 98 1 : 96 , p l . 1 , Fi g .

infaus tum (Reeve, [Cernohor sky, 1 967 : 1 83 , pl . 45, F ig .

in ter r up tum (Anton , [Cernohor sky, 1 978 : 1 12, p l . 37, F ig .

in ter s tn‘

atum (Sower by , [Kay, 1 979 : 320 , F ig . 1 08P].lau tum (Reeve, [Cernohor sky, 1 972 : 1 77, pl . 52 , F ig .

leucozon ias (Deshayes , Ladd , 1 977 : 64 , pl . 29 , F ig . 1 1 ; H o locene.

lucu len tum (Reeve, [Cer nohor sky, 1 967 : 1 72 , pl . 4 1 , F ig .

m ica (Reeve, [Cer nohor sky, 1 967 : 1 72 , pl . 41 , F ig .

m ichau i (Crosse and Fischer , [Cernohor sky, 1967 : 1 71 , pl . 41 , F ig 288 ; as V. in ter taen iatum

(Sower by ,m icrozon ias (Lamar ck , [Cer nohor sky, 1 967 : 1 83 , pl . 45 , F ig .

m i llecos tatum (B roder ip , [Cer noho r sky, 1 967 : pl . 41 , F ig . 8 21 ; as Pus ia adam son i (Reeve,modes tum (Reeve, [Kay, 1979 : 322 , F ig . 1OSH ].moeller i (Kuster , [Cer nohor sky, 1 972 : 1 78 , pl . 51 , F ig .

obel is cus (Reeve, [Cer nohor sky, 1 967 : 1 74 , pl . 4 1 , F ig .

obtus isp inosum (Sower by , [Cer nohor sky, 1 967 : 1 74 , pl . 42, F ig .

pacificum (Reeve, Ekdale et al . 1 979 : as V . speciosum (Reeve, [Cer nohor sky, 1 967 : 1 74,

pl . 42, F ig .

pagodu la (H er v ier , [Cernohor sky, 1 981 : 95 , pl . 1 , F ig .

pat ri ar chal is (Gmel in , [Cernohor sky, 1 967 : 1 84 , p l . 45,F ig .

po lygonum (Gmel in , [Cernohor sky, 1 978 : 1 08 , pl . 36 , F ig .

roseum (Broder i p , Ladd , 1 977 : 65 , pl . 20 , F ig . 1 5 ;'Pleistocene.

rubr um (Br oder ip , [Kay, 1 979 : 33 1 , F ig . I 1 0E].

sangu isugum (L innaeus , [Cernohor sky, 1 967 : 1 76 , pl . 43 , F ig .

speciosum (Reeve,Ekdale et al . , 1 979 ; [Cer nohor sky, 1 972 : 1 77 , pl . 52 , F ig .

s uav is (Sou ver bie, [Cer nohor sky, 1 978 : 1 1 6 , pl . 40 , F ig .

tuberosum (Reeve, [Cer nohor sky, 1 967: 1 86 , pl . 45 , F ig .

tur ben (Reeve, [Cer nohor sky, 1 978 : 1 1 7 , pl . 39 , F ig .

tu r r iger um (Reeve, [Cer nohor sky, 1 972 : 1 75 , pl . 52 , F ig .

tus um (Reeve, [Cernoho r sky, 1 972 : 1 78 , p l . 52 , F ig .

un ifasc ial is (Lamarck , [Cer nohor sky, 1 967 : 1 80 , pl . 44 , F ig .

vu lpecu la (L innaeus , [Cer nohor sky, 1 967 : 1 80 , pl . 44 , F ig .

zelotyp um (Reeve, [Cer nohor sky, 1 972 : 1 77, pl . 52 , F ig .

Fam i ly TURRIDAEAnaci thar a nan is ca (H er v ier , F i g . 4I.

A . notabi lis (E . A . Sm ith , F ig . 4J.

A . s t r icta H edley , 1 922; [F ig 4H ].Car inapex m inut is s ima (Gar r ett , [Kay, 1 979: 344 , F ig .

C lavus bi l ineatus (Reeve, F ig . 4L .

can icu lar is (Ri iding , [Cern ohor sky, 1 972 : 1 85 , pl . 53 , F ig .

laetus (H inds , [Cer nohor sky, 1 978 : 1 53 ,pl . 55 , F ig .

lam bert i (Mont r ouz ier , [Cer nohor sky, 1 978 : 153 , pl . 55 , F ig .

nodulosa (Pease, [Kay, 1 965 : 80 , pl . 1 4 , Figs . 9 ,

Opalus (Reeve, [Cer noho r sky, 1 978 : 1 52 , pl . 54 , F ig .

p ica (Reeve, [Cernohor sky, 1 978 : 152 , p l . 54 , F ig .

T

0

0

0

0

0

0

TSpecies not now l iv ing at Enewetak ; hor izons r ecorded by Ladd (1 966 , 1 972 , 1 977 , (This table con t inued on next page. )

122 KAY AND JOH NSON

TABLE 1 (con t ’d)

Fam i ly TURRIDAE (cont ’d)C . un iz onal is (Lamar ck , [Cernohor sky, 1 972 : 1 86 , pl . 53 , F ig .

C . viduus (Reeve, [Cer nohor sky, 1 978 : 1 59 , pl . 54 , F ig .

C lavus sp . A . ; F ig . 4M .

C lavus sp . 8 ; F ig . 4K .

Daphnella aureo la (Reeve, [Cer nohor sky, 1978 : 159 , pl . 57 , F ig .

D . flammea (H inds , [Cernohor sky, 1 978 : 158 , pl . 56 , F ig .

D . cf . D . gr anata H edley , 1 922; [H edley , 1 922 : 328 , p l . 58 , F ig .

D . olyra (Reeve, [Cernohor sky, 1978 : 1 59, p l . 57 , F ig .

D . r is soides (Reeve, [Powel l , 1966 : 1 24 , pl . 19,F ig .

E tr ema alphon s ianum (H er v ier , [H edley , 1922 : pl . 47, F ig .

E . crass i labr um (Reeve, [Cer nohor sky, 1978 : 1 56 , p l . 56 , F ig .

E . cf . E . polydesma H edley , 1 922 ; [H edley , 1 922 : 282, pl . 47 , F ig .

E . s calar ina (Deshayes , [Deshayes , 1 863 : 1 09 , pl . 1 2, Figs . 1 2

E . t r igonos tom um (H er v ier , F ig . 4R .

Euci thara angios toma (Pease, [Kay, 1 979 : 352, F ig .

E . celebens is (H inds , [Cer nohor sky, 1 972 : 1 89 , p l . 54 , F ig .

E . ci thara (Gould , F ig . 4N .

E . debi lis (Pease, F ig . 4P .

E . mar shallens is (Ladd , Ladd, 1982 : 68 , pl . 22, Figs . 1 5— 1 9 ; M iocene .

E . ret icu lata (Reeve, [Cernohor sky, 1972 : 1 86 , pl . 54 , F ig .

E . cf . E . s tr icta H edley , 1 922; [H edley , 1 922 : 304 , pl . 51 , F ig .

E . s tr ombo ides (Reeve, [Cernohor sky, 1 972 : 1 86 , p l . 54 , F ig .

E . vex i l lum (Reeve, [H i r ase, 1 951 : pl . 1 15, F ig .

Eucithara sp . ; F ig . 4P .

Eucyclotoma t r icar inata (Kiener , [Cernohor sky, 1 978 : 1 60 ,pl . 57, F ig .

Ir edalea pygmaea (Dunker , [Cernohor sky, 1 972 : 1 86 , p l . 55 , F ig .

Kerm ia cf . K . bifasc iata (Pease, [Kay, 1979 : 360 , F ig . 1 1 8F].K . can is t ra (H edley , [Cer nohor sky, 1 978 : 1 62, pl . 57, F ig .

K . cf . K . concinna (H edley , F ig . 45 .

K . daedalea (Gar r ett , [Kay, 1 979 : 36 1 , F ig . 1 18I].K . edychr oa (H er v ier , F ig . 4U .

K . cf . K . gr an icos tata (Reeve, F ig . 4Q .

K . macu losa (Pease, [Cernohor sky, 1978 : 1 61 , pl . 57 , F ig . 1 1 ; as Kerm io bam ardi Br azier ,L ienardia ap icu lata (Mont r ouz ier , [Kay, 1 979 : 353 , F ig 1 1 6L].L . cr ass icos tata (Pease, [Kay, 1 979: 354 , F ig . 1 I 6E].L . cf . L . [alsar ia H edley , 1 922; [Cer nohor sky, 1978 : 1 56 ,

pl . 56 , F ig .

L . goubin i (H er v ier , F ig . 4V .

L . idiomorpha (H er v ier , F ig . 4W .

L . r os eot incta (Mont r ouz ier , [Cer nohor sky, 1 978 : 156 , pl . 56 , F ig .

L . r ubida (H inds , [Cer nohor sky, 1 972 : 189,pl . 54 , F ig .

Lophio toma acuta (Per r y , [Powel l , 1 964 : 303 ,pl . 1 80 ,

Figs . 1 ,

L . albina (Lamar ck , [Powel l , 1 964 : 306 ,pl . 1 80 ,

Figs .

TL . en iwetoken s is Ladd , 1982; Ladd , 1 982 : 62 , pl . 19, Figs . 2 , 3 ; ear ly Miocene.

Lovellona at r amen tosa (Reeve, [Kay, 1979: 348 , F ig . 1 15A].Macteo la in ter rupta (Reeve, [Cer nohor sky, 1978 : 1 57 , pl . 56 , F ig .

M . seges ta (Chenu , [Kay, 1 979 : 355, F ig . 1 16M].Macteo la sp . ; not fi gur ed .

M icr odaphne mor r i son i Rehder , 1 980 ; [Rehder , 1 980 : 88 , pl . 1 1 , Figs : 5 ,

M itr o lum na s tephen i (Melvi lle and Standen , [Cernohor sky, 1 978 : 1 63 , p l . 58 , F ig .

M . metu la (H inds , [Cer nohor sky, 1978 : 1 62, p l . 58 , F ig . 2; as M itromorpha lachryma (Reeve,

Nodotoma har ucoa (Bar tsch , [Johnson , 1 964 : pl . 5 , F ig . 4 ; as Clathu rella lacunosa Gould ,Phi lber t ia nexo (Reeve, [Cernohor sky, 1 978 : 1 62 ,

pl . 58 , F ig .

P . cf . P . cras s i labr um (Reeve, F ig . 4T .

P . r ufo l i r ata (H er v ier , [H er v ier , 1 896 : 99,pl . 2 , F ig .

P . t incta (Reeve, [Cer nhor sky, 1978 : 1 62, pl . 57 , F ig .

Tr i tonotu r r is cum ingi i (Powys , [Kay, 1 979 : 363 , F ig . I 1 8Q].T . r obi llardi (H . Adam s , [Powel l , 1 966 : pl . 20 , F ig .

Tur r idr upa albofasciata (Sm ith , [Powell , 1 967 : 41 6 ,pl . 303 ,

Figs . 3 ,

TSpecies no t now l iv ing at Enewetak ; hor i zon s r ecorded by Ladd (1966 , 1 972 , 1977, (This table con t inued on next pas

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124 KAY AND JOHNSON

TABLE 1 (con t ’d)

Fam i ly CON IDAE (cont ’d)C .

0

0

0

0

0

0

0

0

0

0

0

O

O

O

O

O

O

O

O

O

O

m i l iar is H wass in Brugui er e, 1 792; Lev i ten , 1974 ; Kohn and Lev i ten , 1976 ; Kohn , 1980a; Lev iten and Kohn ,

1 980 ; [Cernohor sky, 1 978 : 1 32 , pl . 44 , F ig

morelet i Cr os se, 1 858 ; [Kay, 1 979 : 375 , F ig . 1 22F].mus icus H wass in Brugu ier e, 1792 ; [Cer nohor sky, 1 967 : 224 , pl . 57, F ig .

m us tel inus H wass in B rugu iere, 1792 ; [Cer nohor sky, 1 978 : 1 27, pl . 5, F ig .

n ussatella L innaeus , 1 758 ; (Kay, 1 979 : 376, F ig . 1 22D].obscur u s Sower by , 1833 ; (Kay, 1 979 : 376 , F ig . 1 22C].omar ia H wass in B rugu ier e, 1 792 ; [Cernohor sky, 1978 : 1 23 , pl . 1 , F ig .

per tus us H wass in B rugu iere, 1 792 ; [Kay, 1 979 : 378 , F ig . 1 23C].

pu l icar ius H wass in Brugu iere, 1 792 ; Kohn , 1980a; Kohn , 1 980b; [Kay, 1979: 378 , F ig . 1 23C].

quer cin us So lander in L ightfoot , 1 786 ; (Kay, 1 979 : 379 , F ig . 1 23J].rattus H wass in B rugu iere, 1 792 ; Ber n stein , 1 974 ; Kohn , 1 980a; Lev i ten and Kohn , 1980 ; [Kay, 1 979:379 , F ig . I Z3E].ret ifer Menke, 1 829 ; Lev iten and Kohn , 1 980 ; [Cer nohor sky, 1 978 : 1 22,

pl . 1 , F ig .

sangu ino len tas Quoy and Gaimard, 1834 ; [Cer nohor sky, 1 967 : 226 , pl . 58 , F ig .

scabr iuscu lus D i llwyn , 1 8 1 7 ; [Cernohor sky, 1 967 : 226 , pl . 58 , F ig .

spon sal is H wass in Bruguier e, 1 792; Bern s tein , 1974 ; Lev i ten , 1 974 ; Kohn and Lev i ten , 1 976 ; Kohn , 1980a;

Levi ten and Kohn , 1980 ; [Kay, 1979: 380 , F ig . 1 23G].s tr iatus L innaeus , 1 758 ; [Kay, 1979 : 380 , F ig .

sugi llatus Reeve, 1 844 ; Kohn , 1 980a; Kohn , l 98ob; [Cernohor sky, 1 967 : 228 , pl . 59, F ig .

tenu is t r iatus Sower by , 1 858 ; [Cernohor sky, 1 967 : 230 , p l . 59 , F ig .

terebra Bo r n , 1 778 ; [Cernohor sky, 1978 : 1 40 , pl . 48 , F ig .

tes su latu s Bor n , 1 778 ; [Cer nohor sky, 1978 : 1 25 , pl . 3 , F ig .

tex ti le L innaeus , 1 758 ; [Cer noho r sky , 1 978 : 1 21 , pl . 1 , F ig .

tu lipa L innaeus , 1758 ; [Cer nohor sky, 1 978 : 137 , p l . 7, F ig .

var iu s L innaeus , 1 758 ; [Cer nohor sky, 1 978 : 135 , pl . 7 , F ig .

vex i l lum Gmel in , 1 791 ; (Kay, 1 979 : 38 1 , F ig . 1 2a].

vi rgo L innaeus , 1 758 ; [Cernohor sky, 1 978 : 13 1 , pl . 43 , F ig .

vitu l inus H wass in B ruguier e, 1 792 ; [Kay, 1 979: 382 , F ig . 1 23K].Fam i ly TEREBRIDAEH as tu la lanceata (L innaeus , [Cer nohor sky, 1 967 : 2 1 0 , pl . 54 , F ig

H .

H .

H .

pen ici l lato (H inds , [Cer nohor sky, 1 967 : 21 2, pl . 54 , F ig .

so lida (Deshayes , M i l ler , 1966 ; as Ter ebra; [Cer nohor sky, 1967 : 212,pl . 54 , F ig .

s tr igi lata (L innaeus , [Cer nohor sky, 1 967 : 21 2, pl . 54 , F ig .

Ter ebr a aff in is Gr ay , 1 834 ; M i l ler , 1 966 ; Ladd , 1 982 : 82 ,pl . 30 ,

Figs . 4 , 5 ; Quater nary .

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi

fi amoena Deshayes , 1 859 ; [Cernohor sky, 1 967 : 1 96 , pl . 49 , F ig .

areo lata (L ink , [Cem ohosky, 1 967 : 1 96 , pl . 49 , F ig .

argus H inds , 1 844 ; M i ller , 1 966 ; [Cernohor sky, 1967 : 1 96 ,pl . 49 , F ig .

baby lon ia Lamarck , 1 822 ; M i l ler , 1 966 ; [Cer nohor sky, 1 967 : 1 98 ,pl . 49 , F ig .

cer i th ina Lamar ck , 1822; [Cernohor sky, 1 967 : 198 , pl . 49 , F ig .

chlorata Lamar ck , 1 822 ; [Cer nohor sky, 1 967 : 1 98 , pl . 49 , F ig .

cingu l i/era Lamar ck, 1 822 ; M i ller , 1 966 ; [Cer nohor sky, 1967 : 198 , pl . 49, F ig

co lumel lar is H inds , 1 844 ; [Cer nohor sky, 1 967 : 1 98 , pl . 49 , F ig .

consper sa H inds , 1844 ; [Cer nohor sky, 1 967 : 1 98 , pl . 49 , F ig .

cr enu lata (L innaeus , Mi ller , 1966 ; [Cer nohor sky, 1 967 : 1 99 , pl . 50 , F i g .

dim idiata (L innaeus , M i ller , 1966 ; [Cernohor sky, 1967 : 1 99, p l . 50 , F ig .

fel ina (Di llwyn , Mi l ler , 1 966 ; as Terebra t ign'na Gmel in , 1 791 ; [Cer nohor sky, 1 967 : 199,pl . 50 , F ig .

lenes tr ata H inds , 1 844 ; [Cernohor sky, 1 967 : 200 , pl . 50 , F ig .

fun icu lata H inds , 1 844 ; [Cernohor sky, 1 967 : 200 , pl . 50 , F ig .

gu tt ata (Rod ing , M i l ler , 1 966 ; [Cer nohor sky, 1 967 : 202 , pl . 50 , F ig .

k i lbu r n i Burch , 1965 ; [Cernohor sky, 1 967: 202 , pl . 50 , F ig .

maculata (L innaeus , Mi l ler , 1 966 ; [Cer nohor sky, 1 967 : 204 , p l . 51 , F ig .

nebu los a Sower by , 1 825 ; M i ller , 1 966 ; [Cernohor sky, 1 967 : 204 , pl . 51 , F ig .

paucis t r iata (E . A . Sm i th , (Cem ohor sky, 1 967 : 205 , p l . 51 , F ig .

per tusa (80 m , [Cernohor sky, 1 967: 205 , pl . 52, F ig .

quoygaimardi Cem ohor sky and B r atcher , 1 976 ; [Cernohor sky, 1978 : 1 46 , pl . 52, F igsubu lata (L innaeus , M i l ler , 1 966 ; [Cernohor sky, 1 967 : 205 , pl . 52 , F ig .

text i l is H inds , 1 844 ; [Cer nohor sky, 1 967 : 206 , pl . 52, F ig .

undu lata Gray , 1 834 ; [Cer nohor sky, 1967 : 208 , pl . 52 , F ig .

(Th is table con t inued on next page,

MOLLUSCA 125

TABLE 1 (con t ’d)Fam i ly TEREBR IDAE (cont ’d)TTer ebr a (St r ioterebr a) sp . A . Ladd , 1 982 ; Ladd, 1982: 82 , pl . 30 , F ig . 3 ; late Miocene.

TTerebr a (Oxymer is) sp . 8 . Ladd, 1982 ; Ladd , 1 982 : 84 , pl . 30 , F ig . 14 ; late M iocene .

Terenolla pygmaea (H inds , [Cernohor sky, 1 978 : 1 50 , pl . 53 , F ig .

[T he r ecords of T . cance l lata Quoy and Gaimard, 1 832 , T . laevigata Gr ay , 1 834 , and T . p icta H inds , 1 844 ,

cited by M i ller ( 1 966) have not been confi rmed ]Subc las s OPISTH OBRANCH IAOr der ENTOMOTAENIATAFam i ly PYRAMIDELL IDAEChemn itz ia cf . C. gabr ieli (H edley , F ig . 5A .

Chr ysalida alveata (A . Adams , F ig . SE .

Con t r ax iala obl iqua Laseron , 1 956 ; [Ponder , 1 985 : 1 08 , F ig 75A— B].Cos tabieta hor r ida (Gar r ett , [Cer nohor sky, 1 978 : 48 , Figs . 1 3 , 1 4 ; as R is so ina].Evalea debi lis (Pease, [Pease, 1 868 : 292 , pl . 24 , F ig .

E . denses t r iata (Gar r ett , F ig . 58 .

E . peasei (Dau tzenberg and Bouge, (Kay, 1979 : 407 , F ig . 1 32A].H er vier a g l i r iel la (Melv i ll and Standen , [Kay, 1 979 : 407, F ig . 1 32C].Mi r alda gemm ifera (Dau tzenberg and Fischer , [Dau tzenberg and Fischer , 1906 : 1 95,

pl . 6 , F ig .

Odos tom ia ex i l is (Ga r r ett , F ig . 5C.

0 . gu l icki P i lsbry, 1 9 1 8 ; [Kay, 1 979 : 4 1 1 , F ig . 132J].O . ox ia Watson , 1 886 ; (Kay, 1 979 : 4 1 , F ig .

OtOp leu ra m it ra lis (A . Adam s , [Cer nohor sky, 1 972 : 201 , pl . 57 , F ig .

O . nodicincta (A . Adams , [Cer nohor sky, 1 972 : 20 1 , pl . 57 , F ig .

Pyram idel la pus i l la A . Adams , 1 854 ; [Dal l and Bar tsch , 1906 : 324— 325, pl . 24 , F ig .

P . sulcata (A . Adams , [Cernohor sky, 1 972 : 200 , pl . 57 , F ig .

Pyrgu l ina cf . P . densecos tata (Gar r ett , F ig . 5D .

P . oodes (Watson , [Kay, 1 979 : 41 4 ,F ig . I 3ZN].

Syr no la lutea (Gar r ett ,S . po lita (Pease,

Tu r bon i lla cf . T . infan tu la Dal l and Bar t sch , 1 906 ; [Dal l and Bar tsch , 1 906 : 338 ,pl . 20 , F ig .

T . l i r ata (A . Adams , [Kay, 1 979 : 414 , F ig . 1 33F].T. or thop l icatu lata Nomur a, 1 936 ; (Nomura, 1 936 : 74 , pl . 1 2,

Figs . 1 0 1a, 10 1b].T . p seudomala Nomur a, 1 938 ; (Nomur a, 1 938 : 29, pl . 4 , F igs . 33a, 33b].T . cf . T . ten u is s im a H edley , 1 909 ; [H edley , 1 909 : 450 , pl . 42 , F ig .

Order SOLEOL IFERAFam i ly ONCH ID I IDAEOnch idiella eve l inae Mar cus and Burch , 1965 ; Mar cus and Bur ch , 1 965 : 253 ,

Figs . 36 — 39 .

Peron ia per son i i (Cuv ier , Mar cus , 1 965 : 264 .

Or der CEPH ALASPIDEAFam i ly ACTAEON IDAEPupa alveo la (Souver bie, (Cer nohor sky, 1 972 : 204 , pl . 58 , F ig .

P . su lcata (Gmel in , [Cernohor sky, 1 972 : 203 ,pl . 58 , F ig .

Pupa sp . ; F ig . 5H .

[T he record of Acteon var iegatus (B rugu iere, 1 789) ci ted by Ekdale et al . (1 979) has not been confi rmed ]Fam i ly RING ICUL IDAER ingicu la cf . R . do lear is Gould , 1 860 ; F ig . 5F .

Fam i ly AMPLUSTRIDAEH ydat ina amp lus t re (L innaeus , Johnson and Boucher , 1 983 : 254 ; (Ber tsch and Johnson , 1 98 1 :

H . phys is (L innaeus , John son and Boucher , 1 983 : 254 ; (Ber tsch and Johnson , 1 981 :

M icromelo guamens is (Quoy and Gaimard, Johnson and Boucher , 1 983 : 255 ; (Bert sch and Johnson ,

Fam i ly SMARAGD INELL IDAEPhaner ophthalm us smaragdinus (Ruppell and Leuckar t , Mar cus and Burch , 1 965 : 238 ; as LathOphthalm us

smaragdin us (Ruppell and Leuckar t , (Kay, 1 979 : 422 ,F ig .

Smar agdinella ca lyculata (B roder ip and Sowerby , Marcus and Burch , 1 965 : 236 — 238 , F ig . 1 ; [Kay, 1 979 : 423 ,

F ig . 1 36D].Fam i ly BULL IDAEBul la vem icosa Gould , 1 859 ; Johnson and Boucher , 1 983 : 255 ; [Kay, 1 979 : 423 , F ig . 1 37E].Cyl ichnatys angu s ta (Gould , F ig . 5M .

TSpecies not now l iv ing at Enewetak ; hor i zons recorded by Ladd (1 966 , 1 972 , 1 977 , (Thi s table cont inued on next page.)

KAY AND JOH NSON

TABLE 1 (con t’d)

Fam i ly ATYIDAEA tys cyl indr icus (H elbl ing , [Cernohor sky, 1 972 : 208 , pl . 59, F ig .

A . debi lis Pease, 1 860 ; [Kay, 1 979 : 424 , F ig . 1 37C].

A . cf . A . kuhn s i P i lsbry , 1 91 7 ; [Kay, 1 979 : 425 , F ig .

A . naucum (L innaeus , [Cer nohor sky, 1 972 : 208 , pl . 59 , F ig .

H am inoea cr ocata Pease, 1 860 ; (Kay, 1 979 : 426 , F ig . 1 37E].cur ta (A . Adam s , [Kay, 1 979 : 427 , F ig . 137I].cymba lum (Quoy and Gaimard,

Marcus and Burch , 1965: 243 ; as Lamp roham inoea cym ba lum(Quoy and Ga imard, [Ber tsch and Johnson , 198 1 :

ga lba Pease, 1 86 1 ; (Kay, 1 979 : 427 , F ig . 1 37G].

cf . H . japon ica Pi lsbry, 1 895 ; F ig . 5G .

linda Marcus and Burch , 1 965 ; Marcus and Bur ch , 1965 : 241 , Figs . 1 2— 1 6 ; [Johnson , 1 982 :

musetta Marcus and Burch , 1 965 ; Marcus and Bur ch , 1965 : 239 , F ig . 6 .

Fam i ly RETUSIDAEColeophys is m in ima (Yamakawa, F ig . 5L .

Fam i ly AGLAJIDAEChel idon u ra h i r undin ina (Quoy and Gaimard, [Kay, 1979: 429 , F ig . 1 38E].

C . inor nata Baba , 1949 ; Johnson and Boucher , 1983 : 255 ; [Johnson , 1 982 :

Odon tog laja guamens is Rudman , 1 978a; [Rudman , 1 978a: 90 , F ig .

Phi l inop s is gardiner i El iot , 1903 ; Johnson and Boucher , 1 983 : 255 ; [Bergh , 1 905 : pl . 3 , F ig .

Fam i ly GASTROPTERIDAEGas trop teron br unneomargi natum Car lson and H off , 1 974 ; [Car lson and H off , 1 974 : 347 , pl . X , F ig .

Fam i ly SCAPH ANDR IDAEActeocina cf . A . hawa iens is P i lsbry, 1 921 ; (Kay, 1 979 : 43 1 , F ig . 1 37M].A . sandwicen s is Pease, 1 860 ; [Kay, 1 979 : 43 1 , F ig . 1 37N].Adamnes t ia japon ica (A . Adams , F ig . SI .

Nakam igawaia spi r a lis Kuroda and H abe,1961 ; F ig . SJ.

Order ANASPIDEAFam i ly APLYSI IDAEAp lys ia dactylomela Rang , 1 828 ; [Kay, 1979 : 437, F ig . 1 3 1A].A . par vu la Gu i lding in Mor ch , 1 863 ; Johnson and Boucher , 1 983 : 255 ; [Kay, 1979 : 439 , F ig . 1 3 1C].A . p u lmon ica Gould , 1 852 ; Mar cus , 1 965 : 266 .

Dolabella aur icu lar ia (L ightfoo t , Johnson and Boucher , 1 983 : 256 ; (Kay, 1 979 : 440 , F ig . 1 40D].Dolabr i/era dolabr ifera (Rang , Marcus and Bur ch , 1 965 : 244 ; [Ber t sch and Johnson , 1 98 1 :Stylochei lus longicaudus (Quoy and Gaimard, Marcus and Bur ch , 1 965 : 244 ; Johnson and Boucher , 1 983 :256 ; [Kay, 1 979 : 440 , F ig .

Order SACOGLOSSAFam i ly PLAKOBRANCH IDAEE lys ia bayer i Mar cus , 1 965 ; Johnson and Boucher , 1983 : 257 ; [Car lson and H off , 1978 : 9 1 , F ig .

margi nata (Pease, Johnson and Boucher , 1983 : 257 ; [Car lson and H off , 1978 : 102 , F ig .

halimedae MacNae, 1954 ; Johnson and Boucher , 1983 : 257 ; [Car lson and H off , 1 978: 99, F ig .

liv ida Baba, 1 955 ; Johnson and Boucher , 1983 : 257 ; [Car lson and H off , 1978 : 1 00 , F ig .

mercier i (Pr uvot -Fol , [Ca r lson and H off, 1978 : F ig .

obtusa Baba , 1938 ; Johnson and Boucher , 1983 : 257; (Baba,1949 : 13 1 , pl . 9 , F igs . 28 ,

ratna Marcus , 1965 ; Johnson and Boucher , 1 983 : 258 ; [Car lson and H off , 1978 : 1 07 , F ig .

vatae R i sbec, 1 928 ; Johnson and Boucher , 1 983 : 258 ; [Car lson and H off , 1 978 : 1 08 , F ig .

P lakobranchus ocellatus van H asselt , 1 824 ; Johnson and Boucher , 1983 : 256 ; [Ber tsch and Johnson , 1 98 1 : 20

Fam i ly H ERMAEIDAESt i l iger smar agdin us Baba , 1 949 ; Johnson and Boucher , 1983 : 256 ; [Baba, 1 949 : 1 29 , pl . 7 , F ig .

Fam i ly CALYPH YLLIDAECyer ce elegans Bergh , 1 888 ; (Kay , 1 979 :

Order NOTASPIDEAFam i ly PLEUROBRANCH IDAEBer thell ina cit r ina (Ruppell and Leuckar t , W i llan , 1 984 : 37

— 40 , Figs 33 — 36 ; [Ber ts ch and Johnson ,

1 98 1 : 26

Ber thella pe l lucida (Pease, Wi llan , 1 984 : 40 — 43 , Figs . 6 , 7 , 1 7, 1 8 , 29 , 37— 39 , 44 .

B . mar tens i (P i lsbry, W i llan , 1 984 : 43— 45 , 5 1 , 52 , Figs . 8 — 1 2 , 2 1

— 28 , 30 — 32 , 40— 42 , 44 .

Pleu r ohedra har aldi Marcus and Mar cus , 1 970 ; Wi llan , 1 984 : 38 — 40 , Figs . 1 — 5 , 13 — 26 , 1 9 , 20 .

I

I

I

I

I

I

mmrn

rn

rn

rn

m

(Thi s table cont inued on next page. )

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1 28 KAY AND JOH NSON

TABLE 1 (con t'd)

Fam i ly CH ROMODORIDIDAE (cont ’d)N . nor ba Mar cus and Mar cus , 1970 ; Johnson and Boucher , 1 983 : 275 ; as Thor unna nor bo ; [Marcus and Mar cus ,1 970 : 1 61 , F ig .

N . pus i l la (Ber gh , [Bergh , 1 874 : p l . 1 , F ig .

R isbecia imper ialis (Pease, [Ber tsch and Johnson , 1981 : 50— 51 ; as Chr omodor is godeffr oyana (Bergh ,

R . t ryon i (Gar r ett , Johnson and Boucher , 1 983 : 268 ; (Gar r ett , 1 873 : 232, pl .Thor unna aus t ralis (R isbec , Johnson and Boucher , 1 983 : 270 ; as H ypeselodor is aus t ralis ; [Johnson , 1982 : 92;

as H yp selodor is aus tralis].T . dan iellae (Kay and Young , (Kay and Young , 1969: 207 , F ig .

T . fur tive Bergh , 1874 ; (Rudman , 1984 :

T . p u rp u repedis Rudman and Johnson , 1 985 ; Rudman , 1 985 : 289— 291 , Figs . I 2H , 27 , 29 , 30 .

Fam i ly H EXABRANCH IDAEH exebr anchus sanguineus (Ruppell and Leuckar t , Mar cus , 1 965 : 271 ; as H . marginatus (Quoy and

Gaimard, Johnson and Boucher , 1 983 : 259 ; [Ber tsch and Johnson , 1 98 1 : cover].Fam i ly DENDRODORID IDAEDendrodon

'

s coronata Kay and Young , 1 969 ; Johnson and Boucher , 1983 : 276 ; (Kay and Young , 1 969 :

21 8 , F ig.

D . elongata Baba, 1936 ; Johnson and Boucher , 1983 : 276 ; [Baba , 1949 : 1 56 , pl . 28 , F ig.

D . er ubescens (Ber gh , Marcus and Bur ch , 1965 : 250 .

D . n igr a (St impson ,Mar cus and Bur ch , 1 965: 250 ; Young , 1 967 : 1 68 ; Johnson and Boucher ,

1983 : 277 ; [Ber tsch and Johnson , 1981 :

D . rubra (Kelaar t , (Or r , 198 1 :

D . tubercu losa (Quoy and Gaimard, [Kay and Young , 1969 : 219, F ig.

Fam i ly PH YLL ID IIDAEPhyll idie pus tu losa Cuv ier , 1 804; Johnson and Boucher , 1983 : 282; [Ber tsch and John son, 1 981 : 74

P . var icose Lamarck , 1 80 1 ; Johnson and Boucher , 1983 : 282 ; [Ber tsch and Johnson , 1981 : 74

Fam i ly GYMNODORIDIDAEGym nodor is ceylon ice (Kelaar t , Johnson and Boucher , 1 983 : 277 ; [R isbec , 1 928 : 1 82, pl . 7 , F ig .

G . citr ina (Bergh , Mar cus and Bur ch , 1965 : 245 ; as Gym nodor is bicolor (Alder and H ancock,Young , 1 967 : 1 69 ; Johnson and Boucher , 1 983 : 278 ; (Baba, 1 949 : 40 , pl . 1 1 , Figs . 37,

G . okinawee Baba , 1 936 ; Johnson and Boucher , 1 983 : 278 ; [Ber tsch and Johnson , 1 98 1 :

G . p lebe ia (Bergh , Johnson and Boucher , 1983 : 279; [Kay and Young , 1969 : 224, F ig .

G . s tr iata (El iot , Johnson and Boucher , 1983 : 279; [Johnson , 1 982 : 93 ; as Analogi um s tr iatum].Nembr o tha cr is tete Bergh , 1 877 ; [Wi llan and Co leman , 1 984 : 12 , F ig .

Roboas t ra gr aci lis (Bergh , (Baba , 1949 : 136 , pl . 1 3 , F ig .

Fam i ly TRIOPH IDAEP locemopher us ceylon icus (Kelaar t , Johnson and Boucher , 1 983 : 281 ; ( Thompson , 1975 : 509, pl . 1a].Fam i ly NOTODORIDIDAEAegi res ci t r inus Pruvot -Fol , 1 930 ; [R isbec, 1 953 :A . p unct i lucens (d

’

Or bigny, [Baba, 1 974 : F ig .

A . vi l losus Far ran , 1 905 ; Johnson and Boucher , 1 983 : 28 1 ; [Edmunds , 1 971 : 379 , F ig .

Fam i ly GON IODOR ID IDAEG on iodor idielle savignyi Pr ti vot

-Fol , 1 933 ; Johnson and Boucher , 1 983 : 282 ; [Baba, 1960: 30 1 , pl .Gon iodor is joubin i R isbec, 1 928 ; Johnson and Boucher , 1983 : 282 ; (R isbec, 1 928 : 1 75, pl . 5 , F ig .

Fam i ly DORIDOMORPH IDAEDor idomorphe gardiner i E l iot , 1 906 ; [Rudman , 1 982 : F ig . 28a].Fam i ly VAYSSIEREIDAEOkadaia e legans Baba, 1 930 ; Young , 1 967 : 1 71 ; 1 969 .

Fam i ly EMBLETONIIDAEEm bleton ia g raci lis R isbec , 1928 ; [Gos liner and G ri ffi ths , 1 981 : 1 42Fam i ly ARMINIDAEDermatobranchus fort unate (Bergh , Johnson and Boucher , 1983 : 282;[Bergh , 1888 : 353 , pl .

Fam i ly BORNELLIDAEBom ella angui l la Johnson , 1 985 ; John son , 1 985 : [Wi llan and Co leman , 1 984 : 50 , F ig .

B . s tell ifer (Adams and Reeve in Adam s , Johnson and Boucher , 1 983 : 282; as Bom ella edem s i i Gr ay , 1 850 ;[Wi llan and Co leman , 1984 : 50 , F ig . 1 61 ; as Bo r nelle edem s i i Gray ,

Fam i ly MARIANINIDAEMar ian ina rosea (Pruvot -Fo l , John son and Boucher , 1 983 : 283 ; [Ca r lson and H off , 1 973 : 1 72 ;

as A r an ucus bif idus Odhner ,(Th is table cont inued on next pag

MOLLUSCA 129

TABLE 1 (con t'd)

Fam i ly CUTHONIDAEPhes t i lle lugubn

'

s (Bergh , Johnson and Boucher , 1 983 : 284 ; [Ber tsch and Johnson , 1 98 1 : 90— 91 ;as Phes t i lle s ibogee Bergh ,

Fam i ly CORYPH ELL IDAEF label l ina al isonae Gos l iner , 1 980 ; Gos liner , 1 980 : 40 ; Johnson and Boucher , 1 983 : 283 ; [Ber t sch and Johnson ,

1 98 1 :

Fam i ly GLAUCIDAEG laucus at lan t icus For ster , 1 777 ; Johnson and Boucher , 1 983 : 285 ; [Ber tsch and Johnson , 1981 :

Fam i ly AEOL IDIIDAEBaeol idia nodosa (H aefelfi nger and Stamm , [Gos liner , 1 980 : 66

Cer ber i lle aff in is Bergh , 1 888 ; (McDonald and Nybakken , 1 975 : 380

Favor in us japon icus Baba, 1 949; Johnson and Boucher , 1 983 : 283 ; [Ber tsch and Johnson , 1 98 1 : 96

F . m i r abi lis Baba, 1 955 ; [Baba, 1 955 : 53 , pl . 1 7, F ig .

H er viella claror Bur n , 1963 ; Marcus and Bur ch , 1 965 : 251 ; Figs . 28— 30 ; Young , 1967 : 172 .

H . m ietta Mar cus and Bur ch , 1 965 ; Mar cus and Bur ch , 1 965 : 251 ; Young , 1 967: 1 71 .

Ph idiena bou r ai ll i (R i sbec, John son and Boucher , 1 983 : 285 ; (Rudman , 1 980 : 1 46, F ig .

P . indica (Bergh , Johnson and Boucher , 1 983 : 285 ; (Ber tsch and Johnson , 1 98 1 : 95 ; as Celon'

e m i l itar is(A lder and H ancock ,

Phyllodesm ium hyal inum Ehr enberg , 1 83 1 ; Johnson and Boucher , 1 983 : 284 ; (Rudman , 198 1 : 22, F i g .

Pteraeol idie ian th ina (Angas , Johnson and Boucher , 1983 : 284— 285 ; [Ber tsch and Johnson , 198 1 : 98

Sp u r i lla major (El iot, Johnson and Boucher , 1983 : 285; [Ber tsch and Johnson , 1981 :

Subclass PULMONATAOr der BASOMMATOPHORAFam i ly MELAMPIDAEMe lampus flavus (Gmel in , [Cer nohor sky, 1 972 : 21 3 , p l . 6 1 , F ig .

Fam i ly SIPH ONAR IIDAESiphonar ia guamens is (Quoy and Gaimard, Marcus and Burch , 1 965 : 256 ; [H ubendick , 1 946 : 41 ,pl . 6 , Figs . 30

S . normal is (Gould, Menge, 1 973 ; Lev iten , 1 974 ; [Cer nohor sky, 1 972 : 21 0 , pl . 60, F ig .

Order ORTH URETH RAFam i ly ACH ATINELL IDAELamell idea pus i lla (Gould ,Fam i ly PUPIL L IDAEGas t r ocop ta pedicu lus (Shutt lewor t h , Reigle, 1964 : 1 28 .

G . ser vi lis (Gould,Or der SIGMURETH RAFam i ly SUBUL INIDAESubu l ine octona (Bruguier e,

Lemellax is gr aci l is (H utton , Reig le, 1 964 : 1 28 .

Subo rder AULACOPODAFam i ly CH AROPIDAETVatu s i le en iwetokens is (Ladd , Ladd , 1 958 : 1 90 , p l . 30 , Figs . 9— 12; Miocene g ; as Ptychodon ; So lem ,

1 982 : 1 95 , Fi gs . 85e— f .

lass B IVALV IAFam i ly ARCIDAEAnedar a an ti quata (L innaeus , [Cer nohor sky, 1 972 : 215 ,

pl . 6 1 , F ig .

A r ea ave l lana Lamar ck , 1 8 1 9 ; (K i lbum , 1 983 : 51 4, Figs . 2 ,

A . ven t r icose Lamarck , 1 81 9 ; [Kay, 1 979 : 500 , Figs . 1 6 1C— D].Bar bat ie amygdalum tos tum (R6ding , (Ki ra, 1 962 : 122, pl . 43 , F ig . 14 ; as Barbatie bicoloreta

decussata (Sower by , (Kay, 1 979: 500 , Figs . 1 6ZG — H].divar icata (Sower by , (Kay, 1 979 : 501 , Figs . 1 61E— C].nu tt ingi Dal l , Bar tsch , and Rehder , 1 938 ; [Kay, 1 979 : 503 , Figs . 1 62I— J].tene l la (Reeve, [Kay, 1 979 : 504 , Figs . 1 62C— D].cf . 8 . wendt i (Lamy, F ig . 6A .

Fam i ly GLYCYMERIDIDAEG lycymer i s r eevei (Mayer , F ig . 68 .

Fam i ly MYT IL IDAEBo tu la s i licule (Lamar ck , (Kay, 1979 : 509; Figs . 1 64E— F].

L ithophage z ittell iena Dunker , 1860 ; F ig . 6E .

UU

CD

UJ

CD

UD

TSpecies not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd (1 966 , 1 972 , 1 977 , 1 982) (Thi s table cont inued on next page. )

KAY AND JOHNSON

TABLE 1 (con t ’d)

Fam i ly MYTIL IDAE (con t’d)Myti lus edu lis L innaeus , 1 758 ; on floating p lastic fi sh ing floats ; [Abbott , 1954 : pl . 35, F ig. M].Modio lus aur icu latus Kr auss , 1 848 ; [Salvat and R ives , 1975 : 367, F ig .

M . phi l ipp inar um (H an ley , (K i r a, 1962 : 1 26 , pl . 46 , F ig .

M . vagina (Lamar ck , F ig . 6C.

Sep tifer bryanae (P i lsbry, [Kay, 1 979 : 51 2, F ig . 164P].Fam i ly PINN IDAEA tn

‘

na vex i l lum (80m , (Rosewater , 196 1 : pls . 156 ,

Pinna mur icate L innaeus , 1758 ; Rosewater , 1961 : 1 91 , pl .Str ep top inna seccata (L innaeus , Rosewater , 1 961 : 22 , pl . 169 .

Fam i ly PTERIIDAEElect r oma electr ine (Reeve, F ig . 6D .

P inctada margar i tfer a (L innaeus , [Kay, 1 979 : 516 , Figs . 1 66O— E].Pteri a pengu in (R6ding , [Cernohor sky, 1 978 : 1 77, pl . 62 , F ig .

Fam i ly ISOGNOMONIDAEIsognomon den t ifere Kr auss , 1 848 ; [Salvat and R ives , 1975 : 370 , F ig . 415 ; as I . pect inate (Reeve,

I . incisum (Conr ad, [Kay, 1 979 : 520 , Figs . 1 67C— D (not E

I . legumen (Gmel in , [Kay, 1 979 : 520 , Figs . 1 67G — H ].

I . per na (L innaeus , [Kay, 1 979 : 521 , F igs . 1 67C— D].Melina eph ipp ium (L innaeus , [Cernohor sky, 1 978 : 1 78 , pl . 63 , F ig .

Fam i ly MALLEIDAEMa lleus regu le (For sskal , (Kay, 1979 : 521 , Figs . 1 67I— K].

Fam i ly PECT IN IDAECh lamys cor uscens (H inds , Waller , 1 972 : 23 1 , p l . 1 , Figs . 1 — 7,

C. wi lhelm in i Bavay , 1 904 ; Waller , 1 972 : 236 , pl . 2, Figs . 30— 33 , 36— 37 ; as C . mar shallens is Wal ler , 1 972 .

Comp topall ium vex i l lum (Reeve, Waller , 1 972 : 243 , pl . 4 , Figs . 64— 75 .

TExcell ichlamys spectabi lis (Reeve, Wal ler , 1 972 : 249 ; H o locene; (Wal ler , 1 972 : 246 , pl . 5, Figs . 85

G lor ipall ium pal l ium (L innaeus , Wal ler , 1 972 : 239, pl . 3 , Figs . 45— 56 .

Jux tam us ium coudein i (Bavay , Wal ler , 1 972 : 250 , Figs . 1 1 1 — 1 27 ; as Jux tamu s ium

maldviens is (E . A . Sm i th ,M irapecten m ir if icus (Reeve, (Kay, 1979 : 526 , Figs . 1 69A— B].Pedum spondylo ideum (Gmel in , Wal ler , 1 972 : 254 , pl . 8 , Figs . 1 36 — 1 43 .

Fam i ly SPONDYLIDAESpondy lus hys tr ix Bo lten in Rad ing , 1 798 ; [Kay, 1 979 : 530 , Figs . 1 76G — H ; and see

8 . n icoben'

cus Schr eiberg , 1 795 in Cernohor sky, 1978 : 1 80 , pl . 63 , F ig .

S. l inguaefel is Sower by , 1 874 ; [Kay, 1 979 : 53 1 , Figs . 1 71D — E].S . squemosus Schr eiber , 1 793 ; [H i r ase, 1951 : pl . 15, F ig .

S . tenebrosus Reeve, 1 856 ; (Kay, 1979 : 532, Figs . 171A— C].8 . cf. 8 . vari ans Sower by , 1 829 ; [Cer nohor sky, 1 978 : 1 80 , pl . 64 , F ig .

Fam i ly PLICATUL IDAEP l icat u la s imp lex Gould, 1 86 1 ; [Johnson , 1964 : 149 , p l . 25 , F ig .

Fam i ly L IM IDAEL ima annu lata Lamarck , 1 8 19 ; [Cernohor sky, 1 972 : 220 , pl . 62 , F ig .

L . concen tr ica Sower by , 1 888 ; [Cer nohor sky, 1 978 : 1 8 1 , pl . 65 , F ig .

L . fr agi lis (Gmel in , [Kay, 1 979 : 534 , Figs . I 7ZA— B].L . l ime vu lgar is (L ink , [Cernohor sky, 1 972 : 21 9 , p l . 61 , F ig .

Fam i ly OSTRE IDAEOs t rea cf . 0 . hen leyana Sower by , 1 871 ; (Kay, 1 979 : 537 ,

Figs . 1 73D — E].0 . cf . 0 . sandv icens is Sower by , 1 871 ; [Kay, 1 979: 539 ,

Figs . 1 73A — B].Fam i ly CH AMIDAEChame broder ip i i Reeve, 1 846 ; (Reeve, 1 846 : Chame No .

C . ios toma Conr ad, 1 837; [Cer nohor sky, 1978 : 1 83 ,pl . 65, F ig .

C . lazarus L innaeus , 1 758 ; [Cernohor sky, 1978 : 1 83 , p l . 65 , F ig .

C . pacifica B r oder ip , 1 835 ; F ig . 6M .

Fam i ly LUCIN IDAECodakia punctate (L innaeus , [Cern ohor sky, 1972 : 221 , pl . 62, F ig .

Ctene belle (Con rad, [Kay, 1 979 : 543 , Figs . 1 76A— B].Lucine eden tu la (L innaeus , [Kay, 1979 : 543 ,

Figs . 177A — B].L . cf . L . er ythraea Is sel , 1 869 ; [Bouchet and Danr i gal , 1 982 : F ig .

TSpecies not now l iv ing at Enewetak ; hor i zons r ecorded by Ladd ( 1 966 , 1 972 , 1 977 , (This table con t inued on next pas

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1 32 KAY AND JOH NSON

TABLE 1 (con t ’d)

Fam i ly GLOSSIDAEMeiocerdia tet r agona (Adams and Reeve, (K i r a, 1962 : 1 48 , pl . 53 ,

F ig .

Fam i ly VENER IDAEDos in ia cf. D . m i t icu le V iader , 1937 ; F ig . 6N .

Gaf rar ium pect inatum (L innaeus , (Cer nohor sky, 1 972 : 234 , pl . 67, F ig .

G . tum idum (Rod ing , [Cer nohor sky, 1972 : 234 , pl . 67 , F ig .

G lycodon ta mer ica (L innaeus , [Cer noho r sky, 1 972 : 237, pl . 67 , F ig .

L ioconche cas trens is (L innaeus , [Cer nohor sky, 1972 : 235, pl . 5, F ig .

L . om ata (D i l lwyn , [Cer nohor sky, 1972 : 236 , p l . 67, F ig .

Per ig lyp te puerpera (L innaeus , [Cernohor sky, 1 972 : 233 , p l . 66 , F ig .

P . reticu lata (L innaeus , [Cer nohor sky, 1 972 : 233 , pl . 66 , F ig .

Pitar pe l lucidus (Lamarck , [Cer nohor sky, 1972 : 236 , pl . 68 , F ig .

Venus toreuma Gould , 1 850 ; [Kay, 1 979 : 570 , F ig . 1 84L].

Fam i ly GASTROCH AEN IDAEGas t rochaena cuneiform is Speng ler , 1 783 ; [Kay, 1979 : 570 , F ig . I 8SA].

Fam i ly PHOLAD IDAEMar tes ia s tri ate (L innaeus , [Kay, 1 979 : 573 , F ig . 185C].

Class POLYPLACOPH ORAOr der CH ITON IDFam i ly ACANTHOCH ITON IDAECryp top lax el iot i Pi lsbry , 1 90 1 .

Fam i ly CH ITON IDAETLucu l ine r us sell i Ladd, 1966 ; Ladd , 1 966 : 23 , p l . 1 , Figs . 13 — 1 5; pr obably P l iocene.

Fam i ly ISCH NOCH ITONIDAEIschnoch iton sp .

Fam i ly SCH IZOCH ITONIDAETSchiz ochiton mar shallens is Ladd , 1966 ; Ladd, 1 966 : 22, pl . 1 , Figs . 4— 9 ; ear ly M iocene.

Fam i ly LEPIDOPLEURIDAELep tochiton acum inatus (Th iele,

Class CEPH ALOPODAOrder OCTOPODAFam i ly OCTOPOD IDAEOctopus cyanea Gray , 1 849; (Kay, 1979 : 589 , Figs . 1 948 — E].

Order TEUTH OIDEAFam i ly OMMASTREPH IDAEH yaloteu this pelagica (Bosc, Voss , 1 954 : 365.

Subc lass NAUT ILOIDEAFam i ly NAUT IL IDAENauti lus macromphelus Sower by , 1849 ; Shel ls occas ional ly washed up on windward beaches ; [Cer nohor sky, 1 972 :240 , pl . 7, F ig .

N . pomp i lius L innaeus , 1 758 ; Shel ls occas ional ly washed up on windward beaches ; [Cer nohor sky, 1972 : 240 ,

pl . 7, F ig . 71.

TSpecies not now l iv ing at Enewetak ; hor i zon s recorded by Ladd ( 1 966 , 1 972, 1977 ,

Cass id idae (Abbott , M i tr idae (Cer nohor sky,D r upa (Emer son and Cer nohor sky,

Rhinoc lavis (H oubr ick , Clypeomor u s (H oubr ick ,

H arp idae (Rehder , Gabr ielona (Rober tson ,

P inn idae (Rosewater , T r idacn idae (Rosewater , and L i ttor in idae (Rosewater , Co l lect ionsof op i s thobr anchs made by J . B . Bur ch and W . H ear dwer e descr ibed by Mar cus and Bur ch (1 965) and Mar cus

Young (1 967 , 1 969) r epor ted on hi s own co llec

t ion of opi s thobr anchs , and Johnson and Boucher ( 1 983 )r ecor ded 76 species from co llect ions they made at

Enewetak . Enewetak pect in ids wer e d i scus sed by Wal lerA paper by Ing r am ( 1 947) and a manuscr ipt by

Ing r am and Mo r r i son (MS, USNM) d iscus s the cowr ies of

the Mar shal l Is lands and include r eco r ds fr om Enewetak .Mos t s tud ies of Enewetak mo l lusks have been eco log i

cal l y o r ien ted . As ear ly as 1 956 , Kohn made co l lect ionsand obser vat ions on the gas tr opod genus Conus , and hiswor k on Conus (and that of hi s s tuden ts ) cont inues (Kohn ,

1 967 ,1 97 1 , 1 980a , 1 980b ; Kohn and Lev iten , 1 976 ;

Lev i ten and Kohn , 1 980 ; Lev i ten ,Eco log ical s tud

ies on other gr oups include those on the T r idacn idae(Stasek , 1 963a, 1963b ; Fankboner , 1 971a,

1 971 b , 1972 ;

Lev i ten and Osenber g , Ter ebr idae (M i l ler ,Ner i ta (Stokes ,

Mur ic idae (Menge,1 974 ; Ber ns tein ,

Cyp r aea moneta (Renaud , Str ombidae

MOLLUSCA 133

(Berg , Vermet idae (H opper , 1 975 , Mor u la(Sinc lai r , Stylochei lus (Sar ver , Phes t i lle

(Sw i t zer , Siphonar ia (Menge ,and so ft bot

tom commun i t ies (Ekdale et al . ,Enewetak has also been the s i te of resear ch on the

genet ic var iabi l i ty and the ph ys io logy of the T r idacn idae(Valen t ine and Ayala, 1 974 ; Ayala et al . , 1 973 ; Valen tineet al . , 1 973 ; Fankboner , 1971a, 1 971 b, of phys iolog ical s tud ies oi the Str ombidae (G i llary, 1 974 ; G i llaryand G i llary, and of s tud ies of gr owth and mor tal i tyrates in mar ine snai ls (Tur k , l 979a, 1 979b) .

MODERN MOLLUSCAN F AUNA

N onmar i ne Mol lu sk s

The nonmar ine mo l luscan fauna, compr ised of eightspec ies , con forms to the descr ipt ion of C. M . Cooke Jr .

(in H ar r y , 1 966) of a typ i cal ato l l land snai l fauna :ver y r es tr icted . made up mos t ly of spec ies that areacc idental l y d is tr ibuted by man .

”Thus , of the eight species

we r ecor d , five (both Gas tr ocop ta, both Subu l in idae, and

Lamell idea pus i lla) ar e pr obably advent ive and w idely distr ibuted among the is lands of the Paci fic as a r esult of theact iv i t ies of man (see Chr is ten sen and K i r ch , 1 98 1 ; H ar r y ,1 966 ; and So lem 1 964 for d iscus s ion of Paci fic bas in landsnai ls and thei r d i sper sal by humans) . The r emain ing thr eenonmar ine mo l luscan spec ies are s tr andl ine spec ies :Melampus flavus and Tr uncatel la guer n i i are w idel y dis t r ibu ted on Paci fic i s lands ; As s im inee n itida mar shallens is is

endemic to the Mar shal l Is lands , accor d ing to AbbottThe l i s t of Enewetak nonmar ine mo l lusks is s im i lar

to that comp i led for neighbor ing Rongelap in the Mar shal lIs lands (Reigle, Of the 1 1 spec ies recor ded fromRongelap , three of the five adven t ive spec ies also occur atEnewetak , and the two ato l ls also shar e two of the s trandl ine species .

TABLE 2

Char acter i s t ics of Pac i f i c Is land Mar i ne Mol lu scan Faunas

Enewetak

76 8 1 79

323 32 472

421 44 6 1 1

1 74 1 7 1 49

Total 994

Total mo l lusk species 1 1 1 6

Gas t r opodzB ivalve Ratio90 : 1 0

‘KurodaTKay

TH edley

Q MN o. 96 No.

1lKay and Sw itzer

H awal l i Funafu ti ; Fann i ng“N o. 96 No. N o. 96

1 3 50 6 35 1 1 16 5

34 292 35 154 48 1 49 51

44 344 40 106 33 98 33

8 1 71 20 34 8 32 10

Mar i ne Mol lu sk s

A t leas t thr ee factor s affect the recor ds of the presenceor absence of mar ine mol lusks at Enewetak and, indeed ,on any ato l l . The three factor s ar e the co llect ing effor t ,k inds of hab i tat avai lable, and the chance occur r ence or

r ecr u i tmen t of young to the is land .

G iven 40 year s of co llect ion act iv i ty and the numer ousco l lector s who have v is i ted Enewetak , the appr oximately1 000 species of mar ine mo llusks that ar e r ecor ded as

occu r r ing at Enewetak today are pr obably fai r ly r epr esentat ive of the mo l luscan fauna that occur s between the in tert idal zone and depths of about 30 m . In term s of number sof spec ies , the Enewetak mo llus can fauna is about thatof the i s land of Ok inawa, 2500 m i les nor thwes t ; jus ts l ight ly lar ger than that recor ded for the H awai ian Is lands ,2000 m i les nor theas t ; and two and a half to thr ee t imeslarger than those r epor ted for two Pac i fic ato lls , Far mingand Funafut i (TableThe fauna is dom inated by gas tr opods , and the

gast ropodzbivalve rat io of is the h ighes t of all Paci ficfaunas for wh ich ther e are r ecor ds (Table The gas tr opod r ecor d includes 947 spec ies in 1 09 fam i l ies . Neogas

t r opods compr ise 4496 of the gas tr opod species , mesogast ropods (includ ing H eterogas t r opoda), eu thyneu rans ,

and ar chaeogas t ropods , about 896 of the mo l lusks .

134 KAY AND JOH NSON

The fam i l ies w i th the greates t number of species ar e the

Tu r r idae Con idae M i tr idae Cypraeidaeand the Cos tellar i idae

The bivalve r ecor d includes 30 fam i l ies and 1 1 5

species . Among the bivalves , 51% of the species are infau

nal , 41% ar e epi faunal bys sate forms , and 696 ar e

fr ee- l iv ing species . The fam i ly Tel l in idae has the lar ges tnumber of species fo l lowed by the Vener idaeA r cidae and Car di idae

Com pos i t ion

Ther e are some not iceable anomal ies in the compos it ion of the Enewetak fauna in that sever al well -knownIndo-Wes t Paci fic mo l lusks are consp icuous by thei rabsence. We do not r ecor d the tu rbin id Lep tothyra p icta,

the ner i t id Smar agdia, the r is so id Mereline p is inna, the ver

met id Dendr opoma max ima , the mesogast ropod l impetH ippon ix , the thaid Vex i lla, the buccin id Can thar usfumos u s , or the heterogas t r opod A r chitecton ica. Sever alspecies such as Gyr ineum gyr ineum and Mor um den ison i

wh ich ar e r ecor ded from Kwajalein ar e absen t at

Enewetak .

Some of the absences are exp l icable . I f a habi tat isabsent , then the spec ies occu r r ing in that hab i tat ar e alsoabsen t . Thus Smaragdia associated w i th sea gr ass bedsand gas tr opods associated w i th vo lcan i c subs trates such asH ippon ix and the mur icid Vex i l la (Vermeij et al . , 1 984) arenot expected to occur at Enewetak . Other mor e sub

t le eco log ical r es tr ict ions have been sugges ted by Wal ler( 1972) who has shown that sever al species of EnewetakPect in idae are extremel y res tr icted in thei r hab i tats . Thesespec ies include Comp topall ium vex i llum wh ich l ives in as so

c iat ion w i th ter r igenous sed iments and mar ine gr asses

around elevated is lands and is found on ly as juven i les inthe deeper po r t ions of the lagoon of the ato l l ; and

Jux tam us ium maldivense, wh ich also l ives in the deeperpor tions of the lagoon , in as sociat ion w i th abundan tH al imeda on wh ich i t may depend for attachment and con

cealment .

The absence of gas tr opods such as the r eef-associatedLep to thyr a p icte, Mer el ine p is in na, Dendr opoma max ima,

and A r ch i tecton ica is not eas i ly explained and, for the

presen t , i s attr ibuted to chance as has been sugges ted forcer tain aspects of the mar ine fauna of the L ine Is lands(Kay, the H awai ian Is lands (Kay, 1 979 , the

nor ther n Mar ianas (Vermeij et al . , and among the

i s lands of the Ind ian Ocean (Taylor ,

B iogeog r aphy

W i th one except ion , the subspec ies As s im inee n i t ida

mar shallen s is Abbo tt , 1 958 , all the Recen t mo l lusks iden t ified to spec ies have been r ecor ded elsewher e in the IndoWes t Pac i fic . As Wal ler ( 1 972) has noted , the s ign i ficanceof the latter obser vat ion l ies in the demon s tr at ion that , forPaci fic i s land mo llusks , m i les of open water w i thout s tr ongcur r en t sys tems have not r esulted in m uch d i ffer en t iat ion .

The mar ine mo l lusks of Enewetak ar e clear ly of IndoWes t Paci fi c der ivat ion , and many species have r anges

wh ich span the Indo-Wes t Paci fic fr om the H awai ianIs lands on the eas t to the coas t of eas t Afr ica on the wes t .A componen t of th is fauna i s r es tr icted to the Pac i fic Plate(sen su Spr inger , 1 4 of the 56 cowr ies and five of

the 1 9 s t r ombids are Paci fic Plate endem ics .

Four spec ies requ i r e spec ial mention . Shel ls of two

species of Nauti lus and specimens of M yt i lus edu l is ar e

r epor ted fr om w indwar d beaches , but ther e is no ev idencethat ei ther Nauti lus or M yti lus l ives at Enewetak . Onespecies , Tr ochus n i lo t icus , was intr oduced by the Japaneseabout 1 935 , and these mo l lusks are ver y common .

FOSSIL MOLLUSKSF oss i l Recor d

The Enewetak mo l lus can foss i l recor d was documen tedby Ladd ( 1 966 , 1 972 , 1977, 1 982) who descr ibed and/orr ecor ded 274 spec ies pr imar i l y fr om two dr i l l cor es on the

ato l l . Ladd dealt on l y w i th ch i tons and gas tr opods and,un for tunately , was unable to comp lete hi s s tudy of the

Enewetak foss i l mater ial . Of the gas tr opods r ecor ded byLadd ( 1 966 et ar chaeogas t r opods compr ise 2596;

mesogas t r opods (includ ing H eterogas t ropoda) , 5696; and

neogas t r opods , 1 796 of the foss i l r ecor d .

A lthough r elat ively numerous in terms of species , theEnewetak foss i l recor d is far from complete. Apar t fr omthe eu thyneu rans , bivalves , cephalopods , and scaphopodswh ich have not been cons idered , the neogas t r opods ar e

under r epr esented , and there are biases in other r espects .

There are pr opor t ionatel y mor e lagoonal forms such as r is

so ids and cer i thi ids than ther e ar e seawar d r eef flat speciessuch as cones and mu r ic ids . H igh in ter t idal li ttor in ids and

ner i t ids are not wel l represented . Four of the s ix foss i l ner i t ids are subt idal forms and on ly one l i tt or in id (Tector ius)is r ecor ded . Sti l l another bias is in s i ze— 82% of the mea

su r ements for shel l s i zes ar e less than 10 mm in gr eates td imens ion and the larges t foss i l shel l is on ly 34 mm in

lengt h . Despi te these defic iencies , the foss i l recor d ,together w i th the cu r r en t checkl is t , al lows at leas t thr eeconclus ions :

1 . Species compos i t ion at Enewetak has changedw i th t ime.

2 . Man y species were w idely d i s tr ibuted in the

Paci fic in the M iocene .

3 . Pr ev ious l y w ide spec ies r anges have become r e

s t r icted wi th t ime .

Geolog i cal Recor dThe Enewetak geo log i ca l r eco r d compr ises a Ter t iar y

sequence extend ing back in to the Eocene and a Quaternar y sur face Iimes tone sect ion . Ladd (1 966) r ecogn i zed s ixsubd iv is ions of the Ter t iar y , the late Eocene (Ter t iar y b) tothe Pl iocene (Ter t iar y h) , based on the sys tem developedfor the Eas t Ind ies by van der V lerk and Umbg r ove (1 927)

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KAY AND JOHNSON

F ig . 2 a, Planax is zouatna, heigh t 7 mm ; b, Caecum cf . C.

‘

fol i n i , lengt h mm ; c, Tu r r i tel la sp . A . , hei gh t1 8 mm ; (1, Tun

-i teIIa sp . B . . heig ht 1 7 mm ; e, Sca l lo la ar enosa, hei gh t 3 mm : f, S. ca ledon ica, hei gh t 2 mm ;9 , S. g lar eosa, heig ht 3 mm ; h , Inc l la japon i ca , hei gh t 3 mm ; 1, N anophor a cf. N . leucomys , hei gh t mm

1, Bouchet r iphor a quadr imacu lata, hei ght 3 mm ; k , Verm icu lar ia sp . , hei ght 10 mm ; I. Cer i th i um claviform e,

hei gh t 25 mm ; m , M etax ia sp . , hei gh t 3 mm ; n , Cer i th iops is sp . A hei ght 4 mm ; o, Cer i th iops is sp . 8

hei ght 4 mm ; p, E u thymel la r ega l is , hei gh t 8 mm ; q , Mas ton ia us tu lata. heigh t 3 mm ; r , Mesophore cnodax,

hei gh t 5 mm ; s , N anophor a sp . . heig ht 3 mm .

MOLLUSCA 137

F ig . 3 a, Sty lapex sp . , hei gh t mm ; b. Balcis acicu la, hei gh t 8 mm ; c, H em i leios t r acum sp . , heigh t 2 mm ;(1, Lameller ia sp . , leng th 8 mm ; e, Epi ton i um au r i tum , hei ght 1 9 mm ; f—g , N ever l ta sp . , hei gh t 5 mm ; h- I.

M or u la squamosa, hei gh t 22 mm ; j- k , N ever i te cf. N . pet iver iana, hei ght mm ; l , Pyr ene obtusa, hei ght 1 2mm ; m , E up l ica am i r an t ium , hei gh t 4 mm ; n , Co lumbel la cf . C. molecu l ina, hei ght 13 mm ; o, Pyr ene cf. P.l ivescens , hei gh t 10 mm ; p, P. a lb ino, hei gh t 1 2 mm ; q , Eup l ica cf. E . l iocym a, heigh t mm ; r , E . cf. E .

m inu te, hei ght 6 mm ; s . Sem i nel la d ivar ica ta, heig ht 4 mm ; t, S. deu t zenber g i , height 4 mm ; u, S. subph i lo

d icia, hei ght 4 mm ; v , Pyr ene punctata, hei gh t 1 3 mm ; w , P. abegu ie, hei ght 4 mm .

KAY AND JOHNSON

F ig . 4 a, G r anul ina vi t ree, heigh t 3 mm; b. Kogomou ovate, hei gh t 4 mm : c. Kogomea sp . A height

mm ; d, Tr i ng inel la sp ., hei gh t mm ; e. H e logi nel la m icr os , heigh t 3 mm ; f , H . keppelens is , hei gh t 7 mm ; g ,Kogomea ber n ier i , heigh t mm ; h , Anac i thar a s t r iata, heigh t 3 mm ; 1, A . nan isca, hei gh t 3 mm ; 1, A . note

b i l ls , hei gh t 3 mm ; k , Clavus sp . B ., heigh t 7 mm ; l , C. b i l ineata, hei gh t 10 mm ; m , Clavus sp. A. n .

Euc i thar a ci thar a, heigh t 1 2 mm ; o, E . daedalea, heigh t 6 mm ; p , E . sp . , heigh t 15 mm ; q, Kerm ia cf . K .

g r an icos tata, height 6 mm ; r , E t r ema t r igonos toma, hei ght 2 mm ; s . Kerm ie conci nna, hei gh t 10 mm ; t,Ph i lber t ia cr as s i lab r um , heigh t 1 2 mm ; u, Kerm ia edychr oa, heigh t mm ; v , L ienardia goubi n i , hei gh t 5mm ; w, Kerm ia idiomo rpha, heigh t 3 mm .

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F ig . 6 a, Bar bat ie wendt i , length 6 mm ; b, G lycymer is r eevei , length 1 5 mm ; c. Mod io lus vagi na, mm ; (1.

Elect r oma electr i ne, lengt h 10 mm ; e, L i thophage z i tellel iana, lengt h 40 mm ; f, Tel l ina pudica , lengt h 6 mm ; 9 , T.

ovate , lengt h 37 mm ; h , Leev icar dium pu lcher r im um , lengt h 18 mm ; i , Tel lina inflate, length 1 7 mm ; 1. Sponder vi l ias implex , length mm ; k, Tel li na sp . , lengt h 5 mm ; I , Macoma obl iquan

’

e, lengt h 6 mm ; m , Chame sp . , length 20

mm ; n , Dos i n ia m i t icu le, leng th , 10 mm .

MOLLUSCA 14 1

when the ato l l s tood above the sea as a h i gh i s land .

Leopold ’

s (1 969) repo r t of spor es and po l len ind i cates a

r icher and mor e var ied flora on Enewetak in the M iocenethan now occu r s . The recor d of a h igh i s land foss i l landshel l , Vatus i la (Ladd , 1 958 as Pytochodon ; So lem ,

is ev idence of cond i t ions qu i te d i fferen t fr om those today .Leopo ld ( 1 969) also demons trated conclus ively fromanalys i s of po l len gr ain s that mangr oves occu r red at

Enewetak thr ough the M iocene. The limes tone sect ionsar e all r eef assoc iated . Many of the fos s i l i ferous beds ,

especial ly those r ich in mo l lusks , appear to have beendepos i ted in lagoons , but other env i r onments —

r eef wal l ,open shoal , and off -

reef depos i ts —

are also found” (Ladd ,

M iocene Recor d

The M iocene r ecor d , the r iches t and mos t d iver se of

the foss i l recor ds , includes 1 70 spec ies , 2696 archaeogas

t ropods , 5896 mesogas t r opods (includ ing heter ogas t r o

pods ), and 1 7% neogas t ropods , repr esen t ing 90 genera

and 42 fam i l ies of mo l lusks .

The M iocene fauna i s ver y d i fferen t fr om that found atEnewetak today . The ar chaeogas t r opods are repr esentedby eigh t fam i l ies ; today ther e ar e 1 3 fam i l ies . Tu r bin ids ,t rochids , and ner i t ids compr i se 7 196 of the M iocenespec ies r ecor d ; t r ochids , f is su r ell ids , s tomatell ids , and tu r

bin ids repr esent 6896 of the pr esen t ar chaeogas t r opod

fauna. Among the mesogas t r opods , the r is so ids (29

spec ies ) are the mos t numer ous fam i l y compr is ing 3096 of

the r ecor d ; the cer i th i ids ( 15 spec ies ), and v i t r inell ids

and dial ids ( 12 species ) about 1 396 of the r eco r d . Todaythe cyp r aeids , cer i thi ids , r i sso ids , s tombids , v it r inell ids , and

cymat ids dom inate the mesogas t r opods .

Some of the d i ffer ences between the M iocene and

moder n faunas undoubtedly r eflect b iases in the foss i lr eco r d as such . Other d i ffer ences ind icate the pr esence of

hab i tats such as sea grass beds , mangr ove swamps , br acki sh water lagoons , and h igh is land shorel ines wh ich ar e not

now pr esen t on the atol l . Thus four genera pr esen t in the

M iocene r ecor d ar e ext inct on Enewetak today but ar efound elsewhere in the Indo-Wes t Pac i fic in env i r onmentswh ich do not presen t ly occur at Enewetak . The ner i t idSmar agdia (thr ee spec ies in the M iocene r ecor d) is as soc iated w i th sea gr ass beds at Kwajalein , Mar shal l Is lands , inthe Caro l ine Is lands , and as far eas t as the H awai ianIs lands (Kay, The sess i le l impet- l ike mesogas t r opodH ippon ix (two species in the M iocene r ecor d) is found onthe volcan ic shor es of the H awai ian Is lands , on Guam , and

in the nor ther n Mar iana Is lands (Vermeij et ai . , The

cer i thi ids Ter ebr el ia and Potam ides (five spec ies in the

M iocene recor d) occur in mangr ove assoc iat ions in the

Ph i l ipp ines and elsewher e in the lndo -Malayan ar ch ipelago .

In all , 1 2 species or 796 of the M iocene fauna r ecor ded byLadd (1 966 , et seq . ) are assoc iated w i th habi tats not now

r epr esented at Enewetak .Other ext inct ions may r epr esen t r ange r es tr ict ions for

reasons other than habi tat change . Ner i ta in scu lp ta,

Tectar ius , P r otobar leeia, Pug i l ina, and Eoc ither e ar e not

reco r ded after the M iocene . Ner i ta inscu lp ta is now foundin Indones ia and the Nor ther n Mar ianas but appar en tly noton i s lands to the eas t . On l y one spec ies of Tector ius , T .

g rand inatus , now occur s on the Pac i f ic Plate, and i t is r e

s t r icted to southeas t Po lynes ia (Rosewater ,P rotobar leeia myer s i now sur v ives in F i j i and Aus tr al ia(Ponder , 1983b) . Although the Melongen idae do occur inthe Indo-Wes t Pac i fic , no species of Pug i l ina is known fr omthe Paci fic Plate.

St i l l other ext inct ions ar e even more puzzl ing . A

num ber of spec ies r ecor ded from the M iocene of Enewetaksur v ive on other Paci fic is lands , includ ing Kwajalein .

Among the spec ies are Lep to thyra laeta, L . candida,

L . ver r uca , Mer el ine p is inna, R is so ina con cinna , R . s up

r acos tate, M i t ra turg ida , and Vex i l lum app r ox imatum .

Pos t -M iocen e Recor d

The pos t -M iocene r ecor d cons is ts of 105 species of

mo l lusks : 2396 ar chaeogas t ropods , 6 196 mesogas t r opods ,

and 1 696 neogas t r opods . Mos t of these mo llusks ar e

extan t . Twen ty-three or 2296 ar e ext inct at Enewetaktoday , although some sur v ive elsewhere in the Indo-Wes tPac i fic , among them Synep tocochlea r osacea, Lep tothyra

p icte , and NannOpho re c ingu l ifere.

Spec ies Su r v i val

Ladd (1 966) po in ted ou t ear ly that man y l iv ing mol

lusks in the Mar s hall Is lands have been there s ince lateTer t iar y t ime. Mor e r ecentl y Stan ley (1 979) and Stan leyet al . ( 1980) commented on the gr eat s tabi l i ty of the

Indo-Wes t Pac i fic fauna" and sugges t that species havetended to su r v ive for unusual ly long per iods in the r eg ion .

The Enewetak foss i l r ecor d together w i th the moder nrecor d pr ov ides s tr ong ev idence for the gener al i zat ion : 62spec ies fr om the M iocene r ecor d are ident i fied as l iv ingtoday at Enewetak . They r epr esent 3596 of that r ecord .

F aunal Relat ion s

The closes t t ie in the fos s i l r eco r d between Enewetakand another locale i s w i th B ik in i . About 34% of the

species found in the d r i l l ho les at Enewetak wer e alsofound in dr i l l ho les at B ik in i . Apar t fr om the B ik in i r elat ionshi p ,

the s tr onges t t ie for wh ich there is a publ i shedr ecor d i s w i th F i j i w i th 1396 of the Enewetak M iocenespec ies s har ed . The recor d is less s trong w i th Guam and

Saipan (496 of the M iocene r ecor d) and JavaSever al pos s ibly far -flung r elat ionsh ips were c i ted by

Ladd Among them he notes that H ep locochl ies(one spec ies in the M iocene) i s r ecogn i zed as a wes ter nAmer ican genus , and the tu r bin id Cyn isca (one species inthe M iocene) is known on l y from South Afr ica. Nei ther of

these r elat ions h ips has been subsequently con fi rmed . H ow

ever , the shel ls of San son ia kenneyi , des cr ibed from the

M iocene of Enewetak , ar e ind is t ingu ishable fr om M iocenes hel ls fr om Italy and Pl iocene shells fr om Fr ance (LeRenar d , per sonal commun icat ion) .

142 KAY AND JOHNSON

Fr om the H o locene r ecor d , 6596 sur v ive today on

Enewetak .Of the 140 extinct spec ies l is ted by Ladd (1 966 , et

8496 became extinct in the M iocene, in the

H olocene . The Enewetak r ecor d may be con tras ted wi ththat r epor ted by Newman (1 986) who calculated fr omLadd ( 1982 : Table 4) that of 1 26 extinct ions s ince the

Eocene, 56 or became ext inct in the Plei s toceneand on ly five of the extant spec ies sur v ived fromthe M iocene. Th i s las t figure is too low for Enewetak andmay be pecul iar to the foss i l r ecor d of Fi j i and the New

H ebr ides (Vanuatu) .There is no s ingle patter n of ext inct ion and sur v ival

such as Wel ls (1 982) found for the M idway cor als , whereat the end of the Miocene ther e was complete r eplacementof the M iocene genera. At Enewetak , the Tur bin idae and

R i s so idae show a patter n in wh ich equal number s of

species have been ext in ct s ince the M iocene, occur bo th inthe M iocene and today , and appear on l y in the moder n

r ecor d . The tr och id , ner i t id , and l i t tor in id r ecor ds , in con

tr as t , ar e almos t en t i r ely moder n .

ACKNOWLEDGMEN TSThe checkl i s t of Enewetak mo l lusks (Table 1 ) i s based

on the r efer ences ci ted in the In tr oduct ion , the Enewetakco l lections in the USNM , the EMBL r efer ence col lection(now housed at the 8 . P . B i shop Museum , H onolulu ,H awai i ) , mater ial co llected for an ear l ier checkl is t by A . S.

Ber n s tein and A . M i l ler wh ich has been incorpo rated intothe Enewetak co l lections at B . P . B ishop Museum , and

obser vat ions by the author s . These obser vat ions ar e basedin par t on exam inat ion of pr ivate shell co l lect ions of m i litar y per sonnel s tat ioned at Enewetak between 1978 and

1 983 . We ar e especial ly grateful to Car o l H opper , R ichar dH oubr ick , and the late Joseph Rosewater for thei r contr ibut ions to the checkli s t .

Numer ous people have con tr ibuted to the lis t ingthr ough co l lect ion and/or ident i fi cation of mol lusks fr omEnewetak and by way of thei r ass is tance w i th the

l i ter atur e. Among them are A . Adams , H . Ber ts ch ,W . O . Cer nohor sky, C. Chr is tensen , P . Co l in , the late D .

Devaney , A . Fer rei r a, R . H oubr ick, J . Lamber son ,

J . Le Renar d , V . O . Maes , W . 8 . Rudman , E . Vokes ,and T . Wal ler . L isa Boucher and Lor i Bel l have been of

par t icular help in locat ing unr ecor ded species . Specialthanks go to R ich Nakagawa for the u se of his unpubl ishedl is t ing of Kwajalein shel ls and to Jeannete H ammon , Karen

deG roff , Dave Johnson , D . J . MacDonald, Lee Rousseau ,and Jim and Judy Wedge for addi t ional in format ion and

ass is tance in finding Kwajalein mater ial .The sen ior author is gr ateful to Kenneth Bos s and

Ruth Tur ner of the Museum of Compar at ive Zoo logy ,H ar var d Un iver s i ty ; Ph i lippe Bouchet , Labor ato i r e des

Inver tébr és mar in s et de Malacolog ie, Par i s ; George Dav i s ,Academy of Natur al Sc ience, Ph i ladelph ia; Oliver Pagetand Er har d Wawr a, Natu ralhi stor i ske Museum ,

V ienna; thelate Joseph Rosewater , Depar tment of Inver tebrates , U . S.

Nat ional Museum , Sm i thson ian Ins t i tut ion ; and J . D . Taylor , Mo l lus ca Sect ion , B r i t i sh Museum (Natural H is tor y),London , for perm i s s ion to ut i l i ze the co l lect ions in thei rcare.

REF ERENCES

Abbott , R . T . , 1 954, Amer ican Seashel ls , D . Van Nos tr and Co . ,

lnc. , New York .1 958 , The Gas tr opod Genus As s im inee in the Ph i l ippines ,Proc . Acad . Nat . Sci . , 1 10: 21 3-278 .

1 959 , The Fam i ly Vas idae in the Indo Pacifi c, Indo -PecificMo l lusca, 1 : 15-32 .

1960 , The Genus Str ombus in the Indo-Pacific , Indo -Pacif icMo l lusca, 1 : 33 - 1 44 .

1 96 1 , The Genus Lam bis in the Indo Paci fic , Indo-Pacif icMo l lusca, 1 : 147- 1 74 .

1967 , Str om bus (Canar ium) wi lson i New Species fr om the

lndo Pac i fic , Indo -Pecif ic Mo llusca, 1 : 455-456 .

1 968 , The H elmet Shells of the Wor ld (Cass ididae),Indo -Pacif ic Mo l lusca, 1 : 1 -202 .

Afshar , F . 1969 , Taxonom ic Rev is ion of the Supr aspecific

Gr oups of the Cr etacous and Cenozo ic Tel l in idae, Geo l . Soc .

Amer ica Mem .,1 1 9 , 21 5 pp .

Ayala, F . J. , D . H edgecock, G . S. Zumwalt , and J . W . Valentine,1973 , Genetic Var iation in Tr idacne max ima, an Ecologi calAnalog of Some Unsucces s fu l Evo lutionar y L ineages ,

Evo lut ion , 27: 1 77- 191 .

Baba, K ., 1 949 , Op is thobranch ia of Sagem i Bay Co l lected by H is

Majes ty the Emperor of Japan , Iwanam i Shoten , Tokyo .

1 960 , The Genera Oken ia, Gon iodor idielle and Gon iodor is ,Publ . Seto Mar . B io l . Lab . , 8: 299-302 .

1 974 , New D i s tr ibutional Record of Aegi res p unct i lucen s

(d’

Or bigny , 1 837) fr om Sado Is land, Japan (Nudibr anch ia:Dor idacea: Aeg i ret idae), Veliger , 1 7: 1 1 - 1 2 .

Berg , C. J. , 1 974 , A Comparative Etho log ica l Study of StrombidGas tr opods , Behaviour , 51 : 274-322 .

Bergh , R . , 1874 , Neue nacktschnecken der sudsee, malacolo

gische unter suchungen , J . M us . Godeff roy, 2: 91 -1 1 6 .

1 888 , D ie Pleu r oleur iden , eine fam i l ie der nudibr anchiaten

gas t r opoden , Zoo l . Jahrb . Abt . Sys t . , 3 : 348-364 .

1 905 , D ie Opis thobr anch iata der Siboga-Expedition ,

Siboga-Exped . Rep ts .

, Mon .

, 50: 1 -248 .

Bern stein , A . S.,1 974 , D iet and Competi tion for Food Among

the Pr edator y Gas tr opods of L imes tone Benches in H awai iand En iwetok , Ph .D . disser tation , Un iver s i ty of Or egon .

Ber tsch , H . , and S. John son , 1 98 1 , H awai ian Nudibr anchs , Or iental Publ . Co . , H ono lu lu .

Boss , K . J . 1969, The Subfam i ly Tellinae in South Afr icanWater s (B ivalv ia, Mo l lusca), Bu ll . M u s . Comp . Zoo l . , H ar var d,138 : 81 - 1 62 .

Bouchet, P . , and F . Danr igal , 1 982, Napo leon ’

s Egyptian Campaign ( 1 798- 1 80 1 ) and the Sav igny Co l lection of Shel ls ,N aut i lus , 96 : 9-24 .

Burgess , C . M . 1 970 , The L iving Cowr ies , A . S. Bar nes and Co . ,

New Yor k .Car lson , C. H . , and P . J . H off , 1973 , Exter nal Descr iption of a

L iv ing A r anucus bif idus (Odhner , 1 936) (Op isthobranch ia:Dendr onotacea) , Veliger , 15: 1 72- 173 .

1 974 , The Gas t rop ter idae of Guam , wi th Descr iptions of

Four New Species (Op is thobr anch ia: Cephalasp idea), Publ .Seto Mar . B io l . Lab

, 21 : 345-363 .

https://www.forgottenbooks.com/join

144 KAY AND J

Cypraeidae Based on the Co l lections of the Amer icanMuseum of Natural H is tor y , Amer . Mus . Novitates , 1336 : 1 -4 .

and J . P . E . Mor r ison , n .d . , Cypr aeidae of the Mar shal lIs lands , MS. , U . S. National Museum .

Ir edale, T . , 1 91 7 , On Some New Species of Mar ine Mol luscafr om Chr istmas Is land, Indian Ocean , Proc . Malaco l . , Soc . ,

Lond . , 1 2: 33 1-334 .

Johnson , R . l . , 1 964 , The Recent Mo l lusca of Augustus AddisonGou ld, B u ll . U . S . Nat l . Mus . , 239, 1 82 pp .Johnson , S . , 1 982 , L iv ing Seashe l ls , Or iental Publ . Co . , H ono lulu .

1985 , A New Indo-West Pacific Species of the Dendr onota

cean Nud ibranch Bom ella (Mo l lusca : Op i sthobr anch ia) withAngu i l l i form Swimm ing Behav ior , M icrones ica, 1 9: 1 7-26 .

and H . Ber tsch , 1985 , Rediscover y and Redescr iption of

Ros tanga lu tescens (Bergh , 1 905) Comb . Nov . (Gastr opoda:Nudibr anch ia) , Vel iger , 27: 406-41 0 .

and L . M . Boucher , 1 983 , Notes on Some Opi sthobr anchia(Mol lusca : Gas tr opoda) from the Mar shal l Is lands , Including 57New Records , Pec . Sc i . , 37: 251 -291 .

Jung , P . , and R . T . Abbott , 1967, The Genus Terebel lum (Gast ropoda : Str ombidae), Indo -Pacific Mo l lusca, 1 : 445-454 .

Kay, E . A . , 1 965 , Mar ine Mo l luscs in the Gum ing Co l

lection , Br i t ish Museum (Natur al H istor y) Descr ibed byW i l l iam H arper Pease, Bul l . B r i t ish Mus . (Nat . H is t .) Zoo logy,Supp l . 1 , 96 pp .

1971 , The L i ttoral Mar ine Mo l luscs of Fann ing is land, Pac.

Sci . , 25: 260-281 .

1 979, H awai ian Mar ine She l ls , B . P . B i shop Mu s . Spec .Publ . , 64 653 pp .

1 980 , L itt le Wor lds of the Pacific, an Essay on Pacif ic Bas inB iogeogr aphy, Un iver s ity of H awai i , H ar o ld L . L yon Ar bo r etum Lecture, 9: 40 pp .

and M . F . Switzer , 1 974 , Mo l luscan D istr ibution Pattern s inFann ing is land Lagoon and a Compar ison of the Mol lusks of

the Lagoon and the Seaward Reefs , Pec. Sc i . , 28 : 275-295 .

and D . K . Young , 1969, The Dor idacea (Opisthobranch ia;Mo l lusca) of the H awai ian Is lands , Pec . Sci . , 23 : 1 1 2-23 1 .

K i lbur n , R . N . , 1 983 , The Recent Ar cidae (Mo l lusca: B ivalv ia) ofSouther n Afr ica and Mozambique, Ann . Natal Mus . , 25:

51 1 -548 .

K i r a, T . , 1962 , She l ls of the Wes ter n Pacif ic in Co lor , H o ikusha,

Osaka.

Kohn , A . J. , 1967 , Env i ronmental Complex ity and Species Di vers i ty in the Gastropod Genus Conus on Indo-West Paci fic ReefPlatforms , Amer . Nat . , 101 : 251 -259 .

1 971 , D iver s ity , U ti l i zation of Resour ces , and Adapt ive Radiation in Shal low Water Mar ine Inver tebrates of T r opicalOcean Is lands , L imnol . and Oceanog . , 16: 332-348 .

1 980a, Abundance, D iver s ity and Resource U se in an

Assemblage of Conus Species in Enewetak Lagoon ,Pac. Sc i . ,

34: 359-370 .

l 980b , Populat ions of T r opica l Inter t idal Gastr opods Beforeand After a T yphoon , M icr ones ica, 1 6: 21 5-228 .

and P . J . Lev iten , 1 976 , Effects of H abi tat Complex i ty onPopulation Dens ity and Species R ichness in T r op ical Inter tidalPredator y Gastropod Assemblages , Oecologia, 25: 1 99-21 0 .

Kosuge, S. , 1 96 1 , On the Fam i ly T r iphor idae (Gastr opoda) fromAmam i Is lands , Venus , 21 : 308-3 1 6 ; 4 1 3-4 1 5 .

1 962 , Descr ipt ions of 1 0 New Species and 1 New Sub

species of the Fam i ly T r iphor idae (Mo l lusca) fr om Shiono

m isak i , K i i Pen insula, Central Japan with a L i st of H ither toKnown Species , Bu l l . Nat l . Sc i . M us .

, Tokyo , 6 78-89 .

1963 , On the Fam i ly T r iphor idae (Gas tropoda) fr om Amam i

ls lands , Venus , 22: 240 -257 ; 257-263 .

LLUSCA 145

1 970 , The Fam i ly L i ttor in idae in the Indo Paci fi c,

Indo -Pecific Mo l lusca, 2: 41 7-534 .

Rudman , W . 1978a, A New Species and Genus of the

Aglajidae and the Evo lution of the Phi linacean Opis thobr anchMo l luscs , J. L inn . Soc . , Zoo l . , 62: 89- 1 07 .

1 978b, The Bor id Op is thobr anch Genera H algerda Berghand Sclerodon

’

s El iot fr om the Indo-Wes t Pacifi c, J . L inn . Soc . ,

Zoo l . , 62: 59-88 .

1980 , Aeo l id Opis thobranch Mo l luscs (G laucidae) fr om the

Indian Ocean and the South-West Paci fic,J . L inn . Soc . ,

Zoo l . ,68 : 139- 1 72.

1 98 1 , The Anatomy and B io logy of A lcyonar ian-FeedingAeo lid Opisthobr anch Mo l luscs and Thei r Development ofSymbios i s wi th Zooxanthel lae, J . L inn . Soc . , Zoo l . , 72

219-262 .

1982 , The Taxonomy and B io logy of Fur ther Aeo l idaceanArm inacean Nudibranch Mo lluscs wi th Symbiotic Zooxanthellae, J. L inn . Soc . , Zoo l . , 74: 1 47- 1 96 .

1 984 , The Chromodor ididae (Opi sthobr anch ia: Mo llusca) ofthe lndc es t Pacific : a Rev iew of the Genera,

J . L inn . Soc . ,

Zoo l . , 81 : 1 1 5-273 .

1 985 , The Chr omodor ididae (Op isthobr anch ia: Mo llusca) ofthe lndo Wes t Pacific: Chromodo r is au reomargi nate, C . ver r i

er i and C . f ide lis Co lour Gr oups , J . L inn . Soc .,Zoo l . , 83

241 -299 .

Sal isbur y , R . , 1983 , Notes on Shell Identi fi ca tions , H awai ian Shel lNews , 31 : 6-7 .

Salvat , B . , and C . R ives , 1 975 , Coqui llages de Polynes ie, Les edi

t ions du Paci fique, Papeete, Tah iti .Sar ver , D . , 1978 , Feeding and Gr owth of the Sea H ar e

Stylochei lus long icauda, MPML An n . Rp t . F Y 1 976,pp .

86-88 .

Sinclai r , M . E . , 1 975 , In i tial Investigations of the Foraging Str ategy of the Inter t idal Gastr opod Mor ula granulate, MPML Ann .

Rept . F Y 1 975 , pp . 89-90 .

So lem , A . , 1964, New Records of New Ca ledon ian Nonmar i neMo l lusks and an Analys is of In troduced Mo ll usks . Pec . Sc i . ,

18: 1 30- 1 37 .

1 982 , Endodon to id Land Snai ls fr om Pacific Is lands ,(Mo l lusca : Pulmonata: Sigmu rethra) Par t I I , Fam i l ies Punct idaeand Char op idae , Zoogeog raphy , Fie ld Mus . of Nat . H is t , Chi

cago , 336 pp .

Spr inger , V . G . , 1 982 , Paci fic P late B iogeogr aphy , wi th SpecialReference to Shor ef ishes , Sm ithson ian Con t r . Zoo l . , No . 367,

1 82 pp .

Stan ley , S. M . , 1 979 , Macroevo lut ion . Patter n and Proces s , W . H .

Fr eeman , San Fr ancisco .

W . O . Addicot t , and K . Chinzei , 1 980 , Lyell ian Curves inPaleonto logy : Poss ibi l i ties and L im itations , Geo logy, 8:

422-426 .

Stasek , C . R . , 1 963a, Synops i s and D i scuss ion of the As sociationof Cten idia and Labial Palps in the B ivalved Mo l lusca , Ve liger ,6: 91 -97 .

1 963b, Or ientation and Form in the B ivalved Mo llusca , J .Morph , 1 1 2: 195-214 .

Stokes , D . R . , 1966 , Some Aspects of the B io logy of Ner i tidGas tr opods at En iwetok , M .Sc . (P lan U n iver s i ty of H awai i ,46 pp .

Swi tzer , M . F . , 1971 , Obser vat ion and Co llection of Opi sthobr anch Mo l luscs , EMBL Ann . Rep . F Y 1 971 , p . 26 .

Talmadge, R . R . , 1963 , In sular H al iot ids in the Wes tern Pacific(Mo l lusca : Gastropoda) , Ve liger , 5: 1 29- 1 39 .

Tay lor , J . D . , 1 971 . Reef Associated As semblages in the Wes tern

146 KAY AND JOH NSON

Ind ian Ocean , Regional Var iation in Indian Ocean Coral Reefs ,D . R . Stoddar t and C . M . Yonge, Symp . Zoo l . Soc . , London ,

N O . 28, 584 pp.

Thompson , T . E . , 1 975, Bo r id Nudibranchs from Eastern Au s

tr al ia (Gas tr opoda, Opi s thobranch ia) , J . Zoo l . , 1 76 : 477-51 7 .

Tur k , T . R . , 1 9793 , Patterns of Gr owt h and Mor tal i ty Rates of

Mar ine Snai ls , MPML Ann . Rep . F Y 1 977, 45-54 .

1 979b, Pattern s of Gr owth and Mor tal i ty Rates of Mar ineSnai ls , MPML Ann . Rep . F Y 1 978 , 83-86 .

Valentine, J . W . , and F . J . Ayala, 1 974 , On Scient i ficH ypotheses , K i l ler Clams , and Extinctions , Geo logy , 2: 69-7 1 .

D . H edgecock , G . S. Zumwalt , and F . J . Ayala, 1 973 , MassExtinctions and Genetic Po lymorph isms in the

“

K i l ler Clam ,

”

Tr idacne, Geo l . Soc . Am . Bu l l . , 84: 341 1 -3414 .

van der Vlerk ,I . M . , and J . H . F . Umbg rove, 1 927 , Ter t iai r e

gidsfo ram in iferan van Neder landsch Oos t-Indie,Wetensch .

Mededeel, 6: 1 -3 1 .

Vermei j, G . , E . A . Kay, and L . G . Eldr edge, 1984 , Mo l luscs of

the Nor thern Mar iana Is lands wi th Special Reference to the

Selectiv ity of Ocean ic D isper sa l Bar r ier s , M icrones ica, 19:

27—55 .

Vokes , E . H . , and A . D’

Att i l io , 1 982 , Rev iew of the Mur icidGenus A tt i l iosa (Mo l lusca: Gastr opoda) , Ve liger , 25: 67-71 .

Voss , G . L . , 1 954 , Decapodous Cephalopod Mo l lusks from the

Mar shal l Is lands , Pec . Sci . , 8 : 363 -366 .

Wal ler , T . R . , 1972 , The Pectin idae (Mo l lusca : B ivalvia) o

En iwetok Ato l l , Mar shal l Is lands , Veliger , 14: 221 -264 .

Wal ls , J . G . , n .d. , Cone She l ls , T . F . H . Publ icat ions , lnc . , Nep

tune Ci ty , New Jer sey .Wel ls , J . W . , 1 982, Foss i l Cor als fr om M idway Ato ll , Geo l . Sur v .

Pr of . Paper 6806 , 7 pp .

Wi llan , R . C. , 1 984 , The Pleu robr anchidae (Op is thobr anch ia:Notaspidea) of the Mar shal l Is lands , Centr al-West Paci ficOcean , Vel iger , 27: 37-53 .

and N . Co leman , 1 984 , Nudibranchs of Aus t ralas ia, Australas ian Mar ine Photo Index , Sydney .Young , D . K . , 1 967 , New Records of Nudibranch ia (Gastr opoda:Opi sthobr anch ia : Nudibr anch ia) fr om the Central and Wes tCen tral Paci fic wi th a Descr iption of a New Species , Vel iger ,10: 159- 1 73 .

1 969 , Okadaic e legans , a Tube-Bo r ing Nudibranch Mo l luscfr om the Cen tral and Wes t Central Paci fic, Amer . Zoo l . , 9:903 -907 .

Zeig ler , R . F .

, and H . C . Por r eca , 1 969 , O l ive Shel ls of theWor ld, Zeigler and Por r eca , W . H en r ietta, New York .

Zmar zly, D . L . , 1 984 , Di str ibution and Eco logy of Shal low-WaterCr ino ids at Enewetak Ato l l , Mar shal l Is lands , wi th an Anno

tated Check l ist of Thei r Symbion ts , Pac . Sci . , 38 : 1 05- 1 22 .

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143 SAMUELSON AND NISH IDA

TABLE 1

Checkl i s t of Enewetak In sects and Related A r th r opods

Class ARACH N IDAOr der SCORPION IDA

Fam i ly ISCH NURIDAEH orm u r us aus treles iae Fabr icius , 1 775 : Townes , 1 946 .

Or der PSEUDOSCORPION IDAFam i ly ATEMNIDAEOretemnus samoenu s whar ton i Chamber l in , 1947: Beier , 1957 .

Fam i ly CH ERNETIDAEH ap lochem es ins u lanus Be ier , 1957: Beier , 1 957 .

Fam i ly CH TH ONIIDAELechyt ia sakagam i i Mor ikawa, 1 952 : Beier , 1 957 .

Order ARANEAEFam i ly ARANEIDAE

‘A raneus theis (Walckenaer ,

Fam i ly CLUB ION IDAE‘Ch i racan th ium diver s um Koch , 1 873 .

Fam i ly D IPLURIDAEMas ter ia hi r s u te Koch , 1873 : Roewer , 1 963 .

Fam i ly EUSPARASSIDAE‘H eteropoda venetor ia (L innaeus ,Fam i ly LINYPH I IDAE

‘Er igone sp .

Fam i ly CONOPIDAEOpopaea foveo late Roewer , 1963 : Roewer , 1963 .

‘Opopeea Iene Suman , 1 965 .

Fam i ly PHOLCIDAE‘Smer ingopus elongatus (V inson ,

Fam i ly SALT ICIDAE‘F laci llu la m inu te (Be r land,

‘Vit ia albipalp is Marp les , 1957 .

Fam i ly SCYTODIDAEScytodes s t r iat ipes (Koch , Roewer , 1 963 .

Fam i ly TH ERID I IDAELat rodectus geometr icus (Koch , Townes , 1946 .

Or der ACAR ISubo rder GAMASIDAFam i ly DERMANYSS IDAE

‘Or n ithonyssus sp .

Suborder IXODIDAFam i ly ARGASIDAEOm ithodor os capens is Neumann , 1 901 : Bushman , Parker and Johnson , 1963 .

Suborder ACTIN EDIDAFam i ly ERYTH RAEIDAEBaleu s t ium sp .

Beleu s tu im sp . : Townes , 1 946 .

Class D IPLOPODA (MYR IAPODA in par t)Order PROTEROSPERMOPH ORA

Fam i ly STRONGYLOSOMIDAE‘Ox idu s gr aci lis (Koch ,

Class INSECTAOr der TH YSANURA

Fam i ly LEPISMAT IDAENot iden ti fied : Townes , 1 946 .

Order COLLEMBOLAFam i ly ENTOMOBRYIDAE‘Sei ra bipunctate (Packard ,Fam i ly ISOTOMIDAE‘Is o tom u r us t r icusp is

'

Bo'

r ner , 1 906?

‘New Enewetak reco rd . (Th i s table cont inued on next page. )

INSECTS AND ALLIES (ARTH ROPODA) 1 49

TABLE 1 (con t ’d)

Order ODONATAFam i ly L IBELLUL IDAEPan te la flavescens (Fabr icius , Jackson , 1 968 .

‘Tramee lacerata H agen , 1 86 1 .

Order BLATTODEAFam i ly BLATTELL IDAE

‘B latt el la l itu r ico ll is (Wa lker ,

‘Lupper ia notu lata (Stal ,Fam i ly BLATT IDAEMelenozos ter ia soror (Brunner ,

‘Neos tylopyga r hom bifo lia (Sto l l ,

‘Pen

‘

p laneta amer icana (L innaeus ,‘Per ip laneta au s t r eles iae (Fabr icius ,Fam i ly PYCNOSCEL IDAE

‘Pycnoscel is s u r inamens is (L innaeus ,

Order ORTH OPTERAFam i ly ACRID IDAEA iolopu s thelass in us temu lus (Fabr icius ,A io lop us temulus (Fabr icius , Townes , 1946 .

Fam i ly GRYLL IDAE‘Or nebiu s bimecu latus (Sh i rak i ,G ryllodes s igi llatu s (Walker , Townes , 1 946 .

‘Speonemobiu s sp . prob . t ig r inus (Saussur e,

Fam i ly TETT IGON I IDAEPhis is pa l lida (Walker , Townes , 1 946 .

Order DERMAPTERAFam i ly CARCINOPH ORIDAEEuborell ia annu l ipes (Lucas , Br indle, 1 972 .

Fam i ly CH ELISOCH IDAEChel isoches mor io (Fabr icius , Townes , 1946 .

Fam i ly LAB IDUR IDAELabidura r iper ie (Pal las , Townes , 1946 ; B r i ndle, 1 972 .

Or der PSOCOPTERAFam i ly CAECIL I IDAECaeci l ius cesar um Badonnel , 193 1 : Thor n ton et al . , 1972 .

Fam i ly ECTOPSOCIDAEEctop socus maindr on i Badonnel , 1935 : Thor nton et al . , 1 972 .

Fam i ly LEPIDOPSOCIDAECyp tOphen ie marginata Thor nton , Lee and Chui , 1972 : Thorn ton et al . , 1 972 .

Fam i ly PSOC IDAEPtycta angu lata Thor nton , Lee and Chui , 1 972 : Thor nton et al . , 1 972 .

Order MALLOPH AGAFam i ly MENOPONIDAEActom ithophi lu s bico lor (P iaget ,Actom i thophi lus cer u leus (T immer rnann , Woodbury , 1 962 .

Actor n i thophi lus incisus (P iaget ,Actor n ithoph i lus och r aceus (N i tzs ch,Aus t r omenopon at r ofu lvum (P iaget ,Colpocephalum angu lat icep s Piaget , 1 880Fam i ly PH ILOPTER IDAEH al ipeu r us m i r abi lis Thompson , 1940 .

Pect inopygu s gr ec i licor n is (Piaget ,Quadr acep s bi r os t r is (Giebel ,Quadracep s hopkins i T immermann , 1 952 .

Quadraceps separates (Kel logg and Kuwana,

Seem unds son ia albemar lens is (Kel logg and Kuwana,

Seem unds son ia puellu le T immermann , 1965 .

Seemunds son ia r emo te T immermann , 195 1 .

Seemunds son ia s nyder i (Kel logg and Paine,

Tr abecu lus hexakon (Water ston ,

‘New Enewetak r ecord . (Th i s table cont inued on next page

150 SAMUELSON AND NISH IDA

TABLE 1 (con t ’d)

Order TH YSANOPTERAFam i ly PH LAEOTH RIPIDAEH ap lothr ip s gowdeyi (Frankl in ,

Order H ETEROPTERAFam i ly ANTHOCORIDAEPhysop leurella mundu la (Wh i te, Her r ing , 1 967 .

Fam i ly COREIDAEL iorhyss us hyal inus (Fabr icius , Gross , 1963 .

Fam i ly CYDNIDAEGeotomus pygrneeus (Dal las , Ruckes , 1963 .

Fam i ly GERR IDAEH alobates m icens (Eschscholtz , Cheng, 1 981 .

H ermetobates sp . : Cheng , 1977 .

Fam i ly LYGAE IDAE‘Germalus sp .

Nys ius p icipes U s inger , 1937 : U s inger , 1949 , 1951 ; Bar ber , 1958 .

Nys ius pu lchel lus Stal , 1 859 : U s inger , 1949, 1951 ; Bar ber , 1 958 .

Pachybrechius n igr iceps (Dallas , Bar ber , 1958,

Fam i ly MIR IDAECampylomma en iwetok Schuh , 1 984 : Schuh , 1 984 .

Campylomma mar shallens is U s inger , 1952 : Schuh , 1984 .

Tr igonotylus doher tyi (D is tan t, Carvalho , 1956 .

Fam i ly NABIDAENabis caps iform is Ger rnar , 1 839 : Gr oss , 1 963.

Fam i ly PENTATOMIDAEOechal ie schellenbergi i (Guer in-Ménevi lle,

Ruckes ,Oechel ie consocial is (Boisduval) : U s inger , 1952.

P letynopus melaoan thus (Boisduval , Ruckes , 1963 .

Or der HOMOPTERAFam i ly APH ID IDAEAphis gossyp i i G lover , 1 855 : Ess ig , 1956 .

Aphis creccivo ra Koch , 1 854 .

Aphis medicagin is Koch , 1854 : Ess ig, 1956 .

Fam i ly CICADELLIDAEBalclutha incisa (Mats umur a, L innav0ur i , 1975 .

Balclutha saltuella (Ki r schbaum , L innavour i , 1975 .

Exi t ianus fusooner vos us (Motschulsky,Ex i t ien us cap ico le Sta], 1 855 : L innavou r i , 1 960 (in part) .

Exit ian us p lebeius (Ki r kaldy, L innavour i , 1960.

Ex it ian us cap ico le Sta], 1 855 : L innavour i , 1 960 (in par t).Oros ius ar gen tatu s (Evans , L innavour i , 1975.

Nesaloha oen ton is Oman , 1943 : Oman , 1 943 .

Oros ius oan ton is (Oman , Ghaun', 1 966 .

Reci l ie affin is (Os born , L innavour i , 1975 .

Recil ia hoppon is (Mats um ur a , L innavour i , 1975 .

Super fam i ly COCCOIDEACerop las tes ci r r iped iform is Coms tock , 188 1 : Bear ds ley , 1966 .

Coccus hesper idum L innaeus , 1 758 : Bear ds ley, 1 966 .

Icerya par ches i Mas ke ll , 1878 : Bear ds ley , 1966 .

Lep idosaphes eseki i Takahas hi , 1939: Bear ds ley, 1966 .

Phenacoocu s solan i Fer r is , 1918 : Bear ds ley, 1 966 .

Pseudococcus m icr oadon id um Bear ds ley, 1966 : Beards ley , 1966 .

Sai ssetia ooffeae (Walker , Beards ley, 1966 .

Fami ly DELP HACIDAESogatella ko lophon (Ki rkaldy, Fennah , 1 971 .

Fam i ly DERBIDAELamen ia oal iginea Char on Fennah , 1 956 : Fennah , 1 956 , 1 971 .

Or der COLEOPTERAFam i ly ANTH ICIDAEA n thicu s vexeto r Wer ner , 1 965 : Wer ner , 1 965 .

‘N ew Enewetak reco rd . (T h is table cont inued on next page .)

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152 SAMUELSON AND NISH IDA

TABLE 1 (con t ’d)

Fam i ly SPH INGIDAE‘Agr ius con vo lvu li (L innaeus , R iotte, in pr epa rat ion .

Cephonodes armatus Rothsch i ld and Jordan , 1 903 : Townes , 1 946 .

‘Cephonodes p icus (Cr amer , R iotte, in prepar ation .

G nathothl ibus er otus (Cramer ,

Chr om is er otus (Cr amer , Townes , 1 946 .

Fam i ly TORTR ICIDAEAdoxophyes fescicu lena (Walker , Clarke, 1976 .

Order D IPTERAFam i ly AGROMYZIDAEOph iomyia co r nuta (de Meijere, Spencer , 1 963 .

Pseudonepomyza sp icate (Ma l loch , Spencer , 1 963 .

Fam i ly CAL L IPH OR IDAEChr ysomya megacephale (Fabr icius , James , 1 962 .

Phaen ic ia cup r ine (W iedemann , James , 1 962 .

Fam i ly CANACEIDAEN oc t icanece mar shallen s is W i r t h , 1 951 : W i r th , 1 951 .

Fam i ly CERATOPOGON IDAEDasyhelea esaki i Tokunaga, 1 940 : Tokunaga and Mu r achi , 1959 .

Dasyhelea flevescens Tokunaga and Mu rachi , 1 959 : Tokunaga and

Mu r achi , 1 959 .

Dasyhelea flavibasel is Tokunaga, 1 940 : Tokunaga and .Mar achi , 1959.

Dasyhelea peli l iouen s is Tokunaga, 1 940 : Tokunaga and Mur achi , 1959 .

For cipom yia tu thi ll i Tokunaga, 1 959 : Tokunaga and Mu r achi , 1 959 .

Fam i ly CH IRONOMIDAEClun io tu th i ll i zTokunaga , 1 964 .

Pon tomyia natans Edwards , 1 926 : Cheng and H ashimoto , 1 978 .

°

Tenytar s us halophi lee Edwards , 1 926 .

°

Telmetogeton pus i l lum Edwards , 1 935 .

Thelessomye mar i t ime W i r t h , 1 947 : Tokunaga, 1964 .

Fam i ly CH LOROPIDAE‘Cadremo pal lida (Loew,

‘“

Gau rex"bico lor ipes (Ma lloch ,

Fam i ly EPH YDRIDAE‘Allot r ichoma alium Cr esson , 1 929 .

‘D iscocer ine mere Cresson , 1 939 .

‘H ecamede per s im i l is H endel , 1 91 3 .

‘H os t is guamens is Cres son , 1 945 .

‘Plecop s idelle cynocephala Ker tes z , 1 90 1 .

Fam i ly H IPPOBOSCIDAE°

O lfer s ia Sp in ifera (Leach ,Fam i ly MIL ICH IIDAE°

Desmometopa var ipa lp is Mal loch , 1 927 .

‘M i l ichiella lecteipenn is (Loew ,

Fam i ly MUSCIDAEA ther igona flavipalp is Mal loch , 1 928 : Snyder , 1 965 .

A ther igona (Acr itochaeta) or ien ta lis Schiner , 1 868 .

A ther igona exc ise (Thompson , Snyder , 1 965 .

Musca (Musca) domes t ica L innaeus , 1 758 : Snyder , 1965 .

Fam i ly PHOR IDAED ip lonevra (Dohr n iphora) cor nuta (B igot , Beyer , 1 967 .

Megesel ia (Megasel ie) s celar is (Loew , Beyer , 1 967 .

Fam i ly SARCOPH AGIDAEBoettcher isca ker nyi (H ardy , Souza Lopes , 1 958 .

Parasar cophage (L iosar cophaga) m isere(Walker , Souza Lopes , 1958 .

Phytos ar cophage gr ess i tt i (H al l and Bohar t , Souza Lopes , 1 958 .

Fam i ly SCATOPSIDAE‘H ol0 p leg ie guamens is (Johannsen , 1 946) or near ?

Fam i ly SCIAR IDAEB r adys ie t r i t ici (Coqu i llet t , Steffan , 1 969 .

‘New Enewetak record . (Th is table cont inued on next page

INSECTS AND ALLIES (ARTHROPODA) 153

TAB LE 1 (con t ’d)

Fam i ly SCIAR IDAE (cont ’d)Corynop tere lat l

'

s tylete (H ardy , Steffan , 1 969 .

P las tosciere let ipons H ardy , 1 956 : Steffan , 1969 .

Fam i ly STRAT IOMYIDAEB r achycer a ven t r a lis Thomson , 1 869 : James , 1 962 .

Fam i ly SYRPH IDAEIshiodon scu teller is (Fabr icius , Sh i r aki , 1963 .

Fam i ly T IPUL IDAEL imon ia (D icranomyia) pect inungu is Tokunaga, 1 940 : A lexander , 1 972 .

Order H YMENOPTERAFam i ly ANTH OPH OR IDAEXy locopa sonor ina Sm ith , 1 879 .

Fam i ly BRACON IDAEZele sp . : Townes , 1 946 .

Fam i ly EULOPH IDAEH em ip ter senus sem ielbiclavus (Gi r ault , Yosh imoto and Ish i i , 1965 .

Fam i ly EVAN I IDAE‘Szep l igetelle ser icea (Cameron ,

Fam i ly FORMICIDAEMonomor ium phar aon is (L innaeus , Townes , 1 946 .

Fam i ly ICH NEUMON IDAEEchthromorpha egr es tor ie ins idiator (Sm ith , Townes , 1 958 .

Fam i ly MEGACH IL IDAEMegach i'le di l igens hedleyi Rainbow, 1 897: Krombe in , 1950 .

Megach i le (Eu t r l'cher eee) fu llaweyi Cockerel l , 1914 : Krombein , 1 950 .

Fam i ly SPH ECIDAE‘Chelybion bengalense (Dahlbom ,

P ison p unct if rons Shuckard, 1 838 : Krombein , 1949 .

So l ier ella peckham i (Ashmead ,So l ierella rohwer i (Br idwel l , Krombein , 1949 .

‘Tachysphex bengalens is Cameron, 1 889 .

Fam i ly VESPIDAE‘Odyner us sp .

Pechodyner us nas iden s (Latr ei l le, Townes , 1946 .

‘Po lis tes fu scatus eu r ifer (Saus sure,

‘Ropel idia marg inata (Lepelet ier ,

‘New Enewetak record .

expect fewer ind igenous insect species on Enewetak thanon , for in s tance, Ar no Ato l l . H owever , the introduct ion of

exot ic p lant and insect species on Enewetak compl icatesthe picture.

The mos t notab le aspect concer n ing the env i ronment atEnewetak is obv ious ly the man-made alterat ions and dis tu r

bances over man y par ts of the ato l l . These had extr emeimpacts on the ind igenous b iota and tended to becomemor e sever e as the sequence of human occupat ion p ro

g r essed on the ato l l . The adven t of M icr ones ians and the

developmen t of a nat ive cultu re were fo llowed by co lon i zat ion and by tr ade w i th non -M icr ones ian elemen ts . Thus , atleas t cer tain i s lets wer e affected by the changes assoc iatedw i th the ear l ier phases of human occupat ion , includ ing thees tabl ishment of copr a plan tat ion s . The later per iodsincluded cons tr uct ion of m i l i tar y bases and ai r fields , warfar e , cleanup operat ions fo l lowing warfare, and tes t ing of

nuclear dev ices and assoc iated act iv it ies . The devas tat ionby war far e that occur r ed on cer tain i s lets du r ing Wor ldWar I I was subsequen tl y magn i fied by appl icat ions of insec

t ic ides (Townes , 1 946) and c leanup operat ions in wh ichen t i r e i s lets wer e c leared by bulldoz ing . One of the ear lypr og r am s o f the U . S . m i l i tar y was to es tabl ish vegetat ivegrowth on the bar e sand to ho ld i t down (B r yan , 1 944 ;

per sonal commun icat ion ) .The pos twar tes t ing of nuclear dev ices accounted for

fu r ther and somet imes comp lete devas tation of cer taini s lets . Some s i tes were also bulldozed fo l low ing tes ts . A ll

these pos twar act iv i t ies and the tr aff ic of per sonnel andmater ials ,

main l y thr ough H awai i , faci l i tated the impor tat ion of exo t ic plants and insects . Many species of plan tswer e in ten t ional l y in tr oduced or r eintr oduced to Enewetakfrom nur ser y s tock v ia Kwajalein (Fosber g , Over 50

species of exot ic plan ts wer e r epo r ted for Enewetak dur ingthat per iod (St . John , 1960) mar k ing a dr amat ic increaseof exot ics over prev ious per iods when ,

for example, on ly24 exot ics were es t imated fo r a lus her par t of the Mar shal lIs lands (Ar no) in 1 850 (H athaway , The p lan ts insome measur e may ser ve as an ind i cat ion of the num ber sof insects and r elated g r oups that wer e also in troduced to

154 SAMUELSON AND NISH IDA

TABLE 2

Col lector s of In sects and Related Gr oups on Enewetak A to l l

N ame Mon th Year

A l len , R . Aug . 1 945

Bohar t , R Jan . 1 945

Bryan , E . H . , Jr . Aug . 1 944

Feb . 1 975

Bushman , J . B . Mar .

— May 1 962

Lamber son , P .

Mo r r ison , J .

Oak ley , R . G . May 1 946

Tuth i l l , L . D .

Wheeler , M . R .

Young , F . N .

Enewetak dur ing the same per iods . H ence , the ind igenous(includ ing endem i c) biota of Enewetak — the species thatrepr esen t the tr ue unadu l terated character of the

atol l — pr obably suffered profoundly through both phys icalimpacts on the env i ronment and bio log ical impacts ,includ ing com pet i t ion for space and r esou r ces fr om adven

t ive form s .

In v iew of the d is tur bances to Enewetak A to l l , i t seem s

r elevan t to note the t imes and places wher e species wer eco llected or obser ved , ther eby pr ov id ing a summar y of thef i r s t four decades of en tomo log ical r ecor ds fo r the ato l l .Th is in format ion is prov ided in the section on Co l lect ionReco r ds .

F aun a

Townes ( 1946) es t imated that a large dr y ato l l l i keEnewetak m ight have 500 spec ies of insects . Th i s es t imate

could be somewhat h igh . The ind igenous insect fauna of

Enewetak would be expected to be less than that of Ar noAto l l . The fauna of A r no was also es t imated at ar ound500 spec ies (U s inger and LaR iver s , and at leas t296 spec ies have alr eady been recor ded fo r that atol l(G ress i tt , Co le ( 1 949 , 1 951 ) lis ted 56 species of

insects col lected from B ik in i A to l l in 1947 . B ik in i l ies inthe same zone as Enewetak and would be expected to sup

por t a near ly iden t ica l fauna . Co le’

s sur vey was the on lyone conducted for B ik in i and

,un for tunatel y , i t occur r ed

after appl icat ions of DDT on the i s lets and after the fi r s tatom ic bom b tes ts there . These per tur bat ions mus t havehad sever e effects on the fauna and on Co le’

s r esults .

Co le ’

s l i s ts wer e adm i ttedly incomplete; however , 35

spec ies l i s ted by him , includ ing 1 2 spec ies of ants , haveno t yet been r epor ted for Enewetak . Mos t of these 35

species pr obably also occur o r have occur r ed on

Enewetak .

Enewetak , Run i tEnewetak?Enewetak , Enjebi , Japtan , Medr en

A lembel , Anan ij, B iji le, Enewetak ,Enjebi , lkur en , Jap tan , Lojwa,

Medr en , Run i t

Aej, Enewetak , Enjebi , lnedral ,Jap tan , Jin im i , Medren , M ijikadrek ,

R ibewon

Enewetak , lkur en , Jap tan , Medl enEnewetak , JaptanEnewetak , Jap tanJaptan

Enjebi

Enewetak , JaptanEnjebi , Run i t

Enewetak7

Enjebi

B iken , lku r en , Kidr inen , L OU], Lujor ,Mut

Aomon , Enjebi , Jap tan (and other swi th Townes ?)

Bokombako , Elugelab, Enjebi

AejEnewetakAomon , Enewetak , Enjebi , lku r en ,

Japtan

Anan ij, lku ren , Jap tan ,Medr en

Japtan , Medren

Nan”

Medr en

Jap tan

Enjebi , lku r en , Japtan , Medr en

Medr en

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1 56 SAMUELSON AND NISH IDA

TABLE 4

Taxa Endem i c to the Mar shal l Is lands

A r achn idaPseudoscorp ion idaGarypu s or natus Beier

— Gar yp idae (B ikin i )Xeno lp ium ocean icum latum Beier — Olp i idae (Lae , Ujae , U t i r ik)

In sectaH eteropteraR ip tor tus sai ler i U s inger

— Coreidae (Kwajalein)Campylomma en iwetok Schuh— M i r idae (Enewetak)

H om opteraUgyop s k inberg i megas Fennah

— Delphacidae (A i l ing lapalap ,

Jalu i t )

Co leopteraCeres ium r obu s tum Gres s itt— Cerambycidae (A r no)Ceres ium un ico lor mar shellum Gr ess i tt — Cer ambycidae (Majur o)Oops is mar shallens is Gress itt— Cer ambycidae (var ious ato l ls )Prosop lus hibisc i Gress i tt— Cerambycidae (Majur o , U t i r ik)Pr osop lu s major Gr ess itt— Cer ambycidae (Ar no)Melanoxen thus lar iver s i Van Zwaluwenbu r g

— Elater idae (Ar no)Simodactylus mar shallens is Oh i r a — Elater idae (Kwajalein )Char iotheca cos tata Kulzer — Tenebr ion idae (Ar no , Ai l ing lapalap)Tegalus angus tu s Kulzer — Tenebr ion idae (Ar no)

D ipteraNoct icanece mar shallen s is W i r th — Canaceidae

Dasyhelea n ig r is t igmate Tokunaga and Mu r achi

Ceratopogon idae (A i l inglapalap)Dasyhelea sp . No . 1 Tokunaga and Mu r achi — Ceratopogon idae(Ar no)For cipomyia tu thi lli Tokunaga— Cer atopogon idae (Enewetak)Clun io tu thi ll i Tokunaga— Ch i ronom idae (Enewetak)Cr icotop us sp . No . 1 Tokunaga— Ch i ronom idae (Kwajalein )Plas tosciere jalu lten s is Steffan — Sciar idae (Jalu i t )Scythropochroa t r isp inose Steffan — Sciar idae (Kwajalein )L imon ia beards leyi A lexander — T ipul idae (Namu , Namor ik)L imon ia sen t ifer a A lexander — T ipu l idae (Namu , Ar no)

H ymenopteraCi r r osp l

'

lo ideus fu llawayi Yosh imoto and Ish i i — Eu lophidae

(Ebon )

ato l ls wher e they ar e ind igenous in the cen tr al and eas ter nCaro l ines (Yap , T r uk , Ponape , Kos rae) . None of the

Mar shal ls or Gi lber ts ar e cor r espond ingl y close to a h ighis land . The h igh is lands of the Caro l ines are cen ter s of

endem i sm w i th r elat ivel y r ich faunas and may have beensour ces for man y of the forms now s tr ictly l im i ted to ato l ls .

Thus , the pr esence of i so lated h igh i s lands in the v icin i tyof ato l ls may have cons ider able influence on the nature of

an ato l l fauna .

Factor s influenc ing an ato ll fauna would include the

above, as we l l as the deg r ee of iso lat ion of the atol l , landarea , cl imate, fr equency of typhoons , effects of w ind and

sea cu r r en ts , and pr esence of m ig r ato r y bi r ds . F inal ly ,human act iv i t ies have cons ider able bear ing on the natur e

TABLE 5

D i s t r i but ion of In sects and

A l l ies En dem i c to the Mar sha l l Is lan ds

Ra l i k RatakCha i n Chai n B oth Si n g le

G r oup (Wes ter n ) (Eas ter n ) cha i ns ato l l

of an ato l l fauna; these affect faunal compos i t ion thr oughenv i r onmen tal al ter at ion and al low incr eased oppor tun i tiesfo r es tabl i shment of adven t ive spec ies . D i r ect ai r and sh ipp ing r outes between the Mar shal ls and H awai i have obv i

ou s ly perm i tted movemen ts of species between the two

is land gr oups w i th in the pas t 40 year s (Table Of the

adven t ive—

elements on Enewetak that also occur in H awai i ,we note that they r epr esent around 2896 (54 of 191 ) of

the ato l l ’s fauna . Fu r thermor e, when cons ider ing the

Enewetak adven t ives alone, we see that a h igher pr oport ion , 6696, also occu r s in H awai i (54 of H awai i , as anexamp le of a cen ter of commer ce in the Paci fic , has w i tnes sed a mar ked incr ease of es tabl ished adven t ives over

r ecen t decades (cf . Bear ds ley , 1 962 , and that i sdue lar gely to incr eased ai r tr aff ic .Gr ess i tt ( 1 954) s tated that ato l ls and low is lands in the

Caro l ines , Mar shal ls , and G i lber ts have r elat ively poor butun i form faunas . T h i s appear s so w i th in the Mar shal ls , atleas t at the gener ic level and fr equently at the specieslevel , when the adven t ive form s es tabl i shed on Kwajaleinand Enewetak A to lls are d i scoun ted .

Zoogeog r aphy

The geograph ical aff in i t ies of major gr oups r epr esen tedin the Mar s hal l Is lands , or otherw i se in M icr ones ia, are

on ly par t ly ascer tained at th i s t ime. Man y gr oups ar e not

yet s tud ied to the po in t wher e they can be analyzed defin it ively . T h i s i s also tr ue for the faunas of adjacen t r eg ions ;all have influences on the M icr ones ian fauna thr ough pas tor p resen t movemen ts of spec ies .

In the fo l low ing d i s cus s ion , the general geogr aph icald i s tr ibut ions ar e given for all spec ies r epor ted for the

Mar shal l Is lands to class i fy the poten t ial inven tor y and

sou r ces of the species for Enewetak Ato l l . The number s of

species coun ted for the Mar shal ls and Enewetak are pr ov ided at the beg inn ing of mos t of the fo l low ing sect ion s .

A lso , mos t major g roups occur r ing in M icr ones ia are sum

mar i zed to g ive some per spect ive to the Mar shal l Is landsfauna ; thi s inc ludes , when known , the number of spec ieso r subspec ies r epo r ted for M icrones ia and the per centage

INSECTS AND ALLIES (ARTH ROPODA) 157

TABLE 6 TABLE 7

D i s tr i bu ti on of Mar shal ls’

Fauna Res t r i cted to Taxa Repor t ed for the Mar shal l s and Res t r i ct edthe Car ol i nes (Cen tr al and Eas tem ‘

yMar shal Is - G i l ber ts to t he Cen t r al and Eas ter n Car ol ines (Excludi ngSector of the Pac i f i c O cean Palau )—Mar shal ls - G i lber ts Sector of the Pac i f i c Ocean

A r achn idaPseudosco rpion idaLechyt ia sakegem l l Mo r ikawa (CA ,

M Mar cus )Nes idiocher nes macu latus Beier (K ,

M)G r oupT Opi l iones

Zalmox is s o li tar ie (Roewer ) (P ,M)A r achn ida

In sectaPseudoscorpion idaOp i l iones

In secta

i i

Totals

‘Central and Easter n Caro l ines all ato l ls and h igh is lands ‘

CA , Caro l ine Ato l ls ; G , G i lber ts ; K , Kos r ae; M ,Mar shal ls ; P ,

E of Palau ; th is div i s ion i s the Feder ated States of M icr ones ia . Ponape; T , T ruk ; Y , Yap ,

TTable 4 l is ts taxa in co lum n 1 , Table 7 l ists taxa in

co lum n s 2 and 3 .

Psocopter aEctop socu s v i l losu s Thor nton and Wong (T , P , K , M)H eteroceeci l ius cempenu le Lee and Thor nton (P , CA , M ,

G)H em ipteraNys ius p icipes U s inger

— L ygaeidae (M , G Wake)Campylomma mar shallens is U s inger

— M i r idae (M , G Kanton )H omopter aMyndus ap ical is (Metcalfl— Cix i idae (CA , T ,

P, K ,M)

Myndus p r aecan us Fennah— Cix i idae (P , M)Pseudococcu s mar shal len s is Beards ley— Pseudococcidae (K ,

M)Pseudococcus m icr oedon idum Beards ley

fi Pseudococcidae

(T , P , M , G)Ugyop s super c i l iate Fennah

— Delphacidae (CA ,P

, K ,M)

F lecc ia dione Fennah— Der bidae (CA , K ,M)

Co leopter aPeremeopedus m in imus (B lai r ) — Cerambycidae (CA , T ,

P,M)

P rosop lu s a . at lan t icus B reun ing— Cerambycidae (P , M)P r osop lus xyOIOp u s (Kar sch)— Cerambycidae (Y , CA ,

M,G)

B r on t ispa chalybeipen n is (Zacher )— Chr ysomel idae (P , K ,M)

Scym n us uncinus Chapin — Coccinel l idae (M ,G)

Eobie new species Macnamara and Ar nett Oedemer idae(M , G Ocean)

Cr yphalomorphus nubi lu s Wood— Sco lyt idae (CA , K , M)U loma ler iver s i Kulzer — Tenebr ion idae (K , M)

LepidopteraNeg ia h ierat ica H ampson — Noctuidae (Y , P , M , G)

D ipteraDasyhelea f lev ibesal l

’

s Tokunaga— Ceratopogon idae (P? ,M)

Dasyhelea pell ivi t tae Tokunaga— Ceratopogon idae (K ,M)

Sm i tt le yepen s is Tokunaga— Ch i ronom idae (Y , T , CA ,M)

Aedes mar shallen s is Stone and Bohar t— Cul icidae (CA , K ,M

,G)

M lcr odr osoph i la er reto r Wheeler and Takada— Dr osoph i l idae(P , K , M)

Scholas tes car o l inens is Ender lein — Platys tomat idae (P , M)Cephalochrysa r ugu lose James

— Str at iomyidae (P , K ,M)

H ymenopteraPseudeuco i la g r es s itt i Yosh imoto — Cyn ipo idea (T ,

M)Parechalcu r a macu lata Watanabe— Euchar idae (CA ,

M)H em ip tar sen us car o linen s is Yosh imoto and Ish i i — Eu lophidae

(Y , CA , M , G)

1 58 SAMUELSON AND NISH IDA

TABLE 8

Pr opo r t ion s of Indigenou s and Adven t i ve Spec ies forG roups Rep r esen ted on Enewetak A tol l

Indig Al so

Total ch ous , foundspecies i n cludi ng Adven . i n

Gr oup r epor ted endem i c ti ve H awai i

‘For ty-one ar e indigenous to Enewetak ; 54 are adventive.

of those endemic to M icrones ia . These f igur es w i l l cer tain l ychange as s tud ies con t inue on th is and adjacen t faunas .

ARACH N IDA

The M icr ones ian arachn id fauna is par t ial l y s tud ied ,though the spider s and m i tes r emain incompletel y su r

veyed. The Amblypyg i jus t enter the M icr ones ian ar ea

at Palau , w i th one Papuan — Malayan spec ies repor ted ,Char on g r ay

'

i Ger vai s (Takash ima,1 950 )

Scor p ion ida

Mar shal ls 2, Enewetak l — The H o rm u r us we l is t forEnewetak is w idely di s tr ibuted in the Pac i fic and alsooccu r s fr om Ch ina to Aus tral ia . The tr op icopo l i tanIsomet r us macu latus (DeGeer ) i s r epo r ted fo r the Mar shal ls ,G i lber ts , Palau ; i t is common ly found in dwel lings but i s not yet r epor ted for Enewetak . M icr ones ianfauna, s tud ied by Chap in compr i ses thr ee spec ies .

A second H orm u r u s of main l y Aus tr al ian — Melanes ian di s t r ibu t ion r eaches Palau .

Pseudos cor p ion idaMar shalls 1 0 , Enewetak 3 — Seven of the 1 0 pseudo

scorp ions r epor ted for the Mar shal ls ar e r es tr icted to

M icr ones ia. Two of these are endem ic to the Mar shal ls but

A r aneae

Acar i

Many add i t ional r ecor ds ar e expected for the Mar shal ls ;subor der s ar e tr eated ‘below .

Gamas ida (Mesos t igmata) : Mar shal ls 4 , Enewetakl — Lae laps ech l

‘

dn ina Ber lese and L . n u ttall i H i r s t arer epor ted for the Mar shal ls but no t Enewetak ; both are

as soc iated w i th Rettus and ar e cosmopo l i tan or near ly so .

The O r n i thonys sus we l is t for Enewetak is not ident i fied ,but the on ly spec ies thus far r epor ted for M icr ones ia is the

ar e not repor ted for Enewetak ; they ar e Garypus or natus

Beier (B ik in i ) and Xenolp ium ocean icum latum (Beier ) (Lae,Ujae, U t i r ik) . The Lechyt ia we l is t for Enewetak is re

s t r icted to ato l ls or low is lands, including the Carol ines(U l i th i ) and Mar cus ls land . Nes idiochem es maculatus Beieri s res tr icted to the eas tern Carol ines (Kos rae) and

Mar shal ls and i s not yet r epor ted for Enewetak . Threespecies ar e ei ther w idespread in M icrones ia or at leas tr ecor ded fr om d is tant local i ties. One of these i s the

H ap locher nes we l i s t for Enewetak . The thr ee extraM icrones ian taxa compr ise Or etem nus samoenus wher ton i

Chamber lin r epor ted for Enewetak , wes tern Car o lines , and

Ok inawa; the nom inate O . s . samoen us Beier of main lyPo l ynes ian di s tr ibut ion ; and a species of Geogarypus alsooccu r r ing on Taiwan . M icrones ian fauna, stud ied by Beier

compr i ses 45 spec ies or“

subspecies of wh ich all

ar e con fined to the Pac i fic bas in .

Op i l iones

Mar shal ls 1 , Enewetak 0 -On ly one species , Zalmox isso l i tar ia (Roewer ) , i s reco r ded for the Mar shal ls ; i ts dis t r ibu t ion i s res tr icted to the eas ter n Caro l ines (Ponape) andMar shal ls (Jalu i t ) . M icr ones ian fauna, s tud ied by Goodn ightand Goodn ight compr i ses s ix endem ic spec ies infive gener a . One genus , Par esamoe , is endemic to Ponapeand i s r elated to a Samoan genus and also to another inthe Seychel les ; the r emain ing four genera have repr esentat ives in the Malayan subreg ion .

Mar shalls 1 9 , Enewetak 1 2— Spider s repo r ted for theMar shal ls ar e w idespr ead (9 spec ies ), d is tr ibuted in the

Malayan subr eg ion and Paci fic confined to Ocean iaor undeterm ined Of the spec ies we l is t for Enewetak ,the A r aneus , Ch i racan th ium

, F laci llu la, H eter opoda,

Latr odect us , Smer ingop us , and Vi t ia ar e Widespread ; theMas ter ia, two

.

Opopaea, and Scytodes are con fined to

Ocean ia ; and the E r igone i s of unknown d is tr ibut ion , as i t

i s un iden t i f ied . At leas t f ive of the 1 2 sp ider s r epor ted forEnewetak also occur in H awai i , a probable avenue for thein troduct ion of some of the adven t ive spec ies . Roewer

( 1 963 ) tr eated seven fam i l ies of M icr ones ian spider s totaling 1 3 species , 1 0 of wh ich are res tr icted to M icr ones ia ;eight of the spec ies we tr eat for Enewetak ar e not includedin Roewer

’

s tr eatmen t .

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1 60 SAMUELSON AND NISH IDA

B lat todeaMar shal ls 9 , Enewetak 7— A ll the cockr oaches

r epor ted for the Mar shal ls are w idespr ead in the Pac i f icand ei ther include at leas t the Or ien tal r egion (4 species )or occu r ar ound the wor ld The M icr ones ian fauna hasnot been sur veyed , but seven species fr om Enewetak ar epr esent in B ishop Museum. Of these, the B latte l la,

L uppar ia, and Melenozos ter ia we l is t ar e Or iental ; theNeos tylopyge and Pycnoscel is ar e O r iental now spr eadthr ough the tr opics ; and both Per ip laneta are Afr ican ,

w i thone spec ies now tr op icopo l i tan {aus t r ales iae} and one now

cosmopo l i tan (amer icana) . A ll seven repor ted for Enewetakinclude H awai i in thei r d is tr ibut ions . Two add i t ionalspec ies repor ted for the Marshal ls are the cosmopol i tanB lattel la german ice (L innaeus ) and the Or ien talMelenozos ter ia n itida (B r unner ) .

Man todeaMar shal ls 1 , Enewetak 0— On ly one Melanes ian

mant id , Or thodera bu rmeis ter i Wood-Mason , reaches the

Mar shal ls but i t i s not r epor ted for Enewetak . Beier ( 1972)r ev iewed the M icr ones ian fauna, wh ich compr ises five spe

cies . The on l y mant id endem ic to M icrones ia i s from Palau ,and i t has Phi l ipp ine aff in i t ies .

Phasm a todeaMar shal ls 1 , Enewetak O— On l y one species ,

Megacr an ia betes i K i r by , is r epor ted for the Mar shal ls butnot Enewetak ; i t is d is tr ibuted thr ough Melanes ia,

Indones ia, and the Ph i l ippines . The Pac i fic walk ings t ickswer e general ly su r veyed by Nakata At leas t n inespecies ar e r epor ted for M icrones ia, includ ing four endem icto the Car ol ines and one to the Mar ianas , but the gr oupthere remain s poor l y co l lected .

O r thop ter aMar shal ls 1 0 , Enewetak 5— The species reach ing the

Mar shal ls are ci r cum tr opical w idespread in the Old

Wo r ld Or ien tal Indo-Aus tr al ian or res tr icted toOcean ia Although the M icrones ian fauna has not beensu r veyed , five species ar e known from Enewetak , ei therr epor ted ear l ier (Townes , 1946 ; Sugerman ,

1 972a) or

p resen t in B i shop Museum . Of the five we l i s t_

for

Enewetak , the A io lop us i s an acr id id of Indo -Aus tral ian di st r ibu t ion ; the Ph is is is a tett igon i id also found in Po l ynes ia(Samoa , Tah i t i ) ; and there are thr ee gryll ids : the G ryllodesi s c i r cum tropical , the Or nebius is Or iental , and the

Speonem obius i s Ocean ian . The G ryllodes and Or nebius

repor ted fo r Enewetak ar e also found in H awai i . The follow ing ar e repor ted fo r the Mar shal ls but ar e not

def in i tely r eco r ded fo r Enewetak — Acr id idae: Locus taden ica (L innaeus) , L . m ig rator ia man i lens is (Mayne), Oxyahy la in tr icate (Stal) ; G r yllacr ididae : G ryllacr is sp . near

aur an t iaca B r unner von Wat tenwyl ; and Tett igon i idae:Phaner op tera fu r cifera Stal .

Isop ter aMar shal ls 6 , Enewetak O— Three of the term i tes

known from the Mar shal ls ar e ident i fied : Kalotermesr ependus H i l l is r es tr icted to Ocean ia, Cop totermes

formosan us Sh i r ak i is an Or iental species spr ead into the

Pac i fic and elsewher e, and Cr yp to termes br evis (Walker ) istr op icopo l i tan (Cole, 1 951 ; Sugerman , 1 972a, 1972b) .

The las t two are also r epor ted for H awai i . The threeun ident i fied term i tes appear to r epresent G lyp totermes ,Nas u t itermes , and P r or hinotermes . The M icr ones ian faunahas no t been monogr aphed .

Emb iop ter aMar shal ls 1

,Enewetak 0 — Ol igo toma (Apos thon ia)

m icr ones iae Ross , descr ibed from the Mar ianas , is alsor epor ted for the Mar shal ls (Jalu i t ) and G i lber ts (Butar i tar i) .I t and a r elated spec ies found in the Car o l ines and

Po l ynes ia, O . (A) ocean ica Ross , along w i th an

undes cr ibed member of the subgenus from Ponape, all pos

s ibly have Ph i l ipp ines aff in i ties . M icrones ian fauna,

rev iewed by Ross compr ises five species of wh ichtwo appear to be endem ic to M icr ones ia . Two Or ientalspec ies that have spread to other trop ical ar eas also r eachM icrones ia.

Psocop ter a

Dermap ter aMar shal ls 7 ,

Enewetak 3 — All of the earw igs r epo r tedfo r the Mar shal ls ar e cosmopo l i tan spec ies . All but one of

the seven are r epor ted for Kwajalein . Three of the seven

compr ise the total earw ig fauna for the Car ol ine atol ls , andthe gr eater number in the Mar shalls , par t icular ly Kwajalein ,

i s l ikely a r esult of heav ier in ter nat ional traff ic . At leas tfou r of the seven also occu r in H awai i , including the threethat we l is t for Enewetak . M icr ones ian fauna, treated byBr indle compr ises 24 species , w i th endem ismabout 4696.

Mar shalls 1 6 , Enewetak 4 — All of these spec ies range

beyond the Mar shal ls , w i th seven of the 1 6 r es tr icted toM icr ones ia, two extend ing on l y to H awai i , thr ee r es tr ictedto Oceania , one also in the Or iental r egion , and threew idespr ead . Of the fou r psoc ids r epor ted for Enewetak ,one is con fined to M icrones ia, one to Ocean ia , and two are

w idespr ead . On l y the Caeci l ius and Ectop socus of the fourwe l is t are also repo r ted for H awai i . M icr ones ian fauna,

r ev iewed by Thor n ton et al . compr ises 90 species .

Endem ism in the fauna is about 4496, w i th 1 9 endem icspec ies in the Mar ianas , 1 9 more in the Caro l ines , and twoin the Bon in s . The number of endem ics i s d im in ished inthe eas ter n par t of the Caro l ines , w i th three on Ponapeand one on Kos rae. Psocids ar e fai r l y well r epresen ted onCar o l ine ato l ls , w i th one of the n ine spec ies there endemic;f ive of the n ine also occur in the Mar s hal ls .

INSECTS AND ALLIES (ARTH ROPODA)

Mal lophaga

Mar shal ls 22 , Enewetak 1 6— Col lections fr om 1 1 av ianspecies wer e made on Enewetak (Elbel , per sonal commun ication) , and at leas t 1 6 species of chew ing l ice are now

known from the ato l l . These are main ly fr om col lect ionsmade by J . B . Bushman and cowor ker s , subsequen tly s tudied by R . E . Elbel et al . Amer son and Emer son (1971 )repor ted mos t of the same species for the central SouthPaci fic . Ear l ier , Woodbury (1 962) repor ted Actor n ithophi lus cer u leus (T immerman) for Enewetak . Six add itional chew ing l ice r eco r ded for the Mar shal ls , but notEnewetak , ar e included in the Nat ional Museum of NaturalH is tor y col lect ion : Actor n i thophi lus l imosae Kellogg ,

A. umbr in us Burmeis ter , E idmann iella a lbescen s Piaget ,Menopon gal l inae L innaeus , Tr inoton quer quedu lee (L innaeus ) , and Pect inopygus s u lee (Rudow) . Mal lophaga ar e

cjoincident w i th thei r av ian hos ts , and all of the hos t spedies sampled on Enewetak r ange beyond the Mar shal ls ,wi th mos t being w idespr ead over par ts of the Pac i f ic . Th i rteen of the 1 6 spec ies l is ted for Enewetak ar e alsorepor ted fo r H awai i .Anop lu r aMar shal ls 3 ,

Enewetak O— Two l ice r epor ted for theMar shalls are cosmopo l i tan and as soc iated w i th man :

Pedicu lus humen is (L innaeus ) and P th i r us pubis (L innaeus ) .Another spec ies , H op lop leu r a pacif ica Ew ing ,

is assoc iatedwi th Rattus and i s w idespr ead in the Paci fic ; i t is alsorepor ted for the Malayan subr eg ion , Madagascar , Car ibbean , and southeas ter n U n i ted States . M icr ones ian faunacompr i ses s ix species wh i ch ar e mos tl y w idespr ead and

cbincident w i th thei r hos ts ; they are mos tly as soc iated w i thman or domes t ic an imals (Fer r is , 1 959 ; W i lson ,

Th ys anop ter aMar shal ls 2 , Enewetak 1 — These spec ies are ei ther

cosmopo l i tan , the H ap lothr ip s we l i s t for Enewetak , or

Or iental , Taen iothr ip s vitt icor n is (Kar ny) ; the latter i s

r epor ted for Kwajalein by Sugerman (1 972a) . Both are

also r epor ted for H awai i . M icr ones ian fauna remainsincomp letely su r veyed . Kur osawa (1940) tr eated 1 4

species based lar gel y on T . Esaki’

s col lect ions ; at that t ime

no thr ips wer e r epor ted for the Mar shal ls .

H eter op ter aMany of the lar ger fam i l ies r epr esen ted in M icrones ia

have been s tud ied . Among the gr oups r emain ing to be

repor ted upon are the aquat ic and sem iaquat ic fam i l iesoccur r ing in fr esh water ; however , they ar e pr obablylargel y absent on iso lated ato l ls . Fam i l ies tr eated forM icrones ia but lack ing r ecor ds fo r the Mar shal ls includeAcan thosom idae (Ruckes , En icocephal idae (U s inger

and Wygodz insky, Neididae (Gr oss , Platasp i

dae (Ruckes , Saldidae (D rake,and T ing idae

(D r ake, As w i th th is and mos t of the fo l low inginsect or der s , each fam i ly r epr esen ted in the Mar shal ls isr ev iewed

, and when ther e is a r elevan t tr eat i se for thegr oup , the author i s c i ted .

16 1

An thocor idae (H er r ing , Mar shal ls 3 , Enewetak1 — These ar e spec ies of Pac i fic or Eth iop ian — New Wor ldd is tr ibut ion , w i th one r es tr icted to M icrones ia . All thr eear e w idespr ead in M icrones ia. The Physop leu r ella we l i s tfo r Enewetak also occur s in H awai i . M icr ones ian faunacompr i ses 20 spec ies , w ith about 7096 endem i sm .

Ar ad idae (Matsuda and U s inger , Mar shal ls 1 ,

Enewetak 0 — Ch ies top lon ia pygmaea Ch ina,descr ibed

fr om Samoa, i s also repo r ted for Mar shal ls and Car ol ines .

Thr ee species includ ing the one r epor ted fo r the Mar shal lsalso occu r on Caro l ine ato l ls ; all thr ee are w idespr ead inM icr ones ia. M icr ones ian fauna compr i ses 40 spec ies , w i thendem ism about 9396.

Cim icidae (U s inger and Fer r is , Mar shal ls 1 ,

Enewetak O— The bed bug , Cimex hem ip ter us (Fabr ic ius ),i s a tr op icopo l i tan spec ies gener al ly d is tr ibuted in

M icrones ia . It is r epor ted for the Mar shal ls but not fo rEnewetak .

Cor eidae (Gr oss and Schaffner , Mar shal ls 3 ,

Enewetak 1 — T hese are spec ies ei ther endem ic or of

Melanes ian or cosmopo l i tan d is tr ibut ion . The endem icspec ies , R ip to r tus sai ler i U s inger , is known on ly fr omKwajalein . The Melanes ian species , Lep tocor is is o late (D istan t) , occu r s thr oughout the Mar shal ls but is not r epor tede lsewher e in M icrones ia . M icrones ian fauna compr ises 1 5spec ies , w i th about 1 396 endem ism .

Cydn idae (Ruckes , Mar shalls 1 , Enewetakl — G eotom us pygmaeus (Dal las ) is a spec ies of

Malayan — Ocean ian d i s tr ibut ion ,including H awai i . On ly one

“

other spec ies , fr om Guam , is r epor ted for M icr ones ia .

Ger r idae (Cheng , Mar shal ls 2, Enewetak2— Two mar ine species wer e r ecen tly co l lected fr omEnewetak . H alobates m icens (Eschscholtz) is common lyfound on the open ocean ; i t is ci r cum tropi ca l in d is tr ibut ionand i s the on ly member of the genus also occur r ing in theAt lan t ic . The H ermatobetes remains un iden t i fied ; i t is

apparen t ly undescr ibed and of l im i ted d i s tr ibut ion in the

Paci fic .L ygaeidae (Bar ber , Mar shal ls 7 , Enewetak

4 — T hese species ar e res tr icted to M icrones ia or ar e

of Malayan — Melanes ian Indo-Aus tr al ian — Pac i fic or

Or ien tal — Pac i fic (1 ) d i s tr ibut ion . Nys iu s p icipes U s inger ,

one of the species r epor ted for Enewetak , i s r es tr icted toato l ls and low i s lands . The Pechybr achius is the on l yspecies of fou r we l i s t for Enewetak that i s also r epo r tedfor H awai i ; i t is main ly of lndo -Aus tral ian — Paci fic dis t r ibut ion . M i crones ian fauna compr ises 43 species , w i th endem i sm aboutM i r idae (Car valho , 1 956 ; Schuh , Mar shal ls 7 ,

Enewetak 3 — These species ar e tr op icopo l i tan main l yof O ld Wor ld d i s tr ibut ion Or iental — Pac i ficAus tr al ia— Pac i fic o r l im i ted to Ocean ia Of the

Enewetak spec ies , Campylomma en iwetok Schuh i s notablebecause i t i s one of on ly thr ee insects thus far r epor ted asendem ic to the ato l l (the other Enewetak endem ics ar e

m idges : Cer atopogon idae and Ch i r onom idae) . Campylomma mar shallens is U s inger is r es tr icted to the Mar shal lsand G i lber ts , plus one reco r d for Kanton ,

Phoen ix Is lands ;the Tr igonos tylus we l i s t i s tr op icopo l i tan . M icrones ian

162 SAMUELSON AND NISH IDA

fauna compr ises 89 species , w i th endem ism about 8 196.

Four spec ies ar e r es tr icted to ato l ls ; all ar e in

Campylomma: the two ment ioned above, plus C . u l ithiens is

Car valho (Ul i th i ) and C . wakeana Schuh (Wake) .Nabidae (Gr oss , Mar shal ls 1 , Enewetak

l — N abis cap s ifo rm is Germar i s a cosmopol i tan spec ies .

M icr ones ian fauna compr ises four spec ies of wh ich one is

endem i c to the Car ol ines (Palau , Ponape) .Pentatom idae Mar shal ls 2, Enewetak

2— The P latynop us we l is t for Enewetak is r es tr icted toMelanes ia and M icrones ia, and the Oechalia we l is t is of

Aus tr al ian — Pac i fic d is tr ibut ion . M icr ones ian fauna com

pr ises 33 spec ies , w ith endem ism about 5596. All of thesespec ies include h i gh is lands in thei r d is tr ibut ions .

Reduv i idae (Wygodz in sky and U s inger ,

Mar shal ls 1 , Enewetak O— The on l y species r ecor ded fr omi so lated ato l ls i s the one r each ing the Mar shalls , P lo iar iain sol ida (B . Wh i te) , wh ich is Ph i l ipp ines - Pac i fic in

d is tr ibut ion . M icr ones ian fauna compr i ses 30 spec ies , w i thendem ism about 6796.

Vel i idae: Mar shal ls 1 , Enewetak O— H elovel ia mar i t ime

Ber gr o th is mar ine and main ly of Aus tr al ian — Indones iand i s tr ibut ion ; i t i s also r epor ted for B ik in i (Co le ,

H om opter aTh i s i s the on ly large insect or der r easonably s tuc ied in

i t s en t i r ety for M icr ones ia . Some of the fam i l ies ar e wel lr epr esen ted on M icrones ian is lands and have a h igh pr opor t ion of endem ic taxa. The fo l low ing fam i l ies weretr eated fo r M icrones ia but r ep r esen tat ives ar e not r epo r tedfor the Mar shal ls : A ley r odidae (Takahash i , Cer cop i

dae (Synave, Cicad idae (Esak i and M i yamoto ,

and Mem br acidae (Kato ,

Aph id idae (Ess ig , Mar shalls 4 , Enewetak 2— All

ar e w idespr ead or cosmopo l i tan species also r epor ted forH awai i . An add i t ional aph id r epor ted for the Mar shal ls isH ys teroneu r a seter iae (T homas ) (Sugerman , 1 972a) .

M icr ones ian fauna comp r i ses 1 6 spec ies , mos tlyw idespr ead ; endem ics include one spec ies and one sub

species o i Malayan or Or iental aff in i t ies .

Cicadell idae (L innavuor i , 1 960 ,Mar shal ls 1 0 ,

Enewetak 7— The spec ies we l i s t for Enewetak ar e

w idesp read in both hem ispher es of New Wor ld— Pac i ficd i s t r ibu t ion Or ien tal — Pac i fic Aus tr al ia— Pac i f icor l im i ted to Ocean ia At leas t one is r epor ted forH awai i : Reci l ie aff in is (Osbo r n ) . Ba lc lutha luc ide (But ler ) isr epor ted fo r Ar no and Ebon but not Enewetak ; i t i s of

New Wor ld— Paci f ic d i s tr ibut ion . Two undeterm ined Cicadell ids ar e r epor ted fo r Kwajalein by Sugerman (1 972a,

Empoasca, a genus well r epr esen ted in the Pac i ficand Car neocephale, a New Wor ld genus not included inL innavuor i

’

s treatmen t . M icr ones ian fauna compr ises 87species , w i th endem i sm about 6696.

Cixi idae (Fennah , 1 956 ,Mar shal ls 2 , Enewetak

O— Both M yndus ap icel is (Metcalf) and M . p raecanus Fen

nah are endem ic to the cen tral eas ter n Car ol ines and

Mar shal ls . M i cr ones ian fauna compr i ses 29 species , all

endem i c .

Cocco idea (Bear ds ley , 1966 , Mar shal ls 36 ,

Enewetak 7— These ar e mos tl y spec ies of w idespr ead dist r ibu t ion of wh ich 25 also occu r in H awai i . Seven speciesar e r es tr icted to M icr ones ia ; of these, five ar e w idespr eadin M icr ones ia and two ar e endem ic to the eas ter n Carol ines and Mar shal ls . Man y of these scale insects are com

mon on cult ivated plan ts and have become widely distr ibuted . All of the species r epor ted for Enewetak alsooccur on Kwajalein , and five of the seven also occur inH awai i . M icr ones ian fauna compr i ses 1 50 species , w i thendem ism about 2396.

Delphac idae (Fennah , 1 956 , Mar shal ls 9,

Enewetak 1 — These species are res tr icted to M icr ones iaM icr ones ia p lus H awai i or main l y to Ocean ia plus

con t inen ts and cont inental is lands bor der ing the wes ter nPac i fic The Sogatella we l i s t for Enewetak is repor tedfor Aus tr al ia , Ph i l ipp ines , and M icr ones ia. Of the r es tr ictedtaxa, Ugyop s kinberg i mayes Fennah i s endem ic to the

Mar shal ls and occu r s on both i s land chains and

U . s uper c i l iatu s Fennah is l im i ted to the eas ter n Carol inesand Mar shal ls . M icr ones ian fauna compr ises 63 spec iesw i th endem ism about Thr ee spec ies ar e endem ic toato l ls elsewhere: the G i lber ts , Wake, and Ocean Is land , ar ai sed ato l l .

Der bidae (Fennah , 1 956 , Mar s hal ls 3 ,Enewetak

1 — Spec ies o r subspecies r es tr icted to M icr ones ia of

wh ich the Lomen ie we l i s t for Enewetak i s endem ic to theMar shalls ; F lecc ia d ione Fennah i s l im i ted to Caro l ineato l ls , Kos r ae, and Mar shal ls ; and Swezeyia zephr us Fen

nah occu r s thr ough much of M icr ones ia. Lamen ia and

Swezeyia poss ibly have Ph i l ippine aff in i t ies , wh i le F leeciehas t ies to Melanes ia (F i j i ) . M icrones ian fauna compr i ses 56species , w i th endem i sm about 9596. Der bids are wellr epr esen ted on Caro l ine ato l ls , w i th 20 species of wh ichfour ar e endem ic to ato l ls ; another spec ies is endem ic tothe G i lber ts .

Ps yll idae (Tuth i ll , Mar shal ls 1 , EnewetakO— Mesohomotome h ibisci (F roggatt) r anges fr om Aus tr al iaand con t inen tal i s lands bo rder ing the wes ter n Paci fic toOcean ia wher e i t i s w idespr ead ; i t is r epor ted for

Kwajalein (Sugerman , 1 972a) . M icr ones ian faunacompr i ses 22 species , w i th endem ism about 6896. Two

add i t ional psy l l ids bes ides M . hibisci occu r on Car o l ineato l ls but ar e not r epor ted for the Mar shal ls .

Coleopter a

Mos t of the lar ge fam i l ies of beet les have been su r

veyed. The outs tand ing except ion i s the Cur cul ion idae or

weev i ls , a fam i ly wel l represen ted on Pac i fic i s lands , wi thon ly the Guam componen ts of the M icrones ian fauna ade

quately s tud ied . A few species ar e r epor ted for the

Mar s hal ls but not Enewetak . Fam i l ies monographed forM icr ones ia,

but no t yet r epor ted for the Mar shal ls , includethe Can thar idae (W i ttmer , Cr yptophag idae (M ich itaka Chfij6 , Cybocephal idae (Endr b

’

dy-Younga,

Endomych idae (Str ohecker , Lampy r idae(W ittmer , Malachi idae (W i ttmer , 1 958 ,

Pla

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164 SAMUELSON

spread ; on l y one species , Orphin us nes iot icus Beal , fr omGuam , i s endem ic .

Elater idae (Van Zwaluwenbur g ,1948 , 1957; Oh i r a,

Mar shal ls 1 1 , Enewetak 2— All identi fied spec iesar e of Ocean ian d is tr ibut ion . The two species we l i s t forEnewetak are ei ther repor t ed for New Br i tain , eas ter nCaro lines , and Mar shal ls (Simodactylus) or range throughM icr ones ia and Po lynes ia (Conoder us) . Two other speciesar e endemic to the Mar shal ls and ar e r es tr icted to s ingleato l ls : Melonoxon thus lar iver s i Van Zwaluwenbu rg fr omA r no and Simodactylus mar shallens is Oh i ra fr omKwajalein . Two un iden t i fied elater ids , Ampedus sp . and

Anchas tus sp . , ar e repor ted for Kwajalein (Sugerman ,

1 972a, M icr ones ian fauna compr ises 70 spec ies ,w i th endemi sm about 7196.

H is ter idae: Mar shalls 1 , Enewetak O— P lotysomo sp . i s

repor ted for Jalu i t (Schnee, The genus is wel lr epr esented in the Indo-Malayan area, but some speciesoccur in the New Wor ld .

H ydroph i l idae: Mar shalls 1 , Enewetak 0 — At leas t onespec ies , Doctylos ter num abdom ina le (Fabr ic ius ) , is repor tedfor the Mar shal ls (Sugerman , l 972a); i t i s poss ibly thesame as one or both of the Cyclonotum spp . repor ted bySchnee (1 904) for Jalu i t . The spec ies is w idely d i s tr ibutedin the New Wor ld and is es tabl ished in H awai i .Mycetophagidae (M ichi taka Chfijé , Mar shal ls 2 ,

Enewetak 1 — The Typhoeo we l is t for Enewetak is a

cosmopo l i tan species . L itorgus ves t i tus Shar p , r epor ted forAr no and Jalu i t , i s an Ocean ian species found throughoutM icr ones ia and par ts of Polynes ia. Both species are

r epor ted for H awai i . M icr ones ian fauna compr ises thr eespec ies , w i th none endem ic .

N i t idul idae (Gi l log ly , Mar shalls 1 4 , Enewetak6— These are mos t ly w idespread species , but one i s r e

s t r icted to M icr ones ia and Po lynes ia; s ix are cosmopo l i tan ;three are found on var ious cont inen ts ; one is also fr omAus tral ia; two are also from New Gu inea; and one occur sin Polynes ia, Ph i l ipp ines , and Sr i Lanka . M icr ones ian faunacompr ises 39 spec ies , w i th endem ism about 3396. None i s

res tr i cted to ato l ls ; cor r espondence of spec ies among di fferent atol l gr oups is fai r ly h igh w i th s ix of eight of theCarol ine ato l ls species and seven of the n ine Gi lber ts spe

cies also found in the Mar shal ls .

Oedemer idae (Macnamara and Ar nett , in pr eparat ion) :Mar shal ls 8 , Enewetak 6 — These species ar e more or lessof Malayan — Or iental d is tr ibut ion . Of these, on ly one is

r ather res tr icted , occur r ing on l y in the Mar shal ls , Gi lber ts ,and Ocean Is land . The remain ing seven tend to be

w idespr ead in M icr ones ia and include some that also occuron Melanes ian and Po l ynes ian is lands . M icr ones ian faunacompr i ses 1 9 spec ies , w i th mos t r es tr icted to M icr ones ia.

Pr opalt ic idae (John , 1960 ,Mar shal ls 1

,

Enewetak 1 — P r opolt icus insu lar is John is res tr icted to

M icrones ia. M icrones ian fauna compr ises eight spec ies ,w i th endem ism 6396.

Scar abaeidae (Car twr ight and Gor don , Mar shal ls6 , Enewetak O— These spec ies are of the subfam i l y Aphodi inae and ar e ei ther r es tr icted to M icr ones ia near l y

INSECTS AND ALLIES (ARTHROPODA)

of C . jolu i tono Kempny Ch r ysopa basa l is Walker isalso r epor ted for the Mar shal ls (A i linglapa lap) .

T r i chop ter a

The M icrones ian fauna is lar gely unsu r veyed , althougha few species are known to occur in the Mar ianas and

Caro l ines , includ ing r emote Kos r ae (Gr es s i tt ,

Lep idop ter a

The M icr ones ian fauna is poo r ly s tud ied , except theSph ing idae and cer tain m icr olep idopter an fam i l ies . Threefam i l ies that have been tr eated for M icrones ia, but havenot been found as yet in the Mar shalls , ar e Agonoxen idae(Clar ke, Chlidanot idae (Clar ke, and Oeco

phor idae (Clar ke,

Ar ct i idae: Mar shalls 2 , Enewetak l — The U tetheisa we

l is t for Enewetak r anges thr ough m uch of M icr ones ia and

the South Paci fic (Schnee, An undeterm ined l i thoss ine is r epor ted for Kwajalein (Sugerman , 1 972a) .

Cosmopteryg idae: Mar shal ls 1 , Enewetak 0 —An

undeterm ined Tr is sodor u s is r epor ted for the Mar shalls butw i thout ment ion of Enewetak (Townes , Th isspec ies m ight be T . honorer iella (Walshingham ), a

Pandanus leaf per fo r ator r ecor ded from Sr i Lanka and the

Pac i fic , includ ing H awai i .Danaidae: Mar shal ls 1 , Enewetak O— Danaus p lex ippus

(L innaeus ) i s r epor ted for the Mar shal ls (Jalu i t ) by Sei tzi t is also r epor ted by Townes (1 946) but w i thout

speci fic local i ty . Th is is a w idespr ead Nor th Amer icanspecies , also occur r ing in H awai i and elsewhere in the

Pac ific , includ ing Aus tral ia.

Gelech i idae: Mar shal ls 1 , Enewetak O— Stoeber hinus

tes teceus Butler , of Indones ian — Pac i f ic d is tr ibut ion ,

includ ing H awai i , is r epor ted for Kwajalein (Sugerman ,

1 972a) .

Geometr idae: Mar shal ls 1 , Enewetak O— An undeter

m ined Thelassodes i s r epor ted for Jalu i t by Seit z A

number of spec ies occur in the South Pac i fic .H esper i idae: Mar shalls 1 , Enewetak l — The Badem ia

we l i s t fo r Enewetak (Townes , 1 946) i s w idely d i s tr ibutedin the Paci fic , includ ing H awai i (as a recent intr oduct ion) ; i talso occur s in the Or ien tal and Aus tral ian reg ions .

L ycaen idae: Mar shal ls 1 , Enewetak O— The Lomp ides

r epor ted by Sugerman (1 979) is probably L . boet icus (L innaeus ) , a blue butterfly d is tr ibuted thr oughout the Old

Wor ld and Paci fic , includ ing H awai i .Noctu idae (Fukushima, 1947 , in par t) : Mar shal ls 13 ,

Enewetak 2— These spec ies ar e mos tly w idespr ead :An t icor s ia i r r or ata (Fabr ic ius) , Moc is fr ugal is (Fabr icius) ,Neg ia l in teole (G uenée) , Plus io cha lci tes (Esper ) , Spodop teraexempta (Walker ) , S. l itu re (Fabr ic ius ), and S . maur it ia(Bo i sduval) occur th r ough warmer reg ions of the Old

Wor ld and extend in to the Pac i fic ; Achaea janato L innaeus ,Anua co ronata (Fabr icius ) , Bocono man ifes tal is Walker ,

165

Celog r amma fes t ive (Donovan) , and Pletysen ta i llecte

(Walker ) are Or iental or lndo-Aus tr al ian - Pac i fic ; and Neg iah ierat ica H ampson i s r es tr icted to M icr ones ia. The las t i sdescr ibed from mater ial taken in the Mar shal ls and Gi lber tsand is also repor ted for the

hCar ol ines (Yap , Ponape) . Of

the two Spodoptera we l i st for Enewetak , one (exempta) iscommon in H awai i .

N ymphal idae: Mar shal ls 2 , Enewetak 24 Thesebutter fl ies ar e Indo-Aus tralian — Pac i fic (H ypol im nas) or

Aus tr al ian — Paci fic (P rec is) in di s tr ibut ion . The fo rmer , H .

bo l ino , exh ib i ts much var iat ion , and spec imens fr om the

Mar shal ls have r eceived the subspec i fic or in frasubspeci f icnames : incons tons , jolu ite, or pollescens . Clar k (1 951 )tr eated the butter fl ies of the Mar shal ls .

O leth r eut idae (Clar ke, Mar shal ls 3 ,Enewetak

O— These spec ies are ei ther Or iental extend ing in to the

Pac i fic and Ind ian Oceans (2) or are res tr icted to

M icr ones ia H eleono p . physolodes (Meyr ick) and

Str ather ot is leucosp is (Mey r ick) belong to the fi r s t catego r y .The res tr icted spec ies , Icel i ta'

monele Clar ke, i s r epor tedfor the souther n Mar ianas , Mar shal ls , and G i lber ts .

M icr ones ian fauna compr i ses '

58 spec ies , w i th endem ismabout 5596.

Py ral idae: Mar shal ls 7 ,Enewetak O— These ar e mos tly

w idespr ead spec ies : Maresm io tr apezol is Guenée occur sthr oughout the wor ld ; H ymen ia r ecu r val is (Fabr icius ) ,Nocoleie diemenol is G uenée, and Syngam ie f lor idol is (Zel ler )ar e of Old Wor ld— Pac i fic d is tr ibut ion ; H erpetogr ommol icar s isol is (Walker ) i s Or ien tal ; P i letocer o s ign ifer ol is (Wal

lengren ) r anges fr om Aus tr alia to Po l ynes ia (Mar quesas) ;and one is undeterm ined (D iaphen io) . The las t i s a largewor ld-d is tr ibuted genus of warmer r eg ions . All of thesespecies ar e r epor ted for Kwajalein by Sugerman (1972a ,

The H ymen ia appear s to be the on ly one of the

above that i s also r epor ted for H awai i .Sph ing idae (R iotte, in preparat ion) : Mar shalls 4 ,

Enewetak 4— We l i s t all of these spec ies for Enewetak ;they are ei ther r es tr icted to Ocean ia (1 ) or also occur on

con t inen tal i s lands or con t inents bor der ing the wes ter nPac i f ic The r es tr icted one is Cephonodes armatus R .

and J . M icr ones ian fauna compr ises 1 8 species , w i th noneendem ic . All seem to have O ld Wor ld aff in i t ies and man ytend to be w idespr ead in the Eas ter n H em isphere.

A lthough these insects are lar ge-bod ied , they are h ighlyvag i le and ar e able to reach ocean ic is lands far from con

t inen ts ; th is may help to explain the appar ent absence of

endem ic fo rms in M icrones ia .

T ineidae: Mar shal ls 2 , Enewetak 0 — Decedor chis

s imu lons (Butler ) and Opogono omoscopo (Mey r ick) r angefr om Afr ica to Aus tr al ia and in to the Pac i fic , includ ingH awai i . Both ar e based on s ing le r ecor ds for the Mar shal lsand are ten tat ively ident i fied (Sugerman ,

1 972a,

Tor tr icidae (Clar ke,Mar shal ls 1 , Enewetak

1 — Adoxophyes focicu lano (Walker ) main ly ranges thr oughthe Malayan and Melanes ian subr eg ion s , w i th the on lyr ecor ds for M icr ones ia being fr om Kos rae and Enewetak .

M icr ones ian fauna compr i ses seven spec ies , w i th endem i smabout 8696.

166 SAMUELSON

D ip ter a

Wh i le man y of the fam i l ies repr esen ted in M icrones iahave been su r veyed , a few r emain incompletel y s tudied atth is t ime; these include the As i l idae, Cecidomy i idae, Do l ichopodidae, Tach in idae, and var ious acalyptr ate fam i l ies .

Fam i l ies monog r aphed for M icrones ia, but not repr esen tedin the Mar shal ls , include the As tei idae (Sabr osky,

B ibion idae (H ar dy , Clus i idae (Steyskal and

Sasakawa, Coelop idae (H ar dy , Empididae(Quate,

Mycetoph i l idae (Colless , M icr opez i

dae (Aczel , N ycter ibi idae (Theodor , P ipuncu lidae (H ardy , Scenopin idae (H ar dy ,Simul i idae (Stone, St r eblidae (Maa, and

Taban idae (Stone,

Agromyz idae (Spencer , Mar shalls 2 , Enewetak2— Spec ies ei ther r anging fr om Melanes ia throughIndones ia to i s lands in the Ind ian Ocean (Ophiomy l'a) or

r es tr icted to the Pac i fic , Aus tr al ia, and Taiwan(Pseudonepomyza) . Both species are w idel y d is tr ibuted inM icr ones ia. M icr ones ian fauna compr ises 1 9 species , w i thendemi sm about 3296. Both species repor ted for the

Mar shal ls also occu r in the G i lber ts and Car o l ine atolls ;however , the Caro l ine ato l ls have three add i t ional speciesnot r ecor ded for the Mar shal ls ; no add i t ional species ar e

r ecor ded for the G i lber ts .

As i l idae : Mar shal ls 2 , Enewetak O— Two un ident i fiedspecies represent ing two gener a (Clinopogon and

Stenopogon ) ar e r epor ted for the Mar shal ls (Co le, 1 951 ;

Sugerman , 1 972a, 1 972b) . Both genera ar e r epr esen ted inthe Or iental r eg ion .

Cal l iphor idae (James , Mar shal ls 6 , Enewetak2— These ar e mos tl y spec ies of broad d is tr ibut ion , beingc i r cumtropical H o lar ct ic and r each ing many otherareas Indo-Aus tral ian rang ing fr om Afr ica to the

Paci fic or Malayan — Pac i fic The two speciesr epor ted for Enewetak ar e c i r cum tr op ical [Phoen iciacup r ine (W iedemann )] or lndo-Aus tral ian (Chr ysom yamegocepholo A ll of the above ar e w idespreadin M icrones ia except the H o lar ct ic Phoen icia ser icate(Meigen) . M icr ones ian fauna compr ises 1 1 species , w i thendem i sm low at around 996 (one endem ic species occur sin the Bon ins ) . The nonendem ic fauna is at leas t par tlyco inciden t w i th man in the Pac i fic . Over half of the

M icrones ian fauna is repr esented in the Mar shalls . A ll thr eespecies r epor ted on Caro l ine ato l ls and three of the fourspecies repor ted fo r the G i lber ts also occu r in the

Mar s hal ls .

Canaceidae (W i r th , Mar s hal ls 2 , Enewetak1 — These are ei ther endem ic to the Mar shal ls (Noct icanecemar shallens is W i r th) or r es tr icted to the Mar shal ls and

eas ter n Caro l ines (Kusaie) (P rocanoce townes i W i r th) . Thelatter i s not yet r epor ted for Enewetak . M icr ones ian faunacompr i ses two spec ies , w i th endem ism

Cer atopogon idae (Tokunaga and Mu rachi ,

Mar s hal ls 23 , Enewetak 5 — These species are mos tl y confined to M icrones ia includ ing three endem ic to the

Mar s hal ls ; fou r also occur on con t inen tal is lands bor der ing

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168 SAMUELSON AND NISH IDA

Sphaerocer idae (R ichards , Mar shalls 4 ,

Enewetak O— Tl iese are species r es tr icted to M icrones iaor otherw ise occur r ing in the Malayan subr eg ion or

cosmopo l i tan M icr ones ian fauna compr i ses 21 species ,w i th endem i sm about The s ing le spec ies r epor ted forCar o l ine ato l ls , as wel l as the two spec ies r epor ted for theG i lber ts , also occu r s in the Mar shalls .

St rat iomyidae, (James , Mar shal ls 4 , Enewetakl — These ar e ei ther is land spec ies r es tr icted to M icr ones iathose occu r r ing also in Melanes ia and i s lands of the

Indian Ocean o r w idespread spec ies of the tr op ics and

subtr op ics T he r es tr icted spec ies , Cephalochrysa

r ugu lose James , occur s in the eas ter n Car o l ines (Ponape,

Kos r ae) and the Mar shal ls . The w idespr ead species ,H ermet ic i l lucens (L innaeus ) , occur r ing in all warm r eg ions ,

is of Amer ican o r ig in . M icrones ian fauna compr i ses 24species , w ith endem ism about 6396. The thr ee speciesoccur r ing on Caro l ine atol ls and the two occur r ing in the

G i lber ts are also found in the Mar shal ls , mak ing the ato l ls t rat iomyid fauna fai r ly un i form .

Syr ph idae (Sh i rak i , Mar shal ls 3 , Enewetak1 — The Ischiodon we l is t for Enewetak is a w idespr eadIndo-Aus tr al ian spec ies occu r r ing all over M icrones ia . A lsorepo r ted for the Mar shal ls ar e Syr i tto or ien tal is Macquar t ,

r ang ing thr ough souther n As ia in to the Pac i fic , and

Er is ta lis call iphor o ides Sh i rak i , r es tr icted to the Caro l inesand Mar shal ls . M icr ones ian fauna compr ises 27 species ,w i th endem i sm about

Tach in idae: Mar shal ls 2 , Enewetak O— The Near ct icLespes io arch ipp ivor o (R i ley) pr obably r eached Kwajaleinv ia H awai i . An “un ident i fied Exor is to, also r epor ted forKwajalein , belongs to a genus wel l r epr esen ted in the OldWor ld . Both species ar e r epor ted by Sugerman ( 1 972a) .

Tephr it idae (H ar dy and Adach i , Mar shal ls 3 ,

Enewetak O— These species ar e ci r cum tr opical [D ioxynosor or cu lo r anging from Afr i ca thr ough As iainto Aus tr al ia and the Paci fic (Spathu l ina acr oleuco

Schiner ) , or Malayan — Pac i f ic in d is tr ibut ion (Docus

fr ouenfeldi Schiner ) . Of these, on ly D . sor orcu lo is r epor tedfor H awai i .

Tethin idae: Mar shal ls 1 , Enewetak O— Dasyr hicnoes so

insular is (Aldr ich) , descr ibed fr om H awai i , is r epo r ted forL ib. Is land (Sugerman , 1 972b) .

T ipul idae (A lexander , Mar shal ls 9 , Enewetakl — Except ing the . c i r cumtr opica l L imon ia um br ote (deMeijer e), all of these spec ies are confined to the Pac i fic ,w ith mos t res tr icted to M icr ones ia; two are endem ic to theMar shal ls . The mos t w idely d is tr ibuted of the pr eced ing ,

Styr ingomyie didyma Gr im shaw , r anging fr om New Gu ineato the Tuamotus '

and nor thwar d in the Pac i fic , was

descr ibed fr om H awai i . The two endem ic spec ies ar e

L im on ia beerds leyi‘

A lexander fr om Namu , K i l i , and

Namor ik and L . sen t ifer o A lexander from Namu and Ar no .

M icr ones ian fauna “ compr ises 71 spec ies , w i th endem ismabout 7296. One of the species fr om the Mar shalls alsooccu r s on Car o l ine ato l ls for wh ich five spec ies are

repor ted , includ ing two that ar e mor e or less endem ic (onemay also occur on Ponape) . The th ree species r epor ted forthe G i lber ts also occu r in the Mar shal ls .

Si phonap ter a(H opk ins , Mar shalls 1 , Enewetak O— On l y

Ctenocephal ides f . fe l is (Bouche) , the cat flea, is r epor tedthus far for the Mar shal ls but not Enewetak . M icrones ianfauna compr i ses s ix spec ies that ar e mos tly cosmopo l i tanspec ies associated w i th man , domes t ic mammals (par t icular ly Car n ivor a) , and rats . One spec ies , a bat flea, i s foundmain l y in the O r ien tal reg ion and r eaches Japan and

apparen tl y Guam (a s ing le recor d) .

H ymenop ter aThe M icrones ian fauna i s par t l y s tud ied but r emains

poor ly sur veyed in general . Gr oups that ar e especial ly inneed of s tudy include the B r acon idae,

Form icidae, and

many fam i l ies of Chalcido idea and Pr octot r upo idea. Famil ies essent ial ly monogr aphed for M icr ones ia but not

r epor ted for the Mar shal ls include Co l let idae (Krombein ,

Ency r t idae (Yosh imoto and Ish i i , H al ict idae(Kr ombein , Mymar idae (Doutt , Sco l i idae(Kr ombein ,

1 949 , Stephan idae (Townes ,and T r ichogrammat idae (Doutt ,

An thophor idae (Kr ombein , 1 950 ,in par t) : Mar shal ls 1 ,

Enewetak 1 X y locopa sonor ina Sm i th is a carpenter beeof Or ien tal aff in i t ies intr oduced to H awai i and elsewher e inthe Pac i fic . M icrones ian fauna compr ises s ix species ,includ ing two of Cer at ino endem ic to the Mar ianas or

Car o l ines .

Bethyl idae: Mar shal ls 1 , Enewetak 0 — An un ident i fiedSc leroderma is r epor ted for Kwajalein (Sugerman , 1 972a) .

The genus is widespr ead and occur s in H awai i .B racon idae: Mar shal ls 4, Enewetak 1 — Fou r gener a

are known for the Mar shal ls , and all ar e s ingle r ecor ds w i thnone of the species iden t i fied (Co le, 1 951 ; Sugerman ,

1 972a , 1 979; Townes , They ar e Chelonus ,Macr ocen tr us , M icr op li t is , and Ze le . On ly the las t isr epor ted for Enewetak . A ll are w idespr ead gener a whosespecies paras i ti ze Lepidoptera. Chelonus and Macrocen tr usar e known fr om H awai i . Watanabe ( 1945) s tud ied the

Esak i col lect ions fr om the Caro l ines and Mar ianas , including thr ee endem ic species of Chelonus and two of

Macr ocen tr us .

Chalc idae: Mar s halls 1 , Enewetak O— An un ident i fiedspecies , poss ibly Cholcis , is r epo r ted for Jalu i t (Schnee,

Cyn ip idae: Euco i l inae (Yosh imoto , Mar shal ls 3 ,

Enewetak 0 — These species , all in Ps eudeuco i la, ar e ei therres tr icted to M icrones ia (2) or also occur in Samoa and

H awai i M icr ones ian fauna compr ises 1 3 spec ies , w i thendem i sm about Two other species of Pseudeuco i la

r epor ted for Car ol ine atol ls ar e known from h igh is lands inthe Caro l ines and elsewher e but ar e not among thoserepor ted for the Mar s hal ls .

Euchar idae (Watanabe, Mar shal ls 1 , Enewetak0 — A s ingle spec ies , Paracholcu re macu lata Watanabe ,

i s

r es tr icted to the Mar s hal ls and Car o l ine ato l ls . M icrones ianfauna compr ises n ine spec ies , w i th all spec ies endem ic .

Eu lophidae (Yosh imoto and Ish i i , Mar shal ls 5,

Enewetak l — These ar e spec ies ei ther res tr icted to

INSECTS AND ALLIES (ARTHROPODA) 1 69

M icr ones ia (4) or also occur r ing in Aus tr al ia, Ph i l ipp ines ,and H awai i The las t is also r epor ted for Enewetak .One of the r es tr icted spec ies , Ci r r osp i lo ideus fu llewayi Y .

and l . , is r epo r ted on l y fr om Ebon . M icr ones ian faunacompr ises 35 spec ies , w i th endem i sm about 6096. On lythr ee of the 1 1 spec ies occu r r ing on Car o l ine ato lls and

one of the two species occur r ing in the G i lber ts are alsopr esen t in the Mar shalls , mak ing the atol l euloph id faunai r regular .

Evani idae (Townes , Mar shal ls 2, Enewetakl — These spec ies are ei ther ci r cumtropical [Evan iaappendiges ter (L innaeus )] or Malayan — Melanes ian and

extend ing far ther into the Pac i f ic , including Enewetak andH awai i (Szep t igetella) . M icr ones ian fauna compr ises thr eespecies , w i th none endem ic .Form icidae: Mar shalls 22 ,

Enewetak l — An ts fr omEnewetak are poor ly sampled , and the 1 3 spec ies l is ted byCole ( 1 949) for B ik in i should also occur there. The so lespec ies we l i s t for Enewetak , Monomor ium phar aon is (L innaeus) , is cosmopo l i tan . Spec ies r epo r ted fo r the Mar shal lsar e w idespr ead Or ien tal mor e or less res tr icted tothe Pac i fic or Aus tral ian At leas t 1 3 of these antsalso occur in H awai i , being main ly “ tr amp species thatar e w idely d is tr ibuted by commer ce. W i lson and Taylor(1 967) tr eated the Polynes ian fauna, wh ich includes man yspecies of pos s ible occur r ence in M icr ones ia .

Ichneumon idae (Townes , Mar shal ls 2 , Enewetak1 — The po lytyp ic Echthromorpha ag res tor ie Sweder us

r anges from Afr ica thr ough As ia into the Paci fic and

includes thr ee subspec ies endem ic to M icr ones ia, plus a

fur ther subspec ies (ins idiator ) of Indones ian — Melanes ian di st r ibu t ion that is r epor ted for Enewetak . The other species ,Tr athalo f lavoo r bi tol is (Camer on ) , is Or iental . Bo th speciesoccur in H awai i . M icr ones ian fauna compr ises 33 species ,w i th endemi sm pr obably less than 6096. On l y 1 296 ar e

r epo r ted for i so lated ato l ls , but none occur s exclus ively onato lls .

Megachi l idae (Kr ombein , Mar shal ls 3 , Enewetak2— Two species of Megachi le r epor ted for Enewetak are

ei ther of Ocean ian d i s tr ibut ion (hedleyi) or poss ibly of

Ph i l ippine or igin and are now found thr oughout M icr ones iaand H awai i (fu llewayi) . A th i r d species , M . umbr ipenn is

Sm i th , is Or iental and was not seen in Micr ones ia unti lafter Wor ld War I I ; i t is now repor ted for the Mar ianas andMar shal ls and also occur s in H awai i . M icrones ian faunacompr ises eight species , w i th endemi sm aboutScel ion idae: Mar shalls 2 , Enewetak O— Two un iden t i

fied species ar e r epor ted fo r Kwajalein (Sugerman ,

they r epr esen t Te lenomus and pos s ibly Calo teleio .

Sphecidae (Kr ombein , 1 949, Mar shal ls 1 1 ,

Enewetak 4 — The species from Enewetak ar e from con

t inental ar eas , pr obably r each ing Pac i fic is lands , includ ingH awai i , thr ough human act iv i t ies . Spec ies r epor ted for theMar shalls ar e widespr ead in the Old Wor ld Or ientalr eg ion New Wor ld or l im i ted to Paci fic i s landsSpec ies in the las t categor y all belong to P ison . N ine of 1 1

spec ies r each ing the Mar shal ls also occur in H awai i , including the fou r r epo r ted for Enewetak . It is pr obable that atleas t some of the sphecids gained en tr y to the Mar shal ls

v ia H awa i i . M icr ones ian fauna compr ises 30 spec ies , w i thendem ism aboutVespidae (Krombein , Mar shal ls 4 , Enewetak

4 — The spec ies of Pechodyner us and Po l is tes we l i s t forEnewetak ar e fr om Nor th Amer ica , in troduced to H awai iand Micr ones ia; the Ropol idie is Malayan . The d is tr ibutionof Odyner us sp . is undeterm ined , but ther e are thr eeendem ic congener s descr ibed fr om the Mar ianas . M icrones ian fauna compr i ses 1 2 spec ies , w i th endem i sm about5896.

COLLECTION RECORDS

In sect s and Al l ies of Enewetak A tol l

Taxa are l i s ted in the same or der as g iven in the

checkl i s t (Table Co l lect ion r ecor ds r epor ted are abbre

v iated and include, when known , the is let , month , year ,and su r name of co l lector . The speci fic is let was unclear insome cases , and these are l is ted as

“ ? Is let” or by'

-

, a

descr ipt ive phr ase (see fi r s t en tr y) . The name of the determ iner is included when the par t icular specimens ci ted werepr ev ious l y unr epor ted for Enewetak .

Class ARACH N IDAOrder SCORPION IDAFam i ly ISCH NURIDAEH ormu r us ous t reles iae Fabr icius

“

On ever y i s land v is ited” : May 1 946 , Townes .

Order PSEUDOSCORPION IDAFam i ly ATEMN IDAEOr etemnus samoenus wher ton i Chamber l inEnewetak Is let : Nov . 1 944 , Dybas and Edgar . Japtan Is let :Oct — Nov . 1 944 , Dybas .

Fam i ly CH ERNETIDAEH ap lochem es in s u lan us Beier

Japtan Is let : Nov . 1944 , Dybas .

Fam i ly CH TH ON IIDAELechyt ia sakegem i i Mor ikawaEnewetak Is let : Nov . 1 944 , Dybas Japtan Is let : Nov . 1 944 ,

Dybas .

Order ARANEAEFam i ly ARANEIDAEA r aneus theis (Walckenaer )Iku r en Is let : Sept . 1975 , Cheng . Medr en Is let : Sept .Cheng — det . by L . J . Pinter .

Fam i ly CLUB ION IDAEChi r ocan thium diver s um KochJaptan Is let : Sept . 1 975, Cheng . Medr en Is let : Sept .Cheng— det . by L . J . Pinter .

Fam i ly D IPLUR IDAEMas ter ia hi r su te KochEnewetak ? Is let : June 1 946 , Mor r ison .

Fam i ly EUSPARASSIDAEH eter opoda venetor ia (L innaeus )Enewetak Is let : Sept . 1 975 , Cheng— det by L . J . Pinter(determ ined from cas t exoskeleton) .

170 SAMUELSON AND NISH IDA

Fam i ly MICRYPHANTIDAEEr igone sp .

Enewetak Is let : Sept . 1 975, Cheng — det . by L . J . Pin ter .

Fam i ly OONOPIDAEOpopaea foveo late RoewerEnewetak ? Is let : Nov . 1 944 ,

Dybas .

Opopaea leno SumanEnewetak Is let : Sept . 1 975 , Cheng . Medren Is let : Sept .Cheng— det . by L . J . P inter .

Fam i ly PHOLCIDAESmer ingop us e longatus (V inson )Enewetak Is let : Sept . 1 975 , Cheng — det . by L . J . P inter .

Fam i ly SALTICIDAEF loci llu la m in u te (Ber land)Jap tan Is let : Sept . 1 975 , Cheng— det . by L . J . P inter .

Vit ia albipalp is Marp lesEnewetak Is let : Sept . 1 975 , Cheng— det . by L . J . P inter .

Fam i ly SICARIIDAEScytodes s t r iat ipes (Koch)Japi an Is let : Nov . 1 944 , Dybas ; July 1 946 , Townes .

Fam i ly TH ER ID IIDAELatr odectus geomet r icus (Koch)Enjebi Is let : Date? , Lamber son — det . by L . J . Pinter . ?Is let :May 1 946 , Townes .

Order ACAR IFam i ly ARGASIDAEOm ithodor os capens is NeumannAej, Anan ij, Boken , Jin im i , R ibewon , and other Is lets : Spr ing1 962 , Bushman or cowor ker s — det . by D . E . Johnson .

Fam i ly DERMANYSSIDAEOr n ithonys sus sp .

? Is let : 1 962 , Bushman or coworker s — det . by N . W i lson .

Fam i ly ERYTH RAEIDAEBaleu s t ium sp .

? Is let : May 1 946 , Townes .

Class D IPLOPODAOrder PROTEROSPERMOPH ORAFam i ly STRONGYLOSOMIDAEOx idus g raci lis (Koch)Enewetak Is let : Sept . 1 975 , Cheng det . by F . G . H owar th .

Class INSECTAOrder TH YSANURAFam i ly LEPISMAT IDAENot ident ified? Is let : May 1 946 , Townes .

O rder COLLEMBOLAFam i ly ENTOMOBRYIDAESei ra bipunctate (Packard)Enewetak Is let : Sept . 1 975 , Cheng— det . by P Bel l inger .

Fam i ly ISOTOMIDAEIsotom u r us t r icusp is Bo r ner

? Is let : Sept . 1 975 , Cheng— det . by P . Be l l inger .

O rder ODONATAFam i ly L IBELLUL IDAEPan tale flavescens (Fabr icius )Enjebi Is let : Date? ,

Jackson .

Tremea lacerata H agen Order PSOCOPTERAMedren Is let : Feb . 1 975 , B r yan —

s ight r ecord by E . H .

Fam i ly CAECIL IIDAEB r yan , Jr . Caeci l ius ces ar um Badonnel

Or der BLATTODEAFam i ly BLATTELL IDAEB lattel la l i tu r icoll is (Walker )Enjebi Is let : Jan . 1 95 1 , Osh i r o . Jap tan Is let : Aug . 1 956 ,

Tuth i ll— det . by S. Asah ina.

Luppar ie no tu lata (Stal)Bokombako Is let : Dec . 1950 , Osh i r o . Japtan Is let : Jan . 1 957,

Tuth i l l ; Feb . 1 957 , Tuth i l l— det . by S. Asahina.

Fam i ly BLATT IDAEMelenozos ter ia so r or (B runner )Anan ij Is let : Aug . 1 956 , Tuth i l l . Bokombako Is let : Dec . 1950 ,

Osh i r o— det . by S. Asahina. Enewetak Is let : Sept . 1 975,

Cheng— det . by G . M . N i sh ida .

N eos tylopyge rhombifo lia (Sto l l)Enewetak Is let : Sept . 1 975 , Cheng— det . by G . M . N ish ida .

Per ip laneta amer icana (L innaeus )Enewetak Is let : Sept . 1 975 , Cheng— det . by F . G . H owar th .Enjebi Is let : Jan . 195 1 , Osh i r o— det . by S. Asahina.

Per ip laneta ous treles iae (Fabr icius )Enjebi Is let : May 1 946 , Oak ley and Townes . Japtan Is let : Aug .

1956 , Tuth i l l ; Sept . 1 956 , Tuth i l l . Medr en Is let : Aug . 1 956 ,

Tuthi l l .Fam i ly PYCNOSCEL IDAEPycnoscel is s u r inamen s is (L innaeus )Anan ij Is let : Aug . 1 956 , Tuth i l l . Elugelab Is let : Jan . 1 951 ,

Osh i r o Enewetak Is let : Nov . 1 944 , Dybas . Enjebi Is let : May1 946 , Oak ley . Japtan Is let : Aug . 1 956 , Tuth i l l ; Jan . 1 957,

Tuth i l l— det . by S. Asahina. Medr en Is let : Sept . 1 975,

Cheng— det . by F . G . H owar th .O rder ORTH OPTERAFam i ly ACR ID IDAEA iolopus thelas s in us temulus (Fabr icius )Aomon Is let : May 1 946 , Townes . Enewetak Is let : May 1 946 ,

Townes ; Sept . 1 975 , Cheng . Japtan Is let : May 1 946 , Townes ;Sept . 1 975 , Cheng — det . by G . M . N ish ida.

Fam i ly GRYLL IDAEOr nebius bimocu letus (Sh i rak i )Enewetak Is let : Sept . 1 975 , Cheng— def . by F . G . H owar th .

G ryllodes s ig i llatus (Walker )Enewetak Is let : Sept . 1 975 , Cheng— det . by F . G . H owart h .

Speonemobius sp . pr ob . t igr inus (Saus sur e)Japtan Is let : Sept . 1 975, Cheng — det . by A . B . Gur ney .Fam i ly TET TIGON IIDAEPhis is pal l ida (Walker )

“

A ll i s lands o r ato l ls v is ited May 1 946 , Townes .

O rder DERMAPTERAFam i ly CARCINOPH ORIDAEEubor ell io onnu l ipes (Lucas )Enewetak Is let : Sept . 1 975 , Cheng— det . by G . M . N ish ida.

Enjebi Islet : May 1 946 , Townes and Oak ley [publ ished as

Nov . 1 947 , Dybas ; Aug . 1 956 , Tuth i l l .Fam i ly CH EL ISOCH IDAEChel isoches mo r io (Fabr icius ) :

“

on ever y i s land v is i ted” : May 1 946 , Townes .

Fam i ly LAB IDUR IDAELabidur a r ipar ie (Pal las ) .Enewetak Is let : Sept . 1 975 , Cheng— def . by G

:M . N ish ida.

Enjebi Is let : May 1 946 , Townes ; Jan . 195 1 , Osh i ro .

?Is let :May 1 946 , Townes ; Jan . 1 95 1 , Osh i r o .

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1 72L

SAMUELSON AN

Townes .? Is let : Jan . 1 945 , R . Bohar t ; May 1 946 , Oak ley .

Aph is croccivo ra KochEnjebi Is let : May 1 951 , Fosberg . Japi an Is let : May 1 946 ,

Oak ley:Fam i ly CICADELL IDAEBa lclutha incisa (Matsu

‘mura)

? Is let : Dec . 1 950 , Osh i r o .

Balclutha saltuella (K i r schbaum )Jap tan Is let : Aug . 1 958 , Tuth i l l .

Ex i t ien us fusconer vos us (Mot schu lsky)Aomon Is let : May 1 946 , Townes . Elugelab Is let : Jan . 1 951 ,

Osh i r o . lkur en Is let : May 1 946 , Townes .

Ex it ienus p lebe ius (Ki rkaldy)Anan ij Is let : Aug . 1 956 , Tuth i l l . Aomen Is let : May 1 946 ,

Townes . Elugelab Is let : Jan . 1 951 , Osh i ro. Enewetak Is let :Sept . 1975 , Cheng . lku r en Is let; May 1 946 , Townes ; Aug .

1 956 , Tuth i l l . Japtan Is let : Sept . 1 975 , Cheng . Medr en Is let :Aug . 1956 , Tuth i l l ; Sept . 1 975 , Cheng— det . by W . C . Gagné .

Or os ius argen tatus (Evans)Aomon Is let : May 1946 , Townes . Enewetak Is let : May 1946 ,

Townes . Japtan Is let : May 1946 , Oak ley ; Aug . 1 958 , Tuth i l l .Reci l l

'

o aff in is (Os bo r n )lku r en Is let : Aug . 1 956 , Tuth i l l .

Reci l io hoppon is (Matsumura)Japtan Is let : Aug . 1 958 , Tuth i l l .

Superfam i ly COCCOIDEACerop los tes ci r r ipedifo rm is Com stockMedr en Is let : Sept . 1957 , Tuth i l l .

Coccus hesper idum L innaeusEnjebi Is let : May 1 946 , Oak ley . Japtan Is let : May 1 946 ,

Townes .

Icer ya p u r chos i Maskel l? “

Nan” Is let : July 1 957 , Tuth i l l .

Lep idosaphes esekli TakahashiJapi an Is let : May 1 946 , Townes .

Phenococcus so lan i Fer r i sEnjebi Is let : May 1 946 , Townes .

Pseudococcus m icroedon idum Beards leyEnjebi Is let : May 1 946 , Townes .

Sais set io coffeoe (Walker )Medren Is let : Sept . 1 957 , Tuth i l l .Fam i ly DELPH ACIDAESogatella ko lophon (K i r kaldy)Japtan Is let : Aug . 1956 , Tuth i l l .Fam i ly DERBIDAELomen ie cal igi neo choron Fennahlku r en Is let : Aug . 1 956 , Tuth i l l . Japtan Is let : May 1 946 ,

Townes [publ ished as Aug . 1 956 , Tuth i l l . ? Is let : Nov .

1 944 , Edgar .

Order COLEOPTERAFam i ly ANTH ICIDAEAn thicus vexator Wer nerEnewetak Is let : Nov . 1 944 , Dybas . Japtan Is let : Nov . 1 944 ,

H agen ; Sept . 1 975 , Cheng — det . by G . A . Samuelson .

Fam i ly CARAB IDAEEgadr omo smer agdu la (Fabr icius )Enewetak Is let : Sept . 1975 , Cheng — det . by G . A . Samuelson .

Enjebi Is let : Jan . 1 95 1 , Osh i r o .

Fam i ly CERAMBYCIDAEP r osop lus at lan t icus ot lon t icus

A

B r eun ing

Elugelab Is let : Jan . 1 951 , Osh i r o .

Fam i ly CH RYSOMEL IDAEAphthono bicolo r ata Jacoby . ? Is let : Sept . 1 975 , Cheng — det . by

INSECTS AND ALLIES (ARTH ROPODA)

Fam i ly SCOLYT IDAECryphalomorphu s nubi lus Woodlku ren Is let : May 1 946 , Townes .

H ypo thenem us er uditus WestwoodJaptan Is let : Nov . 1 944 , Dybas .

Fam i ly TENEBRION IDAEGebien iella car inata (Eschscholtz)? Is let : Nov . 1944 , Edgar .

Conocephalum adp r es s iforme Kas zab

Enewetak Is let : Sept . 1 975, Cheng— det . A . Samuelson .

Order NEUROPTERAFam i ly CH RYSOPIDAECh r ysopa rambu r i SchneiderAomon

_

Is let : May 1 946 , Townes . Bokom bako Is let : Dec .

1 950 , Osh i ro . Jap tan Is let : May 1 946 , Townes .

Order LEPIDOPTERAFam i ly ARCT I IDAEU tetheisa p u lchello ides H ampsonOccur s on ever y is land v is ited May 1 946 , Townes .

Fam i ly H ESPER IIDAEBadem ia exclamat ion is (Fabr icius )? Is let : May 1 946 , Townes .

Fam i ly NOCTU IDAESpodopter a exempta (Walker )? Is let : May 1946 , Townes .

Spodoptera li ture (Fabr icius )“

Occur s on ever y is land v is ited May 1946 , Townes .

Fam i ly NYMPH AL IDAEH ypol imnas bo l ina (L innaeus )

“

On ever y is land v is ited": May 1 946 , Townes .

Precis vi ll ida (Fabr icius )Aomon Is let : May 1 946 , Townes . Enjebi Is let : May 1 946 ,

Townes . Medren Is let : May 1 945 , Young .

Fam i ly SPH INGIDAEAgr i

‘us convo lvu li (L innaeus )_

Enjebi Is let : Jan . 1951 , Os h i ro . Japtan Is let : Aug — Sept 1 956 ,

Tuth i l l . Medren Is let : Aug . 1 956 , Tuth i l l . ? Is let : Aug . 1 967 ,

Knudsen .

Cephonodes armatus Rothsch i ld and Jordan? Is let : May 1 946 , Townes .

Cephonodes p icus (Cramer )Enewetak Is let f May 1946 , Townes and Oak ley . Enjebi Is let :May 1946 , Townes ; May 1 946 , Townes and Oak ley ; Jan .

1 951 , Osh i r o .

Gnothothl l'

bus erotus (Cramer ) : ? Is let : May 1 946 , Townes .

Fam i ly TORTRICIDAEAdox0 phyes fescicu lena (Walker)Elugelab Is let : Jan . 1 951 , Osh i r o . Jap tan Is let : Aug . 1 956 ,

Tuth i l l ; Jan . 1 957 , Tuth i l l .Order D IPTERAFam i ly AGROMYZIDAEOph iomyio cor nuta (de Meijere)Enewetak Is let : Sept . 1 975 , Cheng Japtan Is let : May 1 954 ,

Townes [publ i shed as Aug . 1 956 , Tuth i l l? ; Jan — Feb .

1 957 , Tuth i l l ; Sept . 1 975, Cheng— def . by J . A . Tenor io .

Medr en Is let : Aug . 1 955 , Wheeler .

Pseudonepomyza sp icoto (Mal loch)? Is let : May 1 946 , Townes .

Fam i ly CALL IPHOR IDAEChr ysom ya megocephola (Fabr icius )Bokombako Is let : Dec . 1 950 , Osh i r o . Enjebi Is let : Dec . 1950 ,

Osh i ro .

1 73

Phoen icia cup r ine (W iedemann )Enewetak Is let : May 1 946 , Townes ; Jan . 1 951 , Oshi ro . Enjebi

Is let : May 1 946 , Oak ley .Fam i ly CANACEIDAEN oct icanece mar shallens is W i r thAomon Is let : May 1 946 , Townes . Enewetak Is let : Sept . 1 975 ,

Cheng . Jap tan Is let : May 1 946 , Towhes ; Sept . 1 975,

Cheng— det . by W . W . W i r th .Fam i ly CERATOPOGON IDAEDasyhelea esek l i TokunagaJap tan Is let : Aug — Sept . 1 956 , Tuth i l l ; Jan — Feb . 1 957,

Tuth i ll ;Sept . 1975 , Cheng— det . by W . W . W i r th .

Dasyhelea f lavescens Tokunaga and Mur achi

Enewetak ? Is let : Aug . 1956 , Tuth i l l .Dasyhelea flavibesol is TokunagaJapi an Is let : Aug . 1 956 , Tuth i l l .

Dasyhelea pel i l iouens is TokunagaJap tan Is let : Aug . 1956 , Tuth i l l .For cipomyia (Fo r cipomyia) tuth i ll i TokunagaJapi an Is let : Aug . 1 956 , Tuth i l l .Fam i ly CH IRONOMIDAEClun io tu thi ll i TokunagaEnewetak Is let : Sept . 1 975 , Cheng . Japtan Is let : Sept . 1 956 ,

Tuth i l l ; Sept . 1 975 , Cheng— det . by W . W . W i r th . Medren

Is let : Sept . 1 956 , Tuth i l l .Pon tomyia natans Edwards1 1 30N 1 62 15E ( lagoon ) : date? , Cheng?

Tanytor s us holoph i loe EdwardsJap tan Is let : Sept . 1 975 , Cheng— det . by W : W . W i rt h .

Telmatogeton pus i l lum EdwardsJapi an Is let : Sept . 1 975 , Cheng — det . by W . W . W i r th .

Tholes somyo mar it ime W i r thEnewetak Is let : Sept . 1 975 , Cheng . Enjebi Is let : Dec. 1 950 ,

Osh i r o . lku ren Is let : Sept . 1 975 , Cheng . }Japtan Is let : Aug .

1 956 , Tuth i l l ; Sept . 1975, Cheng— det . by W. W . W i r th .Fam i ly CH LOROPIDAECadremo pa l l ida (Loew)Enewetak ? Is let : Jan . 1 951 , Osh i r o . Enjebi Is let : Aug .

- Sept . ,Whee ler . lku ren Is let : Aug .

— Sept . , Wheeler . Japi an Is let : May1 946 , Townes ; Aug — Sept . 1 956 , Tuth i l l ; Jan - Feb . 1 957 ,

Tuth i l l ; Aug — Sept , Wheeler — det . by C. Sabr os ky.

“

Gau rax”

bicolor ipes (Mal loch)Japi an Is let : Aug

— Sept . 1 955 , Wheeler ; Aug . 1956 , Tuthi l l ;Jan . 1 957 , Tuth i l l- det . by C . Sabr osky (gener ic pos i tionuncer t ain , not Oscinosomo) .

Fam i ly EPH YDRIDAEA llot r ichomo ol ium Cresson

Jap tan Is let : Aug — Sept . 1 955 , Wheeler - det . by W . W . W i r th .D iscocer ino mer e Cr esson

Japi an Is let : Aug . 1 956 , Tuth i l l— det . by W . W . W i r th .H ecamede per s im i l is H endelJaptan Is let : Aug

— Sept . 1 955 , Wheeler .

_

Medr en Is let :Aug

— Sept . 1 955 , Wheeler .

? Is let : Dec . 1 950 , Osh i r o— det . byW . W . W i rt h .

H os t i'

s guamens is Cr esson

Japi an Is let : Aug . 1 956 , Tuth i l l ; Sept . 1975 , Cheng — det . byW . W . W i r th .

P lacop s idelle cynocephala Ker tészEnjebi Is let : Jan . 1 95 1 , Osh i ro — det . by W . W . W i rt h . lkurenIs let : Sept . 1 975 , Cheng . Japtan Is let : Sept . 1 975 ,

Cheng— det . by J . A . Tenor io . Medren Is let : Aug — Sept . 1 955 ,

Wheeler — def . by W . W . W i r th ; Sept . 1 975 , Cheng— det . byJ . A . Tenor io .

174 SAMUELSON AND NISH IDA

Fam i ly H IPPOBOSCIDAEO lfer s ia sp in ifer o (Leach)R ibewon Is let : May 1 962, Bushman — det . by T . C . Maa .

Fam i ly MIL ICH IIDAEDesm ometope ver ip

'

glpis Mal lochi “

Medr en Islet : Aug — Sept . 1 955 , Wheeler .

? Is let : Dec . 1 944 ,

Edgar ; May 1 946 , Townes — det . by C . Sabrosky.

M i l ichiello lecteipen n is (Loew)Enewetak ? Is let : Dec . 1 950 , Osh i r o . Japtan Is let : Aug — Sept .1 955 , Wheeler — det . by C. Sabr osky .

Fam i ly MUSCIDAEAther igona flovipalp is Ma‘l lochElugelab Is let : Jan . 1 951 , Osh i r o . Enjebi Is let : Jan . 1 951 ,

Osh i r o . Japtan Is let : Aug . 1 956 , Tuth i l l .A ther igona (Acr itochaeta) or ien talis SchinerJaptan Is let : Aug . 1956 , Tuth i l l .Musca (Musca) domes t ice L innaeusAnan ij Is let : Aug . 1 956 , Tuth i l l . lku r en Is let : Aug . 1 956 ,

Tuth i l l . Jap tan Is let : Aug . 1956, Tuth i l l . Medren Is let : Aug .

1956 , Tuth i l l . ? Is let : Dec . 1950 , Osh i r o .

Fam i ly PH OR IDAED ip lonevro (Dohr n iphor a) co r nuta (B igot)Jap tan Is let : Aug . 1 956 , Tuth i l l .

Megasel l'

o (Megesel l‘

o) scolor is (Loew)Japi an Is let : Aug . 1 956 , Tuth i l l .Fam i ly SARCOPH AGIDAEBoettcher isce kar nyi (H ardy)Japtan Is let : Nov . 1 944 , Edgar .

Parasar c0 phogo (L iosor cophego) m isero (Walker )Enewetak Is let : Sept . 1 975

:Cheng — def . by J . A . Tenor io .

Enjebi Is let : Jan . 1 951 , Osh i r o . Jap tan Is let : Aug . 1 956 ,

Tuth i l l .Phytoser cophogo g ressitt i (H al l and Bohar t)? Is let : Jan . 1 951 , Osh i r o .

Fam i ly SCATOPSIDAEH o lop leg io guamens is (Johannsen ) or near

lku r en Is let : Sept . 1975 , Cheng— det . by J . A . Tenor io .

Fam i ly SCIAR IDAEB r adys ie t r it ici (Coqu i llett )Japtan Is let : Aug . 1 956 , Tuth i l l .

Corynop ter e lat is tyleto (H ardy)Japtan Is let : Aug . 1 956 , Tuth i l l .

Plas tosciere let ipons H ardyJaptan Is let : Aug . 1956 , Tuth i l l ; Sept 1 956 , Tuth i l l .Fam i ly STRAT IOMYIDAEB rachycer a ven t r al is Thomson

Japtan Is let : May 1 946 , Townes .

Fam i ly SYRPH IDAEIschiodon s cu teller is (Fabr icius )Enewetak Is let : Nov . 1 944 , Dybas ; May 1 946 , Oakley and

Townes . Jap tan Is let : Jan . 1957 , Tuth i l l . ? Is let : Jan . 1951 ,Osh i r o ; date? , Fosber g .

Fam i ly T IPUL IDAEL imon ia (D icrenomyio) pect inungu is TokunagaAnan ij Is let : Aug . 1956 , Tuth i l l . Elugelab Is let : Jan . 1 95 1 ,

Os h i ro . Jap tan Is let : Aug . 1 956 , Tuth i l l ; Sept . 1 975 ,

Cheng— det . by F . G . H owar th .

Order H YMENOPTERAFam i ly ANTHOPH OR IDAEXy locopa sono r ina Sm i thEnewetak Is let : Aug . B r yan ; Feb . B r yans ight r ecords by E. H . B r yan , Jr .

Fam i ly BRACON IDAEZe le sp? Is let : May 1 946 , Townes .

Fam i ly EULOPH IDAEH em ip tor s en us sem iolbiclovus (Gi r ault)? Is let : May 1 946 , Townes .

Fam i ly EVAN IIDAESzep t igetello ser iceo (Cameron )Medr en Is let : Sept . 1 975 , Cheng— det . by G . Nakahashi .

Fam i ly FORMICIDAEMonomor ium pharaon is (L innaeus )Enjebi Is let : May 1 946 , Townes .

Fam i ly ICH NEUMON IDAEEchthr omorpha egr es to r io ins idiator (Sm ith)Bokombako Is let : Dec . 1 950 , Osh i ro . Elugelab Is let : Jan . 1 951 ,

Osh i r o .

Fam i ly MEGACH IL IDAEMegach i le di l igens hedleyi RainbowEnjebi Is let : May 1 946 , Oak ley ; May 1 946 , Oakley and

Townes . Jap tan Is let : May 1 946 , Townes .

Megachi le (Eu t r ichor oeo) fu llewayi Cockerel lEnewetak Is let : Sept . 1 975 , Cheng— determ ined by G . Nak

ahashi . Enjebi Is let : May 1 946 , Oak ley .Fam i ly SPH ECIDAECholybion bengalense (Dahlbom )Enewetak Is let : Sept . 1 975 , Cheng — det . by G . Nakahashi .

Pison p unct if r on s ShuckardJap tan Is let : May 1946 ,

Townes .

So l ierella peckhom i (Ashmead)Enjebi Is let : May 1 946 , Townes and Oakley .

Techysphex bengalen s is Cameron

Japtan Is let : Sept . 1 975 , Cheng— def . by G . Nakahashi .

Fam i ly VESPIDAEOdynerus sp .

Enewetak Is let : Sept . 1975 , Cheng — det . by G . Nakahashi .

Pechodyner us nas idens (Latrei l le)? Is let : May 1 946 , Townes .

Po l is tes fu scotus au r ifer (Saussur e)Jap tan Is let : Sept . 1 975 , Cheng — det . by G . Nakahashi .

Ropal l'

d l'

a marg inata (Lepelet ier )Enewetak Is let : Sept . 1 975 , Cheng . Japtan Is let : Sept . 1 975 ,

Cheng — det . by G . Nakahashi .

ACKNOWLEDGMEN TSWe are indebted to the late Denn is M . Devaney ,

B i s hop Museum , H ono lu lu , fo r encour ag ing ou r par t icipat ion in th i s vo lume . The late Edw in H . B r yan , Jr .

, B i shopMuseum , and Leonar d D . Tuth i l l (r et i red) , Un iver s i ty of

H awai i , H ono lulu , r elated fi r s t-hand in format ion about theenv i r onmen t and human act iv i t ies on Enewetak Ato ll .Rober t E . Elbel , U n iver s i ty of U tah , Salt Lake Ci ty , fu rn i shed r ecor ds fr om ear l ier inves t igat ion s by John 8 . Bushman and other s of the U n iver s i ty on ver tebr ates and ectopar as i tes fr om the ato l l . Lanna Cheng , Scr ipps Ins t i tut ionof Oceanogr aphy , La Jo l la, Cal i for n ia; Rober t E . Elbel ; and

Cur t i s W . Sabr osky and W i l l i s W . W i r th , Nat ional Museumof Natur al H i s tor y , Was h ington , D . 0 ,

prov ided in fo rmat ion on pr ob lemat ical spec ies or determ ined r elat ivel yr ecen tl y co l lected spec imens fr om the ato l l . The fo l lowingalso as s is ted u s in pr ov id ing iden t i f icat ions or r elated informat ion : Syoz i r o Asahina, Tokyo ; Peter F . Bel l inger , Cal i

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176 SAMUELSON A

Fukush ima, l . , 1 947 , Noctuidae of M icr ones ia, Mush i , 18 : 1 -22 .

Goodn ight , C. J. , and M . L . Goodn ight , 1957 , Op i l iones , In sectsMicrones ia, 3 : 71 -83 .

Ghaur i , M . S. K . , 1966 , Rev is ion of the Genus Or os ius D istant(H omopter a: Cicadello idea) , Bu l l . B r . Mus . (Nat . H is t .) En to

mo l . , 18 : 23 1 -252 .

G i l logly , L . R . , 1962, Co leoptera: N it idul idae, Insects M icr ones ia1 6 : 1 33- 188 .

Gr ess i tt, J . L . , 1954 , Intr oduction , Insects M icrones ia, 1 : 1 -257 .

1 955 , Co leopter a : Chr ysomel idae, Insects Micrones ia, 1 7 :

1 -60 .

1 956 , Co leoptera: Cerambycidae, Insects M icr ones ia, 17:

6 1 - 1 83 .

Gr oss , G . F . , and J . C . Schaffner , 1 963 , H eteroptera : Cor eidae(A lydin i by J . C. Shaffner ), Neididae, and Nabidae, In sectsMicr ones ia, 7 : 357-390 .

H ardy , D . E . , 1956 , D iptera: B ibion idae and Scatops idae, Insect sM icrones ia, 12 : 87- 1 02 .

1956 , D ipter a: Dor i laidae (Pipuncu l idae), Insects Micr ones ia,

1 3 : 1 -9 .

1 957 , Diptera: Coelop idae, In sects M icr ones ia, 14 : 41 -46 .

1 958 , Omphral idae (Scenopin idae), In sects Micr ones ia, 13 :

1 1 - 13 .

and M . Adach i , 1 956 , D iptera: Tephr i t idae, InsectsMicr ones ia, 1 4 : 1

-28 .

H athaway, W . H . , 1 953 , The La nd Vegetation of A r no Ato l l ,Mar shal l Is lands , A to l l Res . Bul l . , 16 : 1 -68 .

H er r ing , J . L . , 1967 , H eteroptera: An thocor idae, InsectsM icr ones ia, 7 : 391 -4 14 .

H opkins , G . H . E . , 1 96 1 , Siphonaptera, In sects Micr ones ia, 14:

91 - 1 07 .

Jackson , W . B . , 1 968 , Dr agonfly Populations at En iwetok Ato l l ,B ioscience, 18 : 1 1 23 .

James , M . T . , 1 962, D iptera : St r at iomyidae; Cal l iphor idae,

In sects Micr ones ia, 1 3 : 75- 1 27 .

John , H . , 1 960 , Eine Uber s icht i i ber die Fam i l ie Pr opalt icidae

(Co leoptera), Poc . Insects , 2 : 1 49- 1 70 .

1971 , Co leoptera: Pr opalt icidae, Insects Micr ones ia, 1 6 :

287-294 .

Kato , M . , 1960 , H omoptera: Membr acidae, Insects M icr ones ia, 6 :

345-351 .

Kempny, P . , 1 904 , Bei tr ag zur Neu r opter nfauna der Mar shal lInseln , nebst Beschr eibung zweier neuer Ch r ysopa-Ar ten ,

Ver b . Zoo l . -Bo t . Ges . Wien . , 54 : 352-355 .

Kohls , G . M . , 1 957 , Ixodo idea, Insects Micrones ia, 3 : 85- 1 04 .

Krantz , G . W . , 1 967 , Acar ina: Mesos t igmata. Macr ochelidae,

In sects M icrones ia, 3 : 1 49- 1 54 .

Krombein , K . V . , 1 949 , The Aculeate H ymenoptera of

M icrones ia I . Sco l i idae, Muti l l idae , Pomp i l idae and Sphecidae,

Proc . H awa i i . En tomo l . Soc . , 1 3 : 367-410 .

1 950 , The Aculeate H ymenopter a of M icr ones ia II . Co l let idae, H al ictidae , Megach i l idae, and Apidae, Proc . H awa i i .

En tomo l . Soc . , 14 : 1 0 1 - 1 42 .

Kulzer , H . , 1 957 , Co leoptera: Tenebr ion idae , In sects M icrones ia17 : 185-256 .

Kurosawa, M . , 1940 , Thysanoptera of M icr ones ia, Ten th r edo , 3 :

45-57 .

L ieft inck , M . A . , 1 962 , Odonata, Insects Micr ones ia, 5 : 1 -95 .

L innavuor i , R . , 1960 , H omopter a: Cicadel l idae, InsectsM icr ones ia, 6 : 23 1 -344 .

1 975 , H omopter a: Cicadel l idae, Supplement , Insects

M icrones ia, 6 : 6 1 1 -632 .

L6bl , l . , 1 98 1 , Co leoptera : Scaphidi idae, In sects M icr ones ia, 1 5:

69-80 .

INSECTS AND ALLIES (ARTHROPODA)

Steyskal , G . , and M . Sasakawa, 1 966 , D iptera: Clus udae, InsectsM icr ones ia, 14 : 243 -249 .

Stone,A . , 1 960 , D ipter a: Taban idae, Insects M icrones ia, 1 3 :

51 -53 .

1 964 , D iptera: Simul i idae, Insects M icr ones ia,12 : 629-635 .

Str ohecker , H . F . , 1958 , Co leoptera: Endomychidae, InsectsM icrones ia, 1 6 : 1 05- 1 08 .

Sugerman , B . 1 972a ,Insects and Other Ar thropods fr om

Kwajalein Ato l l (Mar shal l Is lands ) , Pr oc . H awa i i . En tomo l .Soc . , 21 : 271

-286 .

1 972b, Insects of L ib ls land (Mar shal l Is lands ), Pr oc . H awa i i .

En tomo l . Soc . , 21 : 287-289 .

1 979 , Addi tions to the L ist of In sects and Other Ar thr opodsfr om Kwajalein Ato l l (Mar shal l Is lands ) , Pr oc . H awa i i . En tomo l .Soc . , 23 : 147

- 1 51 .

Synave, H . , 1 957 , H omoptera: Cer copidae, Insects Micr ones ia, 6 :

2 1 3 -230 .

Takahash i , R . , 1956 , H omoptera: A leyrodidae, InsectsM icr ones ia, 6 : 1 - 13 .

Takash ima, H . , 1950 , Notes on Amblypygi Found in Ter r itor iesAdjacent to Japan , Pec . Sc i . , 4 : 336-338 .

Theodo r , O . , 1 966 , D ipter a: Nycter ibi idae, Insects M icr ones ia,

14: 275-279 .

Tho r nton , I . W . B . , S. S. Lee, and W . D . Chui , 1972 , Psocop

ter a, Insects M icrones ia, 8 : 45- 1 44 .

Tokunaga, M . , 1964 , D iptera: Ch i r onom idae, Insects M icrones ia,

1 2 : 485-628 .

and E . K . Mu rachi , 1 959 , D iptera: Ceratopogon idae, InsectsM icr ones ia, 1 2 : 1 03 -434 .

Townes , H . , 1 946 , Resu lts of on En tomo log ical Inspect ion Tour of i i

M icr ones ia,Econom ic Sur vey of M icrones ia, 1 4 : 1

-8 1 , U . S.

Commercial Company .1 958 , H ymenoptera: Ichneumon idae, Stephan idae, and

Evan i idae, Insects M icr ones ia, 1 9: 35-87 .

Tuth i l l , L . D . , 1964 , H omoptera : Psy l l idae, Insects M icr ones ia, 6 :

353 -376 .

Uch ida , H . , 1 944 , Co l lem bo la von M ikr ones ien , Bu l l . Tokyo Sci .M us . , 1 7 : 1

-23 .

U s inger , R . L . , 1 949 , War-time D isper sal of Pacific Is land Nys ius

(H em iptera: L ygaeidae), Pr oc . H awa i i . En tomo l . Soc . , 13 :

447 .

1 95 1 , H eter opter a of the Mar shal l Is lands , Proc . H awai i .

En tomo l. Soc. ,1 4 : 3 1 5-321 .

1 952 , New Species and Additional Records of H eter opter afrom the Mar shal l Is lands , Proc . H awai i . En tomo l . Soc .

,1 4 :

51 9-524 .

U s inger , R . L . , and G . F . Fer r is , 1 960 , H eter optera: Cim icidae ,

Ins ects M icr ones ia, 7 : 285-286 .

U s inger , R . and I . LaR iver s , 1 953 , The Insect L ife of Ar no ,

A to l l Res . B u l l . , 1 5: 1 -28 .

U s inger , R . L . , and P . Wygodz insky, 1 960 , H eter opter a: En icocephal idae, In sects M icr ones ia, 7 : 21 9

-230 .

177

Van Du zee, M . C. , 1 933 , New Do l ichopodidae fr om the

H awai ian Is lands (D iptera) , Pr oc . H awai i . En tomo l . Soc . , 8 :

307-357 .

Van Zwaluwenbu r g , R . H . , 1 948 , New Species and New Reco rdsof Elater id Beet les fr om the Paci fic— II I , Pr oc . H awa i i . En tomol . Soc .

,1 3 : 265-276 .

1 957 , Co leoptera: Elater idae, In sects M icrones ia, 1 6 : 1-66 .

Watanabe, C . , 1 945 , Notes on Some M icr ones ian B racon idae,

Mush i , 1 6 : 47-58 .

1958 , H ymenopter a : Euchar idae, Insects Micr ones ia, 1 9:

19-34 .

Wer ner , F . G . , 1 965 , Co leoptera: Anth icidae, Insects Micr ones ia,

1 6 : 255-269 .

Wheeler , M . R . , and H . Takada, 1 964 , D iptera: Drosoph i l idae,

Insects Micr ones ia,1 4 : 1 63-242 .

W i l l iam s , F . X . , 1 93 1 , H andbook of the Insects and Other Invertebr ates of H awa i ian Sugar Cane F ie lds , H ono lulu : H awai ianSugar P lanter s ’

Association , 400 pp .

W i lson , E . O . , and R . W . Taylor , 1967 , The Ants of Po lynes ia,

Pec . In sects Monog r ,14 : 1 - 1 09 .

W i lson , N . , 1 967 , Acar ina: Mesos t igmata. Dermanys s idae, Laelap idae, Sp in tu r n icidae Paras itic on Ver tebr ates , InsectsM icr ones ia, 3 : 1 33 -148 .

1972 , Anoplura, Supplement , Insects M icr ones ia, 8 :

1 45- 148 .

W i r th , W . W . , 1 95 1 , A Rev is ion of the D ipterous Fam i ly Canaceidae, Occas . Pop . Ber n ice P . B ishop Mus . , 20 : 245

-275 .

W ittmer , W . , 1 958 , Co leopter a: Lampyr idae, Canthar idae,

Malachi idae, and Pr ionocer idae, Insects M icr ones ia, 1 6 : 67-74 .

1970 , Co leoptera: Malachi idae, Supplement, InsectsM icrones ia, 1 6 : 279-280 .

Wood, S. L . , 1 960 , Co leoptera: Platypodidae and Sco lyt idae,

In sects M icrones ia,1 8 : 1 -73 .

Woodbur y , A . M . , 1 962 , A Review of the Eco logy of En iwetokA to l l , Pacif ic Ocean , Un iver s ity of U tah , Inst i tute of Env i r on

mental B io log ical Resear ch , pp . 1 - 1 23 .

Wygodz insky, P . , and R . L . U s inger , 1 960 , H eter optera : Reduv iidae, In sects M icr ones ia,

7 : 23 1 -283 .

Yosh imoto , C . M . , 1 962 , H ymenoptera: Euco i l inae (Cyn ipo idea) ,Ins ects M icr ones ia, 1 9: 89- 1 07 .

and T . Ish i i , 1 965 , H ymenoptera: Chalcido idea: Eu lophidae,

Encyr t idae (par t) , Pteromal idae, Insects M icr ones ia, 1 9:

1 09- 1 78 .

Z immerman , E . C . , 1 942 , Cur cu l ion idae of Guam , Bu l l . Bern iceP . B ishop Mus 1 72: 73 - 146 .

1 948 , Notes on Mar ianas Is lands Cur cul ion idae (Co leoptera) ,Proc . H awa i i En tomo l . Soc . , 13 : 305

-3 1 5 .

1 964 , Plotys im u s in Southeaster n Po lynes ia (Co leoptera:Cur cul ion idae : Ot ior hynch inae: Celeu thet in i ) , Poc . Insects , 6 :

747-752 .

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TABLE 1

Check l i s t of Pycnogon i da of Enewetak A tol l

Phy lum ARTH ROPODAClass PYCNOGON IDAOrder PANTOPODAFam i ly AMMOTH EIDAEAmmothella s tau r omata Ch i ld , 1982 .

“ Fam i ly CALL IPALLENIDAECall ipallene sp . cf . novaezelandiae (T hom son ,

Fam i ly ENDEIDAEEndeis nodosa H i lton , 1 942 .

Fam i ly PH OXICH ILID IIDAEAnop lodactylus g landu l ifer Stock , 1 954 .

Anop lodactylus mar shallen s is Ch i ld , 1 982 .

‘New Enewetak and M icrones ian r ecord .

i s lands and ato l ls of M icr ones ia w i l l undoubtedly add manyspec ies to the faunal checkl is t shown in Table 1 . Collec

t ions of cor al r ubble fr om lagoon and outer reef habi tatsw i l l pr obably y ield add i t ional new d is tr ibut ional r ecor ds and

new spec ies . Scr ap ing for foul ing organ isms on p ier p i l ings ,nav igat ion buoys , and sh ip ’

s hulls is another method forsampl ing common pycnogon id habi tats and should incr easeou r knowledge of that fauna . Pycnogon id hab i tats havehar dly been touched on mos t of the ocean ic is lands of the

Pac i fic .

COLLECTION DATA

Ammothella s tau r omata, Anop lodactylus g landu l ifer ,and Anop lodactylus mar shallen s is were all co l lected by theauthor in 1 969 fr om p ier p i l ings on the nor th end of

Enewetak ls land (s i te name Fr ed) . Call ipallene sp . cf .novaezelandiae and Endeis nodosa wer e co llected by theauthor in 1969 and by other s in 1 974 and 1 975 fr omlagoon r ubble and coral r ocks in fai r l y shal low water . The

Call ipallene spec imens were found in r ubble on a lagooncoral kno l l near Jinedr ol Is land (s i te name A lv in) and in

coral r ubble fr om Enewetak Is land . Endeis nodosa camefr om coral rocks in the lagoon off Enewetak Is land and

fr om p ier p i l ings on the lagoon s ide of Run i t l s land (s i tename Yvonne) . No specimens have been repo r ted fr om ther ich outer r eef habi tats of Enewetak Ato l l .

Ammothella s tau romata and Anop lodactylus mar shallens is ar e r ecen tl y descr i bed species (Ch i ld , 1 982) and ar e

known on l y fr om Enewetak Ato l l . Anop lodactylus

g landu l ifer Stock , 1 954 , has the w ides t known d i s tr i but ionof the five species . I t is known in w idely s cattered loca li tiesfr om the Red Sea and the eas t coas t of Afr i ca to Burmaand Singapore and for the fi r s t t ime in the centr al Paci fi cis lands at Enewetak . Endeis nodosa H i lton , 1 942 , has beenknown on ly fr om H awai i , on Oahu Is land , and the

Enewetak spec imens extend i ts known r ange cons iderablyto the wes t . The Call ipallene specimens show s l ight d i fferences from Thom son ’

s (1 884) spec ies novaezelandiae,but

i f they are in fact th is spec ies , the prev ious l y known souther n hem ispher e d is tr ibut ion of C . novaezelandiae (eas tcoas t of Afr ica to New Zealand) would be extended to the

CH ILD

ACKNOWLEDGMEN TSThe author i s grateful to the Atom ic Energy Commis

s ion wh ich operated the Enewetak Mar ine B iolog icalLabor ator y under the management of the Un iver s i ty of

H awai i , to the base suppor t of Kent r on lnc . , and to the

U . 5 . A i r For ce for field suppor t wh i le specimens for theSm i thson ian Ins t i tut ion were co l lected . I also w ish to thankmy fel low d iver s fr om the Sm i thson ian and the U . S.

Geo log ical Su r vey for thei r suppor t and specimencontr ibutions .

nor ther n hem ispher e at Enewetak . The recor d depths forall five spec ies are shallow to l i ttoral .

In general , the four genera have a wor ldw ide dis t r ibut ion , pr edom inan tly in the tr op ics , but there ar e species of

Endeis,Anop lodactylus , and Call ipallene in deep sea habi

tats and even in co ld Antar ct ic water s . Anop lodactylus i s

the larges t genus r epr esen ted in term s of spec ies knownw i th about 75 now r ecogn i zed . It has sever al knownr epr esen tat ives in mos t paper s on the per imeter Pac i ficis land gr oups , including Aus tr al ia and New Zealand , and

appear s in the few repor ts that ex is t on wes ter n and m id

Pac i fic is lands . There ar e close r elat ions of the Enewetakspecies of Anop lodactylus in Japan and the Indian Ocean .

The speci fic food prefer ences of mos t pycnogon idspecies are unknown ; so ther e can be no compar isons of

hab i tat- food pr efer ence among the var ious species .

The genus Ammo thella, w i th 28 known species , istropi cal — temper ate in d is tr ibut ion and has i s land r ep r esentat ives in s catter ed local i t ies around some par ts of the

Paci fic per imeter . Call ipallene, w i th 28 known species , isone of man y genera in a large fam i ly wh ich seems to havean area of pro l i ferat ion , at the gener ic level , in the

Aus tr al - Indones ian ar ea (refer r ing to the is land arc fr omAus tr al ia and New Zealand thr ough the Phi l ippines) . Of the24 genera in the fam i ly Callipallen idae, 1 3 gener a haver epr esentat ives in th is ar ea. One genus (Par apaIlene) hasthe major i ty of i ts known species found in th is area, and

three genera (Pycnopallene,Spasmopallene, and

StyIOpallene) ar e endem ic . Ther efore, i t would not be

surpr is ing at all to find a num ber of coral reef species of

callipallen ids at Enewetak and the Mar shal l Is lands . The

genus Endeis ,‘

w i th 1 7 r epor ted spec ies , has seven fr omthe Aus tr al- Indones ian reg ion .

REFERENCES

Bar nard, J . L . , 1 965 , Mar i ne Amph ipoda of Ato lls in Micr ones ia,

Proc . U . 8 . N at . M us . , 1 77:-4596 52 .

Ch i ld, C . A .,1982 , Pycnogon ida fr om the Mar shal l Is lands , Proc.

B io . Soc . Wash . , 95: 270 28 1 .

H i lton , W . A . , 1 942, Pycnogon ids from H awai i , Occas . Pap .

Ber n ice P . B ishop Mus . , 17: 43-55 .

Stock , J . H . , 1 954 , Pycnogon ida fr om Indo-Wes t Paci fic, Aus ’

t ral ian , and New Zealand Water s , Paper s fr om Dr . Th.

Mor tensen ’

s Pacific Expedition 1914— 16 , Vidensk . Medd.

Dansk Natu r h . Foren . , 1 1 6: 1- 1 68 .

Thom son , G . M . , 1 884 , On the New Zealand Pycnogon ida, wi thDescr ipt ions of New Species , Trans . and Proc . New ZealandIns t itute, 1 6 : 242-248 .

Chapter 1 7

Stoma topod Crus tacea of Enewetak A to ll

MARJORIE L . REAKA‘and

RAYMOND B .

‘Depar tmen t of Zoo logy , Un iver s i ty of Mar y land,

Co l lege Par k, Mar y land 20742 ; TDepar tmen t ofIn ver tebrate Zoo logy, Nat iona l Museum ofNatur al H is tor y

, Sm ithson ian Ins t itut ion ,

Was h ingt on , D . C . 20560

IN TRODUCTIONTh is s tudy pr ov ides a checkl is t and d iscus ses the

biogeog raph ic relat ionsh ips of the 1 2 spec ies of Stomatopoda (man t is shr imps ) now known to occur on EnewetakAto l l . Five species ar e w idel y d is tr ibuted throughout theIndo-Wes t Pac i fic r eg ion , and three species have somewhatmor e res tr icted r anges in the Indo -Wes t Pac i fic and Ind ianOcean . Four spec ies ar e endemic to the Cen tr al Pac i fic(two to Enewetak) , and thr ee taxa need fur ther taxonom icinves t igat ion , wh ich may demons trate fu r ther endem ism .

Stomatopods from Enewetak ar e dwar fed in body s i zecompar ed to thei r mainlaind r elat ives . Smal l s i ze has

s tr ong con sequences for l ife h is to r y and evo lut ionar y patter n s in s tomatopods , and in par t icular is l ikel y to gener ateendem ism . We prov ide in format ion on the color patter nsof the s tomatopods fr om Enewetak , show ing wh ich tr ai tsar e the mos t r el iable ind icator s of spec ies ident i ty for taxonom ic and field resear ch and wh ich trai ts are mos t l ikel yto be in fluenced by body s i ze, sex , or hab i tat . Severalanomal ies in usual ly invar ian t color tr ai ts are found ins tomatopods fr om Aomon Is land , wh ich was closer to

sour ces of r ad iat ion fr om atom ic tes ting than mor e southern is lands in the ato l l . We also summar i ze what i s knownabout the habi tat and fight ing behav ior of the coraldwel l ing man tis shr imps from Enewetak . For each of the

above topics , we ident i fy or d iscus s all Pr ev ious l i teratur eon the s tomatopods of Enewetak .

CH ECKLIST

The s tomatopod Cr us tacea from Enewetak Ato l l wer efi r s t recor ded by Mann ing ( 1 97 1 ) and mor e r ecen tly byMann ing and Reaka these s tud ies included the

descr ipt ion of thr ee new spec ies . H er e we ident i fy fourspec ies (Gonodactylus ter natens is , Pseudosqu i lla ci liata,

P . om ata, and Lys iosqu i l la macu lata) that have no t beenr ecor ded p r ev ious l y from Enewetak , br ing ing to 1 2 the

total num ber of species known from the ato ll (TableWe also ident i fy her e all per t inen t l i ter ature in wh ich part icu lar species of s tomatopods fr om Enewetak have beend i scus sed .

B IOGEOGRAPH IC COMPARISON SPseudosqu i lla ci l iata is the on ly spec ies of those found

on Enewetak that also occur s in the Atlan t ic . Five of the

spec ies known fr om Enewetak (G . ch i r ag r a, G . p latysoma,

P . ci l iata, P . or nata, and L . macu lata) ar e w idel y dis t r ibu ted thr oughout the Indo —Wes t Pac i fic r eg ion . All of thesear e r elat ivel y large species of s tomatopods , and

Pseudosqu i lla has a par t icular ly long lar val d isper sal s tage(Reaka, 1 979a, 1 980a ; Reaka and Mann ing , 1 987a) .

Thr ee spec ies have somewhat more r es tr icted dis t r ibut ions in the Wes ter n Pac i fic , lndo-Malayan , and Ind ianOcean r eg ions than the pr ev ious ly ment ioned gr oup .

H ap tosqu i lla g lyp tocer cus occur s fr om Enewetak to Japan ,

nor ther n Aus tr al ia, and as far wes t as the AndamanIs lands . G onodactylus sm i th i i i s known from Ocean ia and

the South Ch ina Sea and fr om Aus tral ia to the wes ter nInd ian Ocean ; however , populations from the wes ter nInd ian Ocean and Red Sea may be refer able to anotherspec ies , G . acu t i r os t r is De Man . G onodactylus ter natens is

is known w i th cer tain ty on ly from local i t ies betweenSamoa and Indones ia .

Of the r emain ing species , G . esp inos u s and G . in i

cip ien s are known on ly from i s land local i t ies in the Cen tr alPac i fic , wh i le G . insu lar is and G . m icr ones ica apparen tl yare endem ic to Enewetak Ato l l . These four spec ies , as wel las H . g lyp tocer cus (wh ich also has a r elat ivel y r es tr ictedd is tr ibut ion ) , ar e the smal les t species that occur on

Enewetak .

F igu r e 1 shows that the typ ical body s i zes of popu la

t ions of all of the coral-dwel l ing s tomatopods fromEnewetak are d im inut ive compar ed to those fr om manyother geogr aph ic local i t ies , par t icular ly the Indo-Wes tPac i fic main land ar eas . Th is fact is emphas i zed by comparing l ineages of the same o r closely related spec ies in di fferent r eg ions . For example, ind iv iduals of G . sm ith i i attaincons iderably smal ler s i zes in Enewetak than in Thai land or

1 82 REAKA AND MANNING

TABLE 1

Check l i s t of Stomatopoda fr om Enewetak A tol l

Phy lum CRUSTACEA

Clas s MALACOSTRACAOrder STOMATOPODAFam i ly GONODACTYLIDAEG onodactylus ch i ragr a (Fabr icius ) : Mann ing , 1 971 .

G onodactylus esp inosus Bo r radai le: Mann ing , 1971 .

Gonodactylus incip iens Lanchester : Reaka, 1973 ; D ing le et al . , 1973 ;

Caldwell and Br u nenmeis ter , 1 975 ; Reaka, 1 975a; Ca ldwell and Di ng le, 1977; Reaka, 1 979a .

G onodactylus vir idis Serene: Ca ldwel l and D ing le, 1972 .

Gonodactylus ch i ldi Mann ing : Mann ing , 1 972 .

Gonodactylus insu lar is Mann ing and Reaka : Mann ing and Reaka, 1 982

Gonodactylus falcatus (For ské l) : Mann ing , 1971 ; Ca ldwel l and D i ng le, 1972; D ing le et al . , 1 973 ;

Reaka, 1 973 ; Reaka, 1 975a; Reaka, 1975b; Reaka, 1 976 ; Reaka, 1 979a; Mann ing and Reaka, 1 982 .

Gonodactylus m icr ones ica Mann ing : Mann ing , 1 971 ; D ing le et al . , 1 973 ; Reaka, 1 973 .

Gonodactylu s p latysoma Wood-Mason : Mann ing , 1 971 ; Caldwel l and D ingle, 1 972 ; Reaka, 1973 ;

Ca ldwel l and B r u nenmeis ter , 1975 ; Reaka, 1 975a; Reaka, 1 975b; Reaka, 1 979a.

G onodactylu s sm ith i i Pocock : Mann ing , 1 971 ;_

Ca ldwel l and Di ng le, 1 972 ; Reaka, 1973 ; Di ngle et al . , 1973 ;

Ca ldwel l and B r unenmei s ter , 1 975 ; Reaka, 1 975a; Reaka, 1975b; Reaka, 1 979a; Reaka, 1 979b .

‘G onodactylus ter naten s is De Man , 1 902 (new r ecord based on specimen in USNM co l lections) .Fam i ly PROTOSQUILL IDAEH ap tosqu i lla g lyp tocer cus (Wood-Mason ) : Mann ing , 1 971 ; Ca ldwel l and D i ng le, 1 972 ; D ing le et al . , 1 973 ;

Ca ldwel l and Br unenmei s ter , 1 975 ; Reaka, 1 975a; Reaka, 1 975b; Reaka, 1 976 ; Ca ldwel l and D ing le, 1 977 ;

Reaka, 1 979a; Reaka, 1 979b .

Fam i ly PSEUDOSQU ILL IDAE‘Pseudosqu i lla ci liata (Fabr icius , 1 787) (new r ecord based on specimen in USNM co l lection ) .

‘Pseudosqu i lla or nata M ier s , 1 880 (new r ecord based on specimen in R i jksmuseum van Natuu r l ijke H i stor ie, Leiden) .Fam i ly LYSIOSQU ILLIDAE

‘Lys iosqu i lla macu lata (Fabr icius , 1 793 ) (new r ecord based on specimen determ ined by R . K inzie for M id-Paci ficMar ine Labor ator y r efer ence co l lection ) .

N ew records for species at Enewetak .

Aus tr al ia (and we sugges t that th is may be an endem icpopulat ion ) . Ind iv iduals of G . inc ip iens fr om Enewetak ar esmal ler than thei r cognates from Thai land (G . vi r idis) or

Aus tral ia (G . aff in is) . Gonodactylus in su lar is in Enewetak isa dwar f species compared to i t s close r elatives in Thai land(G . m u tatus) , Aus tral ia (G . falcatus) , and H awai i (G . aloha) .

These data cannot be accounted for by co l lecting techn iques , sample s i zes , or lat i tude and temperature . (See

Reaka and Mann ing , 1 987a, for fu r ther d i scus s ion ) . Onel ineage, that of H . glyp tocer cus , does no t f i t th is patter ns ince i t i s not smal ler in Enewetak than in Thai land .

We have d i s cussed elsewhere reproduct ive and l i fe hi stor y tr ai ts and thei r r elat ionsh ip to body s i ze, d i sper salab i l i ty , and evo lut ionar y r ates in s tomatopods , includ ingthose fr om Enewetak (Reaka, 1975a, 1 976 , 1 978 , 1 979a,

1 979b , 1 980a ; Reaka and Mann ing , 1 98 1 , 1 987a) . L i feh i s tor y tr ai ts ar e scaled to body s i ze w i th in r elated l ineagesof s tomatopods , and the l i fe h i s tor y patter ns of spec iesfr om Enewetak con form to these over al l patter n s . In par t ic

u lar , however , smal l body s i ze and r es tr icted d isper sal abi li ty gener ate r apid evo lut ionar y changes and ar e l ikely tobe as soc iated w i th endem i sm . The tendency for dwar f ismand endem ism in the s tomatopods fr om Enewetak prov idesanother example of th i s tr end .

COLOR PATTERN S

Desp i te thei r cr ypt ic hab i ts , s tomatopods often are

flam boyan t l y co lor ed (Reaka , 1 975b, 1 980b, 1 98 1 ; Reaka

and Mann ing , Some of these co lo r patter ns var ydramat ical ly among ind iv iduals of the populat ion ,

wh i leother tr ai ts are invar iable in color . In another s tudy on thecor al-dwel l ing s tomatopods of Enewetak (Reaka and Man

n ing , 1 987b), we pr ov ide the fi r s t quan t i tat ive anal ys is inany cr us tacean of the degr ee of co lor po l ymo rph ism in all

mor pho log ical tr ai ts , and we tes t a ser ies of hypothesesabout the funct ion of co lor patter ns in s tomatopods . Invar i ’

an t and spec ies -

spec i fic co lor tr ai ts can fac i l i tate accurateident i f icat ion of mor pho log ical l y s im i lar species in the fieldand also can be used to resolve taxonom ic d i fferencesamong s ibl ing spec ies (Mann ing , 1 964 ,

1 97 1 ; Mann ing and

Reaka , 1 979 ,1 98 1a, 1 98 1 b, Ther efor e, we pro

v ide her e a summar y of the trai ts that w i l l be ei ther themos t useful or the mos t l ikely to lead one as tray in fieldiden t i ficat ions of the common cor al-dwel l ing s tomatopodsin Enewetak .

The mer al spo t (an often br ightly co lo red oval indentat ion on the dor somed ial su r face of each of the r aptor ialsecond max i l l ipeds ) is one of the mos t r el iable char acter i s

t ics for iden t i fy ing spec ies (Table The meral spo t is

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1 86 REAKA AND MANNING

wh i t ish pink in G . incip ien s , yel low w i th an an ter ior red

in fus ion in G . insu lar is , wh i te in G . m icr ones ica and

G . p latysoma,carm ine r ed su r r ounded by a black-and

wh i te r ing in G . sm ith i i , and cr eam co lor ed in H . glyp to

cer cus . Ou r anal ys is , however , shows that the co lor of the

meral spot r anges fr om blue green i sh to salmon amongind iv iduals clas s i fied as G . incip iens ; morpho log ica l d i fferences , especiall y in the s tr ucture of the telson , amongmember s of th is populat ion also sugges t that th is taxonneeds fu r ther inves t igat ion . Add i t ional ly , indiv iduals in the

Enewetak populat ion of G . sm ithi i have a carm ine meralspot , wher eas mem ber s of other populations of G . sm ith i ibear a dar k purpl ish r ed mer al spot . Th is fact , in combinat ion w i th the obser vat ions that G . sm i th i i fr om Enewetakar e d im inut ive in body s i ze, produce smal ler eggs relat iveto thei r body s i ze, and inves t a smal ler pr opor t ion of thei rbody vo lume in a brood of eggs than do other populat ionsof G . sm ithi i (Reaka, 1 975a, 1 979a; Reaka and Mann ing ,

1 987b) , sugges t that th i s population also is in need of taxonom ic inves t igat ion and may r epr esen t an endem icspec ies .

Sever al other invar ian t tr ai ts also ar e useful for speciesiden t i ficat ions , par t icular ly when used in combinat ion w i ththe meral spots . H ap tosqu i lla g lyp tocercus spor ts un iquel ymaroon an tennules and wh i te antennae among the spec iesat Enewetak . The an tennal s ca les of G . p latysoma ar e

whi te pr ox imall y but gr een d is tal l y . The pr opodu s of the

raptor ial appendage in G . sm i thi i is i r idescent blue pu r ple,

set again s t a br ight yel low backgr ound outl ine where i t

fo lds again s t the mer us . Later al dar k spots char acter i ze thepr opodus of the r aptor ial appendage in G . in cip ien s(although thei r number incr eases w i th depth and type of

hab i tat) . The propodu s and dactyl of the rapto r ial max i ll iped ar e un iquel y r ed in G . m icrones ica. The dacty l ofthe r aptor ial appendage in G . p latysoma i s br ight blu i shblue-

gr een . Gonodactylus p latysoma also i s remar kable forthe co lor ful concen tr ic eye spo ts ”

on each s ide of the

eighth thor acic and s ixth abdom inal segmen ts . On the

thorax th is spot is composed of an outer wh i te band , followed by a pale blue, r ed, dar k blue, and final ly an innermos t r ed orange c i r clet of co lor . On the s ixth abdom inalsegment, the “

eye spot is formed of an outer band of

wh i te, fo l lowed by r ed brown and dar k blue r ings and an

innermos t patch of yel low orange. The basal segmen t ofeach ur opod in G . p latysoma is orange w i th a l ight anddar k blue spot . The basal segmen t of each ur opod inG . sm i thi i bear s a br i l l iant red spot wh ich is consp icuous l yexposed agains t dar k gr een body co lorat ion when the u ro

pads are flar ed (par t icular l y when the telson is co i leddefens ively in fr on t of the body dur ing a fight) ; these spotsvan ish when the uropods are tucked under the telson in a

subm is s ive pos tur e. In add i t ion , G . insu lar is is characteri zed by a l ineage-

spec i fic set of fou r black spots — two on

the s ixth abdom inal segmen t and two on the anter ior telson . The u r opodal exopods (and usual l y the endopods ) areun iquely r ed in G . insu lar is , as are the dor sal car inae on

the s ixth abdom inal segmen t and telson in G . m icr ones ica

(Tab le

In contras t to the invar iable and often un ique tr ai ts di scus sed above, however , the co lor of mos t trai ts var ies

among indiv iduals or w i th sex , developmental s tage, or

habi tat in mos t species . When the r ange of var iab i l i tyamong ind iv iduals is cons idered for all species (Reaka and

Mann ing , 1 987b) , all of the major morpho logi cal tr ai ts canbe r anked fo r relat ive var iabi l i ty of co lor , pr ov id ing an indicat ion of wh ich tr ai ts are l ikely to be the mos t— or the

leas t— rel iable ind i cator s of species identi ty in general .Except for the spec i fic cases iden t i fied in Table 2 as beinginvar iable,

the ur opodal exopods , antennules , antennae,

uropodal endopods , and antennal s cales , respect ively , forma set of tr ai ts that ar e moderately var iable in co lor . Similar ly, when all spec ies are cons ider ed (wi th invar iableexcept ions be ing noted in Table the co lor of the maxi l

l ipeds , walk ing legs , and uropodal for ked tips var ies s t i l lmor e among ind iv iduals ; body color is in gener al the mos tpo l ymorph ic of all trai ts . Descr ipt ions of species and fieldiden t i ficat ions based upon the latter character s ar e mos tlikely to be biased by smal l sample s i zes .

Co lor patter ns var y pr ed ictably w i th sex in some cases(Table The co lor of the body , walk ing legs , and max i llipeds are mo re likely to be gr een ish in males but maysh i ft towar d other hues (especial ly brown ish , yel lowi sh) infemales of some spec ies (G . sm ithi i, G . incip ien s , H . g lyp to

cer cus) . In G . incip iens , the antennae ar e mor e l ikely to bered or or ange in males but clear in females .

The co lor patter n of cer tain tr ai ts also var ies predictably w i th s i ze o r age in some spec ies . The incidence of

blue gr een or gr een colorat ion on the an tennal scales or on

the endopods , exopods , or for ked t ips of the ur opodsincreases in larger indiv iduals of sever al species(G . in cip iens , G . insular is , G . sm ith i i) . The co lor of the

uropodal endopod sh i fts towar d r ed (match ing the r ed of

the invar iant exopod) as indiv iduals of G . insu lar is becomelar ger . A lso , the an tennae and an tennules of G . p latysoma

change fr om clear to red as ind iv iduals matur e . Whi tespeckl ing on the body incr eases w i th s i ze in G . in cip iensand G . sm ithi i .

Sever al co lor character is tics sh i ft w i th habi tat w i th inspec ies , wher e these sh i fts cannot be explained by the dist r ibu t ion s of ind iv iduals of d i fferen t sexes and s i zes in di ffer en t habi tats . In G . in cip iens , the fr equency of wh i tetr an sver se band ing on the body , dar k spots on the

p r opodu s of the r aptor ial appendage, and yel low color at ionon the antennules and d is tal for k of the uropods increasesfr om s hal low m) to deeper to m) habi tats .

The degr ee of co lor po lymorph ism among ind iv iduals alsoincreases w i th depth (antennal s cales , max i l l ipeds , dis talfor ks of the u r opods ) in G . incip ien s . Addi t ional ly , theur opodal endopods in G . incip ien s sh i ft towar d clearco lorat ion in hab i tats that contain assemblages of aggres

s ive spec ies of s tomatopods .

Co lor patter ns also cor r elate w i th habi tat and suscept i

bi l i ty to pr edat ion among species . Because they are h ighlyvag i le or occupy deep , r eefward, or open exposed env i ronmen ts , G . incip ien s , G . p latysoma, and G . m icr ones ica ar e

l ikely to encoun ter cons iderable fish predation . On the

STOMATOPOD CRUSTACEA 187

other hand , G . insular is , G . sm ith i i , and H . g lyp tocer cus

are less vagi le, often occur in more protected and shorewar d habi tats , and undoubtedly suffer less pr edation thanG . incipiens , G . p latysoma, and G . m icr ones ica . Gonodac

tylus p latysoma and G .

“

incip iens are the mos t heav i lyspeck led and mott led species , and transver se banding is

most developed in these two spec ies and G . m icr ones ica.

Green is the pr edominant color in and

r espect ively, of the trai ts of G . incip iens , G . p latysoma,and G . m icr ones ica. Green colorat ion is found on on ly895, and r espect ively , of the trai ts in G . insular is , G .

sm ithi i , and H . glyp tocer cus (see Reaka and Mann ing,1987b, for detai ls ) . Thus , we conclude that fish pr edat ionis likely to be a selective factor promoting drab green ishco lor ation and speck ling and banding patterns that camouflage the individual agains t algae and the grainy calcareoussubs tr ate.

On the other hand , species w i th elaborate aggress ive

displays (see sect ion on behav ior ) bear the mos t br ightlyco lored s tr uctu r es ; 6296 and respect ively , of the

trai ts in G . smithi i and G . insular is , but onlyand r espect ively , of those in nonagress ive species (G .

incip iens , G . m icr ones ica, and G . p latysoma, respect ively)include predominantly r ed and yel low hues . (See Reaka

and Mann ing, 1 987b, for analys is .) H ap tosqu i lla glyp to

cercus is one of the mos t uncolorfu l species , w i th only 9%of Its tr ai ts (the antennu les) bear ing br ight coloration ; indiv iduals of this species show extr emely aggress ive behaviorbu t lack complex displays .

Simi lar ly , br i ght co lor s are concentrated an anter iors tructures associated wi th thr eaten ing and attack ing behavior (see Table 2 and quanti fied data in Reaka and Man

ning , 1987b). Red, maroon , or orangish an tennules are

found in G . sm ithi i , G . m icr ones ica, G . p latysoma, H . glyp

tocer cus , and G . incip iens , r es pectively . Red antennae ar e

fr equen t in G . insular is , G . sm ithi i , and G . p latysoma.

Gonodactylus insular is and G . sm ithi i usual ly bear red or

yellowi sh antennal scales . Max i llipeds often ar e yellowish inG . incip iens , G . insular is , and G . sm ithi i . Gonodactylussm ithi i bear s a red meral spo t , and the carpus and propodus in G . m icr ones ica and G . p latysoma are r ed. Br ightco lor s also character i ze poster ior s tr uctur es used in defens ive combat (red uropodal spo ts in G . sm ithi i ; red or yel

low ur opodal endopods and exopods in G . insu lari s and G .

smithi i , r espect ively ; red telson car inae in G . m icr ones ica) .

Co lor polymorphi sm of the body is greates t in

G . sm ithi i and G . insular is ; G . in sular is and G . incip ienshave the mos t co lor pol ymorph ic appendages . Ther efor e,

co lor po lymorphi sm among ind ividuals in the populat ion i sgr eates t ei ther in species wi th elaborate aggress ive com

mun icat ion graded di splays in G . ins ular is ,G . sm ithi i) or in species that occur in assoc iat ion wi ththese species (G . incip iens ) . These obser vations sugges tt hat co lor pol ymor phi sm may fun ction for indi vi dual r ecogni tion of aggr ess ive communi ca ti on .

Several devian t co lor pat ter ns , par t i cular l y in

G . p latysoma, wer e found on the nor ther n i s land of

Aomon , whi ch was closer to sour ces of radiati on fr om

atomic tes ting than many of the southern i s lands in the

ato l l . Dev iat ions in the red color at ion of the anter ior rap

tor ial appendage, in the blue spots on the orange backgr ound of the basal segment of the uropod , and in the

sequence of concen tr i c color s on the lateral “

eye spo ts ”

were observed in the populat ion of G . p latysoma col lectedfr om Aomon . Compar i sons wi th other populations of th i sspecies col lected fr om far ther south on the ato ll r evealedthat these dev iat ions were not explained by body s i ze, sex ,

or habi tat .

BEH AVIOR

Stomatopods ar e h ighly v isual , act ive predator s thathave some of the mos t complex behav ior known in inver

tebrates (Caldwel l and D ingle, 1 975 ; Reaka, 1980b; Reaka

and Mann ing , The gonodactylids and p r otosqu i ll ids

on Enewetak (Table 1 ) inhab i t pr eformed bur r ows in coral

or calcareous subs tr ate that cannot be enlarged ; thesespecies show the mos t intense and complex aggr es s ive

behav ior . Member s of the genus Pseudosqu i lla also showrelat ivel y elaborate behav ior and s trong aggr ess ion and

occupy ei ther holes in coral or excavat ions under coral .Lys iosqu i ll ids bur r ow in sand , can enlar ge thei r bur r ows ,have les s elabo r ate behavior , and can grow to ver y lar ges i zes (Caldwell and D ingle, 1975 ; Reaka, 198 1 ; Reaka and

Mann ing , 1 98 1 , 1 987a) .The second maxilli ped of s tomatopods i s enlar ged in to

a r aptor ial appendage that is used to smash and s tab cornpet i tor s for bur rows , predator s , or har d-bodied prey .Intense, poten tially lethal fight ing behav ior is obser vedamong member s of the same and di ffer en t species in bo ththe field and labor ator y . Aggr ess ive behavior s includes tr i kes (a smash ing blow wi th the hammer - li ke r aptor ialappendage or a s tab wi th the open , needle-s har p dactyl) ;a var iety of aggres s ive lunges and chases ; and gr adedthr eat displays in wh ich the raptor ial appendages ar e

lowered and spread , expos ing the conspicuous meral spo ts .

Defens ive behav ior s include co i ling the armored telson in

fr ont of the body wi th the ur opods spread wi de. In coraldwel l ing species th is s tan ce may effectively block the

en tr ance of a bur r ow agains t an intr uder . In addi tion , theseman ti s shr imps exh ibi t several r etr eat behavior s , includi ngsubmi ss ive pos tur es in whi ch the head and telson ar e

lower ed and thei r as sociated s tr uctur es ar e tu cked alongs ide or under the body .

The agon is tic behav ior of di ffer ent species of coraldwell ing s tomatopods in Enewetak has been addr essed inseveral s tu dies (Caldwel l and Di ngle, 1972, 1 977 ; Reaka ,

1 973 ; Di ngle, H ighsm i th , Evans , and Caldwell , 1 973 ;

Caldwell and a enmei s ter , We pr ov ide new dataand s yn thes ize all pr evious r esear ch on the aggress ive

behavio r of these man ti s shr imps in a compr ehens ive s tudyelsewher e (Reaka and Mann ing , 1 987b) , but we pr ovi de a

br ief s ummary here .

H ap tosqu i lla g lyp tocer cu s deli ver s the mos t str ikes , butth i s r ela ti vel y unco lor fu l species is exceptional l y pr one to

r etr eat fr om opponents and is les s li kel y to use thr eatenin gdi sp lays than G . sm it h i i o r G . in sular is . Spor ting a yellow

188 REAKA AND MANNING

mer al spot and other br ight co lor at ion , G . ins u lar is usesthe h ighes t num ber of thr eat d isplays in i t s in ter act ions ,s tr ikes relat ivel y o ften , and r arely r etr eats in a contes t ;consequentl y , th is species can be class i fied as a ver yaggr es s ive species . G onodactylus sm ith i i also is h ighlyaggr ess ive, s tr ik ing but also retr eat ing qu i te fr equen tly .Mem ber s of th i s species per form some of the mos t expans ive threat d isp lays , thus expos ing one of the mos tdramat ic mer al spots (carmine sur r ounded by black and

wh i te concen tr ic ci r cles ) seen in any s tomatopod . Interspec i fic tes ts ind icate that H . g lyp tocercus dom inatesG . ins u lar is , and that G . insu lar is w ins con tes ts w i thG . sm ith i i . Gonodactylus inc ip iens and G . p latysoma ar e

char acter i zed by low levels of fighting act iv i ty and subordi

nate soc ial r elat ionsh ips to the other spec ies in Enewetak .

H AB ITAT

Field wor k in 1 972 al lowed u s to quan t i tat ively analy zethe d i s tr ibut ion of shal low water s tomatopods from sever almajor types of habi tat in Enewetak (Reaka, 1 973 ; Reaka

and Mann ing , 1 987b) . These habi tats include:1 . Dead coral r ubble2 . Soft r ubble and vermet id r ocks3 . The calcareous bases of

“

m icroatolls (co lon ies of

Por ites in wh ich the cen ter has d ied and often was r iddledw i th bor ing or gan i sms )4 . Open r eef benches of conso l idated calcareous accre

t ion (Table 3 )D ingle et al . Ca ldwel l and D ingle and

Caldwel l and B r unenmeis ter (1975) prov ide add i t ionalin format ion about the occur r ence of s tomatopods in par t ieular hab i tats on Enewetak .

Cor al r ubble habi tat i s found on the r eef plat form as

isolated heads , in smal l patch r eefs , or along the reefward

algal r idge on the eas t s ide of Enewetak . These habi tatsusual ly ar e m deep and con tain G . inc ip ien s ,G . m icr ones ica

, and H . g lyp tocer cus . Stomatopods alsoare col lected fr om coral r ubble in somewhat deeper habitats (E l m deep) in the channel between Enewetak and

the sandy is lets on the nor theas t s ide of Enewetak ,between these sandy i s lets and Japtan , and on the

nor theas t s ide of Japtan . These r ubble habi tats con tainG . inc ip iens , G . m icrones ica, G . p latysoma (small),G . sm ith i i (smal l) , and H . g lyp tocer cu s . In add i t ion to the

above species , a populat ion now class i fied as G . in su lar is(Mann ing and Reaka, 1 982) was found in the r ubble on the

nor theas t fl ank of the sandy is lets (D ingle et al . , 1973 ;

Reaka, 1 980a) . By the summer of 1972 , however , ther ubble had been par tl y bur ied by sand , and no G . in sular iscou ld be found . Ou r later anal ys is sugges ts some mor pholog ica l d i ffer ences between the populat ion found in 1 971

and G . insu lar is (Mann ing and Reaka, and the pos

s ibi l i ty remains that the former populat ion r epresen ts an

addi t ional (poss ibly now ext inct? ) species in the closelyrelated G . fa lcatus l ineage . Of the spec ies found in deadcoral r ubble, G . incip iens i s the mos t common (4596 of the

40 ind iv iduals co l lected) , fo llowed by H . g lyp tocer cus

smal l G . sm i th i i G . m icrones ica and

smal l G . p latysoma

Stomatopods inhabi t soft r ubble and vermetid r ocks onthe shor eward and reefward s ides m deep) of themoat on the nor th s ide of Aomon . In 1 972 G . insu lar isinhab i ted on ly the shor ewar d s ide of th is moat mdeep) . Gonodactylus incip iens and smal l G . p latysoma and

G . sm ith i i also ar e found here; the di s tr ibut ion of smallG . sm ith i i extends into the somewhat deeper r eefward

ar eas of the moat . Of the s tomatopods co l lected fr om theAomon moat (N G . in su lar is i s the mos t common

fo l lowed by smal l G . sm ithi i G . incip ien sand small G . p latysoma

Stomatopods also can be co l lected fr om the bases of

the m icr oatolls at depths of to m on the reef flatnor theas t of Enewetak . Gonodactylus sm i thi i is abundan t(7296 of the 43 ind iv iduals quan t i fied) and reaches lar gebody s i zes in th is habi tat . A lthough thei r number s wer enot thoroughly quant i fied , ou r field notes indi cate thatG . p latysoma also attains large s i zes and is

“

abundant . ”The smal ler spec ies occur r ing in th i s habi tat , G . incip iensand H . g lyp tocercus , also reach r elatively large s i zes butare less common (9% each of the quant i fied ind iv iduals ;however , quan t i ficat ion was less s ys temat ic in this than inother habi tats , and fu r ther sampl ing is needed) . Ca ldwel land D ingle (1 975) sugges t that large indiv iduals of

G . p latysoma occupy r elatively open caver ns underm icroato lls , wh i le indiv iduals of G . sm ith : and the smal lerspecies inhabi t bor ings fu r ther ins ide the cen ter of the

Por ites co lony . Desp i te thei r abundance, ind iv iduals of

G . sm i th i i are r ar ely obser ved away from thei r pr otectivecoral colony (Reaka, per sonal obser vat ion ) .Ind iv iduals of two species , G . p latysoma and G . in

cip ien s , ar e common ly obser ved mov ing r ap idly about onthe open r eef adjacen t to the m icroato lls , par t icular ly onlow or incom ing t ides . As in the m icr oatolls where theyfind r efuge, the G . incip iens and G . p latysoma obser vedon the open reef flat ar e relat ivel y large in s i ze. Ou r fieldnotes ind icate that indiv iduals of G . p latysoma are par t icu

lar ly abundan t in th is habi tat . The shal low reef benchesm deep) on the eas t s ide of Enewetak and the eas t

s ide of Japtan also y ield s tomatopods . These r eef benchesar e scour ed by wave act ion and fr equen tly are exposed atlow t ides . Stomatopods (G . incip ien s , H . g lyp tocercus) are

co l lected fr om tubular ho les in the so l id r eef subs tr ate or

as they dar t about in the t ide pools at low t ide.

Gonodactylu s incip iens was the mos t common s tomatop'odin th is habi tat or 7896 of the quant i fied indiv iduals) ,although fur ther sampling is needed to assess the relat iveabundance of H . g lyp tocer cus .

In 1 972 a heavy r ainfal l dur ing a m id-day low t idecaused a mass ive r eef k i l l in th i s shal low exposed habi tat(Reaka, 1 980a; Lev i ten and Kohn , Desp i te thei rh igh act iv i ty levels and pr obable h igh metabo l ic rates ,s tomatopods are remar kably to lerant of har sh phys i cal condi t ions (low 0 2, h igh temper atu r es , etc . ; Reaka, per sonalobser vat ion) . Dur ing this reef k i l l , mor tal i ty among the

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1 88 BEAKA AND MANNING

mer al spo t and other br i ght co lor at ion , G . insular is usesthe h ighes t num ber of thr eat d isplays in i t s in teract ions ,s tr ikes r elat ively often , and rar el y r etr eats in a contes t ;consequent ly , th is spec ies can be class i fied as a ver yaggr ess ive spec ies . G onodactylus sm i th i i also i s h ighlyaggress ive, s tr ik ing but also r etr eat ing qu i te fr equen tly .

Mem ber s of th i s spec ies per form some of the mos t expans ive thr eat d i splays , thus expos ing one of the mos td r amat ic meral spots (carm ine su r r ounded by black and

wh i te concen tr ic c i r cles ) seen in any s tomatopod . In terspeci fic tes ts ind icate that H . g lyp tocer cus dom inatesG . in su lar is , and that G . insu lar is w ins con tes ts w i thG . sm i th i i . G onodactylus incip iens and - G . p latysoma are

character i zed by low levels of fighting act iv i ty and subordi

nate social r elat ionsh ips to the other species in Enewetak .

H AB ITAT

Field wor k in 1 972 al lowed us to quan t i tat ively analy zethe d is tr ibut ion of shal low water s tomatopods fr om sever almajor

_

types of habi tat in Enewetak (Reaka, 1 973 ; Reaka

and Mann ing , 1 987b) . These habi tats include:1 . Dead cor al r ubble2 . Soft r ubble and vermet id r ocks3 . The calcar eous bases of

“

m icroatolls (co lon ies of

Por ites in wh ich the cen ter has d ied and o ften was r iddledw i th bor ing organ i sms )4 . Open r eef benches of consol idated calcar eous accre

t ion (Table 31 )D ingle et al . Ca ldwel l and D ingle and

Caldwel l and B r unenmeis ter ( 1975) prov ide add i t ionalin format ion about the occur r ence of s tomatopods in par t ieular habi tats on Enewetak .

Coral r ubble habi tat is found on the reef plat form as

i so lated heads , in smal l patch r eefs , or along the r eefward

algal r idge on the eas t s ide of Enewetak . These habi tatsusual l y are m deep and con tain G . incip ien s ,G . m icr ones ica, and H . g lyp tocer cus . Stomatopods alsoar e co l lected fr om coral r ubble in somewhat deeper habitats m deep) in the channel between Enewetak and

the sandy i s lets on the nort heas t s ide of Enewetak ,between these sandy is lets and Japtan , and on the

nor theas t s ide of Japtan . These r ubble hab i tats containG . inc ip iens , G . m icrones ica, G . p latysoma (smal l) ,G . sm ith i i (smal l) , and H . g lyp tocer cus . In addi t ion to the

above s pec ies , a populat ion now class i f ied as G . insu lar is(Mann ing and Reaka,

1 982) was found in the r ubble on the

nor theas t flank of the sandy is lets (D ingle et al . , 1973 ;

Reaka, 1 980a) . By the summer of 1972 , however , ther ubble had been par t ly bur ied by sand , and no G . insular iscould be found . Ou r later analys is sugges ts some morpholog ical d i ffer ences between the populat ion found in 1 971

and G . ins u lar is (Mann ing and Reaka, and the pos

s ibi l i ty remains that the fo rmer populat ion r epr esen ts an

add i t ional (poss ibly now ext inct? ) species in the closel yr elated G . fa lcatus l ineage. Of the species found in deadcor al r ubble, G . incip iens is the mos t common (4596 of the

40 indiv iduals co l lected) , fo l lowed by H . g lyp tocer cus

smal l G . sm ith i i G . m icrones ica and

smal l G . p latysoma

Stomatopods inhabi t soft r ubble and vermetid r ocks on

the shor ewar d and r eefward s ides m deep) of themoat on the nor th s ide of Aomon . In 1 972 G . insu lar isinhab i ted on l y the shorewar d s ide of th is moat mdeep) . Gonodactylus incip ien s and smal l G . p latysoma and

G . sm i th i i also are found her e; the d is tr ibut ion of smal lG . sm i th i i extends into the somewhat deeper reefward

ar eas of the moat . Of the s tomatopods col lected fr om the

Aomon moat (N G . insu lar is i s the mos t commonfo llowed by smal l G . sm i thi i G . in cip iensand smal l G . p latysoma

Stomatopods also can be col lected fr om the bases of

the m icroatolls at depths of to m on the reef flatnor theas t of Enewetak . G onodactylus sm ithi i is abundant(72% of the 43 ind iv iduals quan t i fied) and reaches lar gebody s i zes in th is habi tat . A lthough thei r num ber s wer enot thor oughly quan t i fied , our f ield notes indicate thatG . p latysoma also attains large s i zes and is

“

abundant . ”The smal ler species occur r ing in th is habi tat , G . incip iensand H . g lyp tocer cus , also reach r elat ively large s i zes butar e less common (9% each of the quant i fied ind ividuals ;however , quant i ficat ion was less s ys temat ic in th is than inother habi tats , and fur ther sampl ing i s needed) . Ca ldwelland D ingle (1 975) sugges t that large ind iv iduals of

G . p latysoma occupy r elatively open caver ns underm icroatolls , wh i le ind iv iduals of G . sm ithi i and the smallerspecies inhabi t bo r ings fur ther ins ide the cen ter of the

Por ites colony . Despi te thei r abundance, ind iv iduals of

G . sm i th i i ar e r ar ely obser ved away fr om thei r pr otect ivecor al co lony (Reaka, per sonal obser vat ion ) .Ind iv iduals of two spec ies , G . p latysoma and G . in

c ip ien s , are common ly obser ved mov ing r ap idly about onthe open r eef adjacent to the m icroatolls , par t icular ly onlow or incom ing t ides . As in the m icr oatolls where theyfind r efuge, the G . incip iens and G . p latysoma obser vedon the open reef flat are r elat ively large in s i ze. Ou r fieldno tes ind icate that ind iv iduals of G . p latysoma are par t icu

lar ly abundan t in th i s hab i tat . The shal low reef benchesm deep) on the eas t s ide of Enewetak and the eas t

s ide of Japtan also y ield s tomatopods . These reef benchesare scour ed by wave action and fr equently are exposed atlow t ides . Stomatopods (G . incip iens , H . g lyp tocer cus) are

co l lected from tubular ho les in the so l id r eef subs trate or

as they dar t about in the t ide pools at low t ide.

Gonodactylu s incip iens was the mos t common s tomatopodin th is hab i tat or 78% of the quant i fied indiv iduals) ,although fur ther sampl ing i s needed to asses s the r elativeabundance of H . g lyp tocercus .

In 1 972 a heavy r ainfal l dur ing a m id-day low tidecaused a mass ive r eef k i l l in this shal low exposed habi tat(Reaka, 1 980a; Lev i ten and Kohn , Desp i te thei rh igh activ i ty levels and probable h igh metabo lic r ates ,s tomatopods are remar kably to lerant of har sh phys i cal condi t ions (low 0 2, h igh temper atur es , etc . ; Reaka, per sonalobser vat ion ) . Du r ing th is reef k i l l , mor tal i ty among the

190 BEAKA AND MANNING

s tomatopods was low compar ed to that in other cr us tacean s and gas tr opods (Reaka and Mann ing ,

1 987b) .

Ou r field s tud ies show that cons iderable m icr ohabi tatover lap , par t icular ly fo r ind ivi duals of s im i lar body s i zes ,occur s among the spec ies of coral-dwel l ing s tomatopods of

Enewetak (Table G onodactylus incip iens i s the mos tubiqu i tous spec ies , and H . g lyp tocercus , G . sm ith i i ,G . p latysoma, G . m icr ones ica, and G . insu lar is , r espec

t ively, occupy increas ingly res tr icted habi tats (number of

s i tes and habi tat type) . Of the 10 s i tes samp ledcompr ehen s ivel y in 1 972 , the mos t common co occur r ingspecies assem blage includes G . incip iens , G . sm ithi i ,G . p latysoma. A lthough not ver y common , G . m icrones ica

occu r s in th is assemblage as wel l . When G . insu lar is is

pr esen t , i t also i s a member of the G . incip ien s , G .

sm ith i i , G . p latysoma as semblage . A lthough H . g lyp tocer cus

fr equen ts a w ide r ange of hab i tat types , th i s spec ies co

occur s mos t often w i th G . incip iens alone, thus form ing a

second majo r species as sociat ion .

ACKNOWLEDGMEN TSF ield wor k was suppo r ted by gran ts fr om the Pauley

Foundat ion and the Enewetak Mar ine B io log ica l Labo r ator y to M . L . Reaka. We thank L . B . H o lthu i s for iden t i fying and D . M . Devaney for informing us of the new r ecor dof Pseudosqu i lla om ata . We ar e grateful to A . J . Kohn forhelpful d iscuss ion , and M . L . Reaka thanks E . O . W i lsonfor pr ov id ing space and fac i l i t ies at the Museum of Com

parat ive Zoo logy , H ar var d Un iver s i ty , dur ing manuscr iptpr epar at ion .

REF EREN CES

Caldwel l , R . , and S. B r unenmei s ter , 1975 , Agon i stic Behav ior of

Stomatopod Crus tacea, M id-Pacif ic Mar ine Lab ., An n . Rept .

,

1 975: 26-27 .

and H . D ing le, 1 972 , Aggr es s ive and Reproductive Behav iorof Stomatopod Crustaceans , En iwetok Mar . B io l . Lab .

,

1 971 — 1 972 Ann . Rept .

1975 , Eco logy and Evo lution of Agon is tic Behav ior in

Stomatopods , Naturwis sen schaf ten , 62: 214-222 .

1977 , Var iat ion in Agon i stic Behav ior Between Population sof the Stomatopod , H ap tosqu i lla g lyp tocer cus , Evo lut ion , 31 :

220-223 .

D ing le , H . , R . C . H ighsm ith , K . E . Evans , and R . L . Caldwel l ,1 973 , Inter specific Aggress ive Behav ior in T ropica l ReefStomatopods and Its Poss ible Eco log ica l Sign ificance,

Oeco log ic , 1 3 : 55-64 .

Lev iten , P . J. , and A . J . Kohn , 1 980 , M icrohabi tat Resour ce U se,

Activ ity Patter ns , and Episodic Catas tr ophe : Conus on T rop ical Inter t idal Reef Benches , Eco l . Monog r . , 50 : 55-75 .

Mann ing , R . B . , 1964 , A New West Amer ican Species of

Pseudosqu i lla (Stomatopoda) , Cr u s taceana, 6 : 303-308 .

1 971 , Two New Species of Gonodactylus (Crustacea , Stomatopoda) fr om the Pacific Ocean , Amer . Mus . Nov. , 2484:

1 - 1 1 .

and M . L . Reaka, 1 979 , Thr ee New Stomatopod Crustaceans from the Pacific Coas t of Costa R ica , Proc . B io l . Soc .

Wash . , 92: 634-639 .

and M . L . Reaka, 1 98 1a, Gonodactylus aloha, a New Sto

matopod Crustacean fr om the H awai ian Is lands , J . Crus t .B io l . , 1 : 1 90-200 .

and M . L . Reaka, 1 98 1 b, Gonodactylus s iamenes is , a New

Stomatopod Crustacean fr om Thai land , Proc . B io l . Soc .

Wash . , 94: 479-482 .

and M . L . Reaka, 1 982, G onodactylus insu lar is , a New

Stomatopod Crustacean from Enewetak Ato ll , Pacific Ocean ,

Proc . B io l . Soc . Wash . , 95: 347-35 1 .

Reaka, M . L . , 1 973 , Var iation in Aggress ive Behav ior in Popu lations of G onodactylus falcatus (Stomatopoda : Crustacea) fr omPopulat ions in Iso lation and in Sympatr y wi th Congener s ,En iwetok Mar . B io l . Lab . , Ann . Rept . 1 972— 1 973 , pp . 51 -54 .

1 975a, The Eco logy and Evo lut ion of Mo lt ing , L ife H is tor yPatter n s , and Agon is t ic Behavior in Stomatopod Crus tacea,

PhD . thes is , Depar tment of Zoo logy , Un iver s ity of Ca l ifor n ia,

Berkeley .1 975b, Mo lt ing in Stomatopod Crus tacea: Stages of the Mo ltCycle, Setagenes is , and Mo rpho logy , J . Morpho l . , 1 46 : 55 80 .

1 976 , Lunar and T idal Per iodici ty of Mo lting and Reproduc

t ion in Stomatopod Crustacea; a Selfish H erd H ypothes is ,B io l . Bu l l . , 150 : 468-490 .

1 978 , The Effects of an Ectoparas it ic Gastropod ,Caledon iella mon tr ouz ier i , upon Mo lt ing and Reproduction of

a Stomatopod Crustacean , Gonodacty lus vi r idis , Ve liger , 21251 -254 .

1 979a, The Evo lutionar y Eco logy of L ife H istor y Patter ns inStomatopod Crustacea, S. E . Stancyk (Ed ), Reproduct iveEco logy of Mar ine Inver tebrates , U n iver s i ty of South Car o l inaPress , Co lumbia.

1 979h, Patter ns of Mo lting Frequencies in Coral—Dwel l ingStomatopod Crustacea, B io l . Bu l l . , 156 : 328-342 .

1 980a,Geog raph ic Range, L ife H i stor y Patter ns , and Body

Size in a Gui ld of Cor al-Dwel l ing Man tis Shr imps , Evo lut ion ,

34 : 1 0 1 0- 1 030 .

l 98ob, On Lear n ing and L iv ing in H o les by Mantis Shr imp,An . Behav , 28 : 1 1 1

- 1 1 5 .

1 98 1 , The H o le Shr imp Sto r y , Nat . H is t 90: 36-43 .

and R . B . Mann ing , 1 98 1 , The Behav ior of StomatopodCrustacea, and It s Relation sh ip to Rates of Evo lution ,

J . Cr us t . B io l . , 1 : 309-327 .

and R . . B . Mann ing , 1 987a, The Sign ificance of Body Si ze,

Lar val D isper sal,and H abitat for Rates of Morpho log ica l Evolu t ion in Stomatopod Crustacea, Sm ith . Con t r . Zoo l . , 448 :1 -46 .

and R . B . Mann ing , 1 987b ,The D istr ibutional and

Behav ioral Eco logy , and the Eco logy of Co lor Patter ns in theStomatopod Crustacea from Enewetak Ato l l , Sm i th . Con tr .

Mar . Sc i . , (in pres s ) .

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192 TITGEN

TABLE 1 (con t'd)Fam i ly IBL IDAE

‘

l lbla cum ingi s ibogae H oek , 1907 .

Fam i ly LEPAD IDAEl Conchoderma aur i tum (L innaeus ,‘l Conchoderma vi rgatum (Speng ler ,il l e pas anat i/era L innaeus , 1758 .

“

l'Lepas anat ifera s t r iata de Gr aaf, 1 952 .

“

l'

Lepas anser ifera anser ifera L innaeus , 1 767 .

TLepas h i ll i (Leach ,Fam i ly MICROLEPADIDAEl M icrolepas spp .

Fam i ly POECILASMATIDAE“

I’

Octolasm is lowei (Darwin ,

“

l’

Tr i lasm is fissum (Darwin ,

Fam i ly SCALPELLIDAEl L ithotrya n icobar ica Reinhardt, 1850 .

Subo rder BALANOMORPH ASuper fami ly CH TH AMALOIDEAFam i ly CH TH AMALIDAESubfam i ly EURAPH IINAETEuraphia hembeli Conrad, 1 837.

¢Euraphia in ter texta (Darwin ,

Super fam i ly CORONULOIDEAFam i ly CORONULIDAESubfam i ly CHELONIBIINAE

‘

l'Chelon ibia caretta (Spengler ,TChelon ibia tes tudinar ia (L innaeus ,Subfam i ly CORONULINAETCoronu la diadema (L innaeus ,Subfam i ly PLATYLEPAD INAETPlatylepas decorata Darwi n , 1 854.

‘l'

P latylepas hexas ty los (Fabr icius ,Fam i ly TETRACLITIDAESubfam i ly TETRACL ITELLINAE

“

l’

Tetr acl i tella divisa (N i lsson—Can tel l ,Subfam i ly TETRACLITINAEi Tesseropora pacif ica (Pi lsbry,

Super fam i ly BALANOIDEAFam i ly ARCH AEOBALAN IDAESubfam i ly ARCHAEOBALAN INAE

‘

l‘Acas ta spp .

Fam i ly BALAN IDAEBalanus amph it r i te amph i tr ite Darwin , 1854 : Zul lo et al . , 1 972 .

Balanus ebum eus Gould , 1 841 : Zullo et al . , 1972 .

‘

I’ Balanus tr igonus Darwin , 1 854 .

TMegabalanus t in t innabu lum (L innaeus ,Fam i ly PYRGOMAT IDAESubfam i ly PYRGOMATINAE

‘

l‘Can tell ius spp .

‘

l‘H i r oa s tubbings i Ross and Newman , 1 973 .

TNobia spp .

Zl Savign ium cr enatum (Sower by ,TSavign ium den tatum (Darwi n ,

‘I‘Savign ium elongat um (H i ro ,

‘

I‘Savign ium m i llepo r um (Darwin ,

Order ACROTH ORACICASuborder PYGOPH ORAFam i ly CRYPTOPH IALIDAE

‘l‘Cryp tophialu s cordylacis Tom l inson , 1 969 .

Crytophialus heterodon tus Tom l inson , 1 969 ; Tom l inson , 1 973 .

‘I'Cryp tophialus r oss i Tom l inson , 1 973 .

“

( Species l ikely to be found at Enewetak Ato ll .t ew Enewetak Ato l l r ecords . (Th is table con tinued on next page )

CIRRIPEDIA 193

TABLE 1 (con t’d)Fam i ly LITHOGLYPTIDAEl Balanodytes talwanus Ut inom i , 1950 .

1 Kochior ine u lu la Tom linson, 1973 :Tt hog lyptes bicornis Aur ivi ll ius , 1 892.

L ithoglyptes sp inatus Tom l inson and

Newman, 1960; Tom l inson , 1973 .

L ithoglyp tes mitis Tom l inson , 1969: Tom l inson ,

Tt hoglyptes wilson i Tomlinson , 1969.

‘l'Weltner ia r eticu lata Toml inson , 1969 .

Order RH IZOCEPH ALASuborder KENTROGONIDAFami ly SACCULINIDAE

‘l’Sacculina actaeae Guer inG anlvet , 1 91 1 .

‘l'Saccu lina bipunctata Kossmann , 1872.

‘l‘Sacculina carp illae Guer i n-Ganlvet , 191 1 .

1'Saccullna lncons tans Boschma, 1 952 .

‘I'Saccullna punctata Boschma, 1 934 .

¢Saccul lnid sp .

‘Systematic hierarchy taken fr om Bowman and Abele

‘l'Specles l ikely to be found at Enewetak Atol l .zi New Enewetak 'Atol l records .

ASCOTHORACICA

Ascothoracicans general ly have been cons idered the

most pr im i tive barnacles , but there is argument to separatethem as a subclass coor d inate w i th the Ci r r ipedia (Grygier ,

The bases for the argument are that the Ascothor aclca have naupl i i that lack “

frontolateral horns ; a relativelyanamorph ic rather than h ighly metamorphic development ;an ascothoracid larva capable of feeding at all stagesrather than a s ingle cypr id stage lacking a complete gu tand functional mouthpar ts ; whol ly pr ehensi le fi r st antennaerather than ones also provided w i th cement glands ; andnatatory thoracic limbs that show no indication of ever

having been used as ci r r i . Fur thermore, i t has r ecen tlybeen shown that the ascothor aclcan sperm , the most gen.

erallzed known for the Cr ustacea, is dist inct from that ofthe remainder of the c i r r ipeds”

(Newman , Thoughthe ev idence is s trong for the separation of the Ascothor a

cica from the Ci r r ipedia, no consensus exis ts . It is includedherein for th is reason and because only one species hasbeen repo r ted fr om the Mar shal l Is lands (U t inom i ,Ascothoracicans are ecto and endoparas i tes of var ious

anthozoans and ech inoderms . Often h ighly mod i fied for a

paras it ic exis tence,they show li tt le resemb lance to the

“ typical” thoraci c bar nacle. They lack calcar eous p lates , donot cement themselves to the subs tratum , and r etain morecharacter is tics of the free-l iv ing ances tor s of the ci r r ipedsthan any other group of cr ustaceans .

Gorgonolaureus bikin iens is U t inom i , 1 962 , is the s ing leascothoraclcan repor ted fr om the Mar shal l Is lands .

Descr ibed as a new genus and spec ies , i t is known on lyfrom B ik ini Atol l and is repor ted to be a par as i te on the

gorgonacean Paracis squamata. Th is shal low-water gor

gon ian has not been r epor ted fr om Enewetak Ato l l (Lang

and Devaney , th is vo lume) . The on ly other descr i bedcongener ic species is Gorgonolau reus muz ikae G rygier ,

198 1 , repor ted from deep water in H awai i (G rygier ,1 98 1b).

TH ORACICA

Thor acicans include the typ ica l bar nacles usual l y r ecogn ized by thei r form and the presence of calcar eous plates .

There are both fr ee-l iv ing and symbiot ic taxa, and they canbe d iv ided morpholog ical ly into pedunculate and sess i leforms . The pedunculate barnacles (Lepadomorpha) are

composed of a peduncle that attaches to the subs tratumand a cap i tulum that is usual ly sur r ounded by calca reousplates . The ses s i le bar nacles (Ver r ucomorpha and

Balanomorpha) lack the peduncle, and the aper tur e is

guar ded by an oper culum formed by one or two pai r s of

calcareous plates .

Man y thoracic bar nacles have w ide d is tr ibut ionalr anges . Th i s is due in par t to the major i ty hav ing plankton ic lar val s tages (Newman and Toml inson , 1 974) and

some , such as sever al ocean ic lepadomorph spec ies , natural l y att ach ing to float ing debr i s . In add i t ion , some of the

natur al d i s tr ibut ions have been greatly extended becausecer tain species tend to “ foul” sh ip bottoms and float ingobjects and can be car r ied gr eat d is tances .

Six of the n ine bar nacle species r epor ted fr omEnewetak Ato l l ar e thoracicans . Th is h igh per centage is

not unusual because thoracicans ar e general l y larger thanspecies in the other bar nacle or der s and have a more obv i

ou s“ bar nacle appearance, thus mak ing them mor e r ecog

n i zable and l iable to be co l lected . The other thr ee spec ies ,belong ing to the Acr othoracica , ar e smal l , bur row in cal

c ium car bonate, and do not have the typ ical barnacle

194 TITGEN

appearance. They wer e found because Toml inson ( 1973 )in ten t ional ly looked for thei r bu r r ows in mo l lusk shel ls .

ACROTH ORACICA

Acr othorac icans ar e smal l , nonpar as i t ic naked bar nacles that bur r ow pr imar i ly in cal car eous subs tr ata . Thesesubs tr ata include l ive coral , coral r ock , and mo l lusk shel ls .

Sexes ar e separ ate , and the bu r r ows ar e cr eated by thefemales . The dwar f males ar e s hor t- l ived , non

-feedingor gan i sm s (Tom l in son , 1969) that are usual ly found near

the attachment d i sc of the female.

Newman and Toml inson (1 974) presen ted an in ter es ting ques t ion on the d isper sal mechan i sms of acr othor acican

bar nacles . The adults ar e poor d isper ser s because they areob l igate inhabi tan ts of ca lcium car bonate . Ther efore,

i t i s

d i ff icu lt to exp lain the d is tr ibut ion of spec ies such as

L i thog lyp tes bico r n is Au r iv i llius , 1 892 , wh ich is found inthe Car ibbean Sea , the Red Sea, Aus tr al ia, Japan , the

Mar shal l Is lands , and the L ine Is lands . The d i fficulty ar isesbecause the naup l iar d i sper sal s tages are usual l y passed inthe egg , and the cypr ids are non

-feed ing and weak swimmer s (Newman and Toml inson ,

A s ingle paper has been publ ished r epo r t ing on

acr othor acicans fr om Enewetak Atol l (Toml inson ,

Not co l lected as bar nacle mater ial , the specimens wer edi scovered in shel ls of the tu r bin id snai l Tu r bo argyro

s tom us L innaeus , 1 758 , housed at the U . S. Nat ionalMuseum of Natur al H i s tor y ’

s D iv is ion of Malaco logy .The th ree species of acr othor acican bar nacles r epo r ted

fr om Enewetak Ato l l have r elat ivel y w ide di s tr ibut ions con

s ider ing thei r pr esumed l im i ted power s of d isper sal .Cr yp top hialus heterodon tus Toml in son , 1 969 , has a

wes ter n Paci fic d i s tr ibut ion . and is known from Aus tr al ia ,

the Mar shal l Is lands (Tom l inson , and Ok inawa(Tom l inson , L i thog lyp tes m it is Toml inson ,

1 969,

appear s to have an insular d i s tr ibut ion , hav ing been co l

lected. at F i j i (Tom l inson , the Loyalt y Is lands ,the

Mar s hal l Is lands , New Caledon ia , N inafou ls landand Samoa (Tom l inson , As ment ioned ,

L i thog lyp tes b ico r n is Au r iv i ll iu s , 1 892 ,has an unusual l y

w ide d i s tr ibut ion fo r an acro thorac ican bar nac le. Over par tof i t s r ari ge i t i s known by i t s s ynonyms L i thog lyp tes sp i

natus Toml inson and Newman , 1 960 , and L i thog lyp tes

ampu l la Au r iv i ll iu s , 1 892 (Newman and Toml in son ,

On ly a few spec ies of acr otho r ac ican s have beenr epor ted fr om the Mar s hal l , Car ol ine , and G i lber t Is lands(Tom l inson , 1 969 , 1 973 ) that have not also been found atEnewetak Ato l l . Due to the cr ypt ic habi tat of bur r ow ingbar nacles and the fact that they ar e seldom looked fo r , i ti s pr obab le that these and/o r o ther acr othor ac icans occurat Enewetak Ato l l .

RH IZOCEPH ALA

Rh i zocephalans ar e endoparas i tes of_cr u s taceans , pr i

mar i ly decapods . The adu lts . are h igh ly mod i fied and bearno r esem blance to other c i r r ipeds . H owever , they ar e clas

s i f ied w i th bar nacles because thei r naup l iar lar vae have thecharacter is t ic fr on to lateral hor ns , and ther e i s a typ i calcypr id lar val s tage.

There ar e no publ i shed accounts of r h i zocephalansfr om Enewetak A to l l or the Mar shal l Is lands . The on lyr ecor d is of a saccu l in id found on a male spec imen of

Ca lathea aff in is Or tmann ,1 892 , co llected at Sta . No .

JWK-41 2 , Engebi Is land , Enewetak Ato l l , found on an

Acr opor a sp . (H aig , per sonal commun i cat ion ) . Appar entl y ,no o ther r h i zocephalans wer e found on the decapods col

lected at Enewetak Ato ll (J . S . Gar t h and A . J . B r uce, per

sonal comm un icat ion) . H owever , Saccu l ina punctataBoschma,

1 934 , and Saccu l ina bip unctata Kos smann ,

1 872 ,have been r epor ted fr om the Caro l ine Is lands .

Saccu l ina carp i l iae Guer in -Gan ivet , 1 91 1 , Saccul ina incon

s tans Boschma, 1 952 , and poss ibly Saccu lina actaeae

Guer in -Gan ivet , 1 9 1 1 , have been r epor ted from the Gi lber tIs lands (Boschma, 1 953 , and an undescr i bedspec ies of Ler naeod is cu s has been r epor ted from H awai i(Edmondson , 1 946 ; Boschma,

A l though pr esen trecor ds ind icate that in fect ion by r h i zocephalans at

Enewetak Ato ll is of low inc idence, some undoubtedlyoccu r s . In gener al , l i tt le wor k has been done on r h i zocephalans in recen t year s , pr obably because of the h is tolog ical wor k necessar y for taxonom ic iden t i fi cat ion .

D ISCUSSIONThe mos t not iceable aspect of bar nacle r esear ch at

Enewetak Ato l l , and the Mar shal l Is lands in gener al , i s thepauc i ty of wo r k that has been done . The mor e obv iousnoncommensal forms have been co l lected and ident i fied ,along w i th a few of the mor e cr ypt ic species . H owever , incompar i son to the Car o l ine Is lands , the smal ler commensal ,bur r ow ing , and par as i t ic form s have been largel y m issed .

Th i s i s char acter i s t ic of s tud ies in wh ich bar nacle r esear chi s inc idental to other wor k being car r ied ou t .

Because the c i r r iped fauna of Enewetak Ato l l is so

poo r ly known ,l i ttle can be said o f i ts b iogeography . Its

fauna ,however , i s Indo-Wes t Pac i fic in or i gin , w i th some

elemen ts of the c i r cum tr op ical -temper ate fauna . EnewetakAto l l i s composed of low i s lands and is located on the

Pac i f ic P late. For these r easons the fauna can be expectedto be less d iver se than at h igh i s lands , such as the Carol ines , or is lands such as Palau and the Ph i l ipp ines that ar enot on the Pac i f ic P late (Spr inger , 1 982 ; W . A . Newman ,

per sonal commun icat ion ) . W h i le some patch iness is to beexpected , the low i s lands of the Mar shal ls would be

expected to have few , i f any, of Spr inger ’

s ( 1 982) T ype 2or 3 endem ics . H owever , some car e mus t be taken in

in ter pr et ing bar nacle endem ic i ty and d is tr ibut ional r anges .

Several of the taxa ar e so poor ly known that d i s tr ibut ionalr eco rds often ind icate wher e a cer tain amoun t of wor k hasbeen done,

no t wher e the natu ral l im i ts to d i s tr ibut ionoccur .

Man y of the spec ies repor ted fr om the Car oline Islandscou ld occur at Enewetak Ato l l . Mos t have r easonab ly w idewes ter n Pac i fic d i s tr ibut ional r anges , and the prox im i ty of

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Chapter 1 9

N onp lank ton ic Copepoda of Enewetak Ato ll

Compi led by DEN N IS M . DEVAN EY (deceased)Ber n ice P . B ishop MuseumH ono lu lu, H awai i 968 1 7The copepod fauna of Enewetak is based on s tud ies

car r ied ou t on s ymbiot ic (commensa l and par as i tic) spec iesas wel l as those found as par t of the plankton commun i ty(Chapter 20 , th is volume) . A checkl is t of copepod fauna isfound in Table 1 .

Copepods assoc iated w i th mar ine inver tebrates at

Enewetak have been r ecor ded s ince 1970 main ly thr ough

the effor ts of A . J . H umes . Twen ty-five of these copepods ,thei r hos ts , and r efer ences are given in Table 2 . Theseinclude two harpactaco id, 1 9 poeci los tomato id (as cyclopo id), and four s iphonos tomatoid (as cyclopo id) forms . As

a result of these publi cat ions , one new genus and 14 new

species were repor ted fr om Enewetak ; addi t ional speciesr emain to be descr i bed (H umes , per sonal commun i cat ion) .Par as i t ic copepods fr om fi shes include two poeci lo

s tomato id (as cyclopo id) and 20 s iphonos tomato id (as

caligo id) species , three of wh ich wer e new to science(Lew is , 1964 ; These copepods and thei r fish hos ts

TABLE 1

Checkl i s t of N onp lank ton ic Copepoda of Enewetak A tol lSubc lass COPEPODAOrder H ARPACTICOIDAFam i ly CANUELL IDAESunar is tes dardan i H umes and H o , 1 969 : H umes , 1 971 b.

Fam i ly TEGASTIDAETesgas tes acroporanus H umes , 1 98 1 : H umes , 1 98 1 .

Order POECILOSTOMATOIDAFam i ly BOMOLOCH IDAENothobomolochus gibber (Shi ino, Lewi s , 1 968 .

Pseudotaeniacan thus sp . : Lewi s , 1968 .

Fam i ly LICHOMOLG IDAEAcan thomolgu s fis s iset iger (H umes and H o ,

H umes 1 973b .

An isomolgu s insolens (H umes and H o ,

H umes , 1973b.

L ichomolgu s t r idacnae H umes , 1 972 : H umes , 1 972b.

Metaxymo lgus acu leatus (H umes and H o ,

H umes , 1 973h.

Octop icola rega lis H umes , 1 974 : H umes , 1974 .

Parador idicola ade lphus (H umes and H o ,

H umes 1973b.

Par amolgu s en iwetokens is H umes , 1 973 : H umes , 1973b.

Paramolgu s os ten tus H umes , 1973 : H umes , 1 973b.

Schedomolgus lobophor us H umes and H o , 1 968 : H umesand Stock, 1973 .

Syns tellicola acan thas ter is (H umes ,Stell icola acan thas ter is H umes , 1 970 : H umes , 1970a.

Synonomy based on H umes

Fam i ly MYICOL IDAEAn thess ius alatus H umes and Stock , 1965 : H umes ,

1972b.

An thes s ius am ical is H umes and Stock , 1 965: H umes1972b.

An thess ius so lidus H umes and Stock , 1965 : H umes ,1972h.

Fam i ly PSEUDANTH ESSIDAEPseudan thess ius coman thi H umes , 1 972 : H umes , 1 972a.

Senar iellus tens us H umes , 1 977 : H umes , 1977 .

Fam i ly SABELL IPH IL IDAEScambicor nus idoneus (H umes and Cressey ,

H umes , 1980 .

Fam i ly TAEN IACANTH IDAEClavisodal is heterocen trot i H umes , 1 970 : H umes , l 970b.

Fam i ly XARIF IDAEXar ifia brevi ramea H umes and Doj i r i , 1982 : H umes and

Do j i r i , 1982 .

Xar ifia sabiu r aens is M isaki , 1978 : H umes and Doj i r i1 982 .

Order SIPH ONOSTOMATOIDAFam i ly CALIG IDAEAnu retes ser ratus Sh l i no , 1954 : Lewis , 1968 .

Cal igu s alaih i Lewis , 1968 : Lewi s , 1 968 .

Cal igus asymmetr icus Kabata, 1 965 : Lewi s , 1968 .

Cal igu s bon ito W i lson , 1905 : Lewi s , 1 968 .

confusas Pi llai , 1961 : Lewi s , 1968 .

‘The higher class ifica tion used herein (to Fam i ly level) fo llows Bowman and Abele

(Th is table con tinued on next page.)

198 DEVANEY

TABLE 1 (con t’d)

Fam i ly CAL IG IDAE (cont ’d)Cal igu s coryphaenae Steens t r up and Lu tken , 1 861 :

Lewis , 1 968 .

Cal igus kapuhi l i Lewis , 1 967 : Lewis , 1968 .

Cal igus laticaudus Shi ino , 1 960 : Lewi s , 1968 .

Cal igus l igatus Lewis , 1964 : Lewis , 1968 .

Cal igu s productus Dana, 1 853 : Lewis , 1968 .

Cal igus pseudokalumai Lewis , 1 968 : Lewis , 1 968 .

Den t igr yps li tus Lewi s , 1 964 : Lewis , 1964 ; 1 968 .

Lepeophthei r us diss im u latus W i lson , 1905; Lewis , 1968 .

p lectropom i Nunes -Ruivo and

Fou rmano i r , 1 956 : Lewis , 1 968 .

Pseudanu r etes pomacan thi Lewis , ‘

1968 : Lewis , 1 968 .

Pseudocal igus s im i lis Lewis , 1 968 : Lewis , 1 968 .

TABLE 2

Fam i ly DISSONIDAED issonus heronens is Kabata, 1966 : Lewis , 1 968 .

D issonu s s im i lis Kabata , 1966 : Lewi s , 1 968 .

Fam i ly EURYPH ORIDAEA lebion gr aci l is W i lson , 1 905 : Lewis , 1968 .

Fam i ly NANASPIDIDAENanasp is manca H umes , 1 973 : H umes , 1 973a; 1980 .

Nanasp is pus i l la H umes , 1 973 : H umes , l 973a; 1 980 .

Nanasp is sp in ifera H umes , 1 973 : H umes , l 973a; 1980 .

Fam i ly PANDARIDAEPandar us cranchi i Leach , 1 8 19: Lewis , 1968 .

Fam i ly STELLICOMITIDAEAs troxynus cu lcitae H umes , 1971 : H umes , l 97 l a.

In ver tebr ate H os t and Sym b ioti c CopepodaRecor ded fr om Enewetak A tol l

H os ts

CN IDARIA

Octocoral l ia (Soft cor als)A lcyonaceaLobophytum pauciflor um

Lobophytum pauciflor umLobophytum pauc iflor umLobophytum pauciflor umNephthea chabr ol i i

Sto lon iferaPachyclavu lar ia vio lacea

Zoanthar iaScler act in ia (Stony corals )Acropor a flor idaAcropor a flon

’

da

Acropo ra flor idaAcropo ra flor ida

CrustaceaCoenobi to idea (H erm it cr abs )Dardanus gu ttatusDardanus lagopodesDardanus megi s tosDar danus s cu tellatus

Octop ico la r ega lis

Copepod s ymbion ts

Acan thomolgu s fis s iset iger

Refer en ces

H umes (1 973b)

H umes ( 1 974)(Th is table continued on next page . )

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200

H os tsCarang idaeCaranyx melampygu s

Car anyx melampygus?

Mu l l idaePar upeneus cyc los tomas?

ChaetodontidaeChaetodon aur igaChaetodon lunu la

Pomacanthidae

Pomacan thus imper ato rAcanthur idaeAcan thurus n igr icauda

(A . gahhm)Acan thur u s o l ivaceusNaso vlam ingi i

Scombr idaeEuthynnus aff in is(E . yaito)

G rammator cyn us bi l ineatus

DEVANEY

TABLE 3 (con t ’d)

Par as i t i c copepod

Cal igus confus as ?Cal igu s coryphaenae

Lepeophtheir u s diss im u latu s

Cal igu s kapuhi l i

Cal igus kapuhi l i

Pseudanu retes pomacan thi

Pseudotaen iacan thus sp .

Cal igu s lat icaudu s

Anur etes ser r atu s

Cal igu s coryphaenae

Den t igryp s li tusD is sonas heronen s is

D is sonus s im i lis

Refer en ces

Lewi s (1968)

Lewi s (1 968)Lewi s (1 968)

Lewi s (1968)Lewi s (1 968)Lewi s (1 968)Cr es sey and CresseyLewi s (1 968)Lewi s (1 968)Lewi s (1968)Lewi s (1 968)Lewi s (1 968)Lewi s (1 968)Lewis (1 964 ; 1 968)Lewi s (1 968)Lewis (1 968)

TCons ider ed m is identi fication of host (B ruce B . Co llette, per sonal commun ication) .

REFERENCES

Bowman , T . A . , and L . G . Abele, 1 982, Class i fication of the

Recent Crustacea, L . G . Abele The B io logy of Crus taceaVo lume 1 Sys temat ics , the Fos s i l Recor d, and B iogeography,Academ ic Pr ess , New Yor k , pp . 1 -27 .

Cr essey , R . , and H . B . Cressey , 1979 , The Paras i tic Copepods ofIndo—Wes t Pacific L i zardfi shes (Synodon tidae) , Sm ithson . Con

tr ib . Zoo l . , 296 : 1 -71 .

1 980 , Paras itic Copepods of Mackerel and Tuna-L ike Fishes(Scombr idae) of the Wor ld , Sm ithson . Con tr ib . Zoo l . , 31 1 :1 - 1 86 .

H umes , A . G . , 1970a, Stell icola acan thas ter i s n . sp . (Copepoda,Cyclopo ida) Associated wi th the Starfish Acan thas ter p lanci(L . ) at En iwetok Ato l l , Publ . Seto Mar . B io l . Lab . , 1 7:

329-338 .

1 970b , Clavisodal is heterocen tr o t i gen . n . et sp . n . , a Cyclopo id Copepod Paras it ic on an Ech ino id at En iwetok Ato ll ,J . Par as i tol . , 56 : 575-583 .

1 971a, Cyclopo id Copepods (Stellicom i t idae) Paras i tic on

Sea Star s fr om Madagasca r and En iwetok Atol l , J . Paras itol . ,57 : 1330- 1 343 .

1 971b, Sunar is tes (Copepoda, H ar pact ico ida) Ass ociatedwi th H erm i t Cr abs at En iwetok Ato l l , Pac . Sc i . , 25: 529-532 .

1 972a, Pseudan thess ius coman thi n . sp (Copepoda,

Cyclopo ida) Associated wi th a Cr i no id at En iwetok Atol l , Pac .

Sc i . , 26 : 373 -380 .

1 972b, Cyclopo id Copepods Associated with T r idacn idae(Mo l lusca , B ivalv ia) at En iwetok Ato l l , Pr oc . B io l . Soc . Wash . ,84 : 345-358 .

1973a, Nanasp is (Copepoda, Cyclopo ida) Paras itic on the

H o lothur ian Thelenota ananas (Jaeger ) at En iwetok Ato ll , J .Paras itol . , 59: 384

-395 .

1 973b, Cyclopo id Copepods (L ichomolgidae) from Octocor als at En iwetok Ato l l , Beaufort ia, 21 : 1 35-1 51 .

1974 , Octop icola r egalis n . sp . (Copepoda, Cyclopo ida,

L ichomolg idae) Associated wi th Octopus cyaneus from New

Ca ledon ia and En iwetok Ato l l , Bul l . Mar . Sc i . , 24 : 76-85 .

1 976 , Cyclopo id Copepods As sociated wi th Aster oid Ech inoderm s in New Ca ledon ia, Sm ithson . Con t r ib . Zoo l . , 217 : 1 - 1 9 .

1977, Pseudanthes i id Copepods (Cyclopo ida) Associatedwith Cr ino ids and Ech inoids (Ech inodermata) in the Tr opicalWes tern Pacific Ocean , Sm ithson . Con tr ib. Zoo l . , 243 : 1 -43 .

1 980 , A Rev iew of the Copepods Associated wi th H olothu r ians Including New Species fr om the Indc acific, Beaufor tia,

30 : 3 1 - 1 23 .

NONPLANKTONIC COPEPODA 20 1

198 1 , A New Species of Tegas tes (Copepoda, H arpac

t ico ida) Assoc iated wi th a Scleract in ian Coral at EnewetakAtol l , Proc . B io l . Soc . Wash . , 94 : 254-263 .

and M . Doj i r i , 1982 ,Xar i f i idae (Copepoda) Par as i tic in lndcr

Pacific Scleract in ian Cor als , Beaufor t ia, 32:

and J . H . Stock , 1 973 ,A Rev is ion of the Fam i ly L ichomolgi

dae Kossmann , 1877, Cyclopo id Copepods Main ly Associated

with Mar ine Inver tebr ates , Sm ithson . Con t r ib . Zoo l . , 1 27:

1 -368 .

Lewi s , A . G . , 1 964 , The Caligid Copepod Genus Den t igryps(Crustacea: Calligo ida) , Pr oc . U . 8 . Nat . Mus . , 1 15: 347-380 .

1968 , Copepod Crustaceans Paras itic on Fishes of En iwetokAto l l , Pr oc . U . 8 . Nat . M us . , 125: 1 -78 .

Ver voo r t , W . , 1 964 , Fr ee-L iv ing Copepods fr om Ifaluk Ato l l inthe Car o l ine Is lands , U . S . Nat . M us . Bu l l . , 236 : 1 -43 1 .

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204 GERBER

TABLE 1

Check l i s t of Plank ton of Enewetak A tol l

Phy lum DINOPHYTA (D INOFLAGELLATES)Class D INOPH YCEAEOr der D INOPH YSIALES

Fam i ly AMPH ISOLEN IACEAEAmph iso len ia spp . : H i r shfi eld et al . 1 957.

Fam i ly DINOPH YSIACEAED inophys is r on tundatum (Clap et Lach) : H i r shfield et al . 1957 .

D inophys is schuett i i (Mur r ay et Wh i tting) : H i r shfi eld et ai . 1 957 .

Phalacr oma per r adictyum Stein : H i r shfield et al . 1957 .

Fam i ly ORNITH OCERCIACEAEOm ithocer cu s sp lendidu s Mur ray and Wh i tting : H i r shfield et al . 1 957 .

Om ithocer cus sp .

Fam i ly CERATOCORYACEAECeratoco rys ho r r ida Stein : H i r shfi eld et ai . 1957 .

Order PYROCYSTALESFam i ly PYROCYSTACEAEPyrocys t is fus iform is Thompsen ex Mur r ay : Ger ber , 198 1 .

Pyrocys t is p seudonoct i luca Thompsen ex Mur ray : Ger ber , 1 98 1 .

D is sodin ium lunu la (Schutt) Tay lor : Ger ber , 198 1 .

Or der PER ID IN IALESFam i ly PER ID IN IACEAE

P rotoperi din ium divergens (Ehrenber g) Ba lach .Per i din ium divergens (Ehr enberg) : H i r shfield et al . 1957 .

P rotOper idin ium depressum (Bai ley) Balach .Per idin ium depressum (Bai ley) : H i r shfield et al . 1957 .

P rotoper idin ium g lobu lum (Stein) Balach .Per idin ium globu lum (Stein) : H i r shfi eld et al . 1 957 .

Per idin ium ocean icum (Vanhoffen) Balach .Per idin ium ocean icum (Vanhoffen ): H i r shfield et al . 1957 .

Fam i ly GONYAULACACEAEGonyau lax bi ros tr is (Stein) : H i r shfi eld et al . 1 957 .

Fam i ly CERATIACEAECerati um candelabr um (Ehr enber g) Stern : H i r shfield et al . 1 957 .

Cerat ium fur ca (Ehr enberg) Clap . et Lachm . : H i r shfield et al . 1 957 .

Cerat ium gibber um Gou r r et : H i r shf ield et al . 1 957 .

Ceratium lineatum (Ehrenberg) Cleve: H i r shfi eld et al . 1 957 .

Cerat ium t r ipos (O . F . Mul ler ) N i tzs ch : H i r shf ield et al . 1 957 .

Cer at ium tr ipos O . F . Mul ler var . B : H i r shfield et al . 1957 .

Phy lum SARCOD INAClass ACTINOPODEA

Order PORULOSIDASuborder ACANTH ARINA

Acan thomet ron sp . : H i r shfield et al .

Acan tholonche sp . A . : H i r shfield et al .

Acan tholonche sp . B : H i r shfi eld et al .

Acan thochiasma fus iform is H aeckel : H i r shfi eld et al .

Acan thos tau r us purpurascens H aeckel : H i r shfi eld et al .

ASp idomma sp . : H i r shf ield et al .

Dodecap s is t r icinata H aeckel : H i r shfield et al .

Doratasp is sp . : H i r shf ield et al .

H exalasp is hel iod iscus H aeckel : H i r shfi eld et al .

H exacolp us infundibu lum H aeckel : H i r shfield et al .

H exonasp is has tataz H aeckel : H i r shf ield et al .H ys t r ichasp is dor sata H aeckel : H i r shfi eld et al .

Phr actasp is sp . : H i r shf ield et al .

Quadr i lonche mesos tau r as H aeckel : H i r shfield et al .

Tes sar apelma concretu i ra H aeckel : H i r shfi eld et al .Tes sar apelma concretu i ra H aeckel : H i r shfield et al .

Zygos tau r us sag i tal is H aeckel : H i r shfield et al .

Zygacan tha sp . : H i r shfield et al . (Th is table cont inued on next page . )

LAGOON PLANKTON

TABLE 1 (con t’d)

Subo rder SPUMELLINAD ip losphaer a hexagonal is H aeckel : H i r shf ield et al .

Euchiton ia lanceo lata H aeckel : H i r shfield et al .

E . muel ler i H aeckel : H i r shfield et al .

F lus t rella sem isp i r al is H aeckel : H i r shf ield et al .

H el iod iscus as ter iscus H aeckel : H i r shfield et al .

H exalonche phi losophica H aeckel : H i r shfield et al .

H exas tylus phaenaxon ia H aeckel : H i r shfield et al .

H ymen ias tr um eucl idis H aeckel : H i r shfield et al .

Monozon ium alatum H aeckel : H i r shfield et al .

Ommatospyr is vi rgi neaH aeckel : H i r shfield et al .

t exagon ium H aeckel : H i r shfield et al .

Panar tus p luteus H aeckel : H i r shf ield et al .

Sphaerozoum sp . : H i r shfield et al .

Spongas ter tet ras Ehr enberg : H i r shfi eld et al .

Spongod iscus ve le lla H aeckel : H i r shfield et al .

Stylospongia sp . : H i r shfield et al .

Tetr apyle sp . : H i r shfi eld et al .

Or der OSCULOSIDASuborder NASSELL INA

Adelocyr t is sp . : H i r shfield et al .

A r chiscen ium sp . : H i r shfield et al .

Call im itr a agnesae H aeckel : H i r shf ield et al .

Corocalyp tr a el isabethae H aeckel : H i r shf ield et al .

D ictyocephalus sp . : H i r shf ield et al .

D ictyoceras sp . : H i r shfield et al .

Eucyr t idium hexagonatum H aeckel : H i r shfield et al .

Eucyr t idium sp . : H i r shfield et al .

Lychnocan ium pyr iforme H aeckel : H i r shfield et al .

Monos tephus sp . : H i r shf ield et al .

Pterocan ium t r icolp um H aeckel : H i r shf ield et al .

Pterocorys campanu la H aeckel : H i r shf ield et al .

Seman t is sp . : H i r shfield et al .

Theocorys sp . : H i r shfield et al .

Tr icyl idium dictyospyr is H aeckel : H i r shfield et al .

Zygos tephan ium sp . : H i r shfield et al .

Phylum CIL IOPHORAClass POLYH YMENOPH ORA

Or der OLIGOTRICH IDASuborder TINTINN INAFam i ly CODONELLIDAE

T in t innOp s is sp . : H i r shfield et al . 1957 .

Codonella cusp idata Kofo id and Campbel l : Go ld and Mor alesCodonar ia ocean ica (B randt) Kofo id and Campbel l : Go ld and Morales

Fam i ly CODONELLOPSIDAECodonellop s is amer icana Kofo id and Campbe l l : Go ld and Morales

Fam i ly CYTTAROCYCL IDAEFare l la azor ica (Cleve) Jorgensen : H i r shf ield et al . 1957 .

Fam i ly PTYCH OCYLIDAEEp ip locyclis sp . : H i r shfield et al . 1 957 .

Poroecus ap icatus Kofoid and Campbe l l : Go ld and MoralesPo roecus ap icu latus (Cleve) Cleve: Go ld and MoralesPoroecus cu r tus Kofoid and Campbel l : Go ld and Morales

Fam i ly EPIPLOCYLIDAEEp ip locyl is labiosa Kofo id and Campbel l : Go ld and MoralesEp ip locyl is pacif ica Kofo id and Campbel l : Go ld and Morales

Fam i ly PETALOTRICH IDAECraterella tor u lata Jorgensen : Go ld and MoralesPetalotn

’

cha major Jorgensen : Go ld and Mor alesFam i ly RH ABDONELLIDAE

P rotor habdonella s imp lex (Cleve) Jorgensen : Go ld and MoralesP rotor habdonella s t r iatu ra Kofo id and Campbel l : Go ld and Morales ( 1977)

205

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TABLE 1 (con t ’d)

Fam i ly RH ABDONEL L IDAE (con t ’d)Rhabdonel la amo r (Gleve) B r andt : Go ld and Mor alesRhabdonella elegen s (Jorgensen ) Kofo id and Campbel l : Go ld and Mor alesRhabdonella inflata Kofo id and Campbe l l : H i r shfi eld et al .

Rhabdonella pocu lu rn; (Os ten feld and Schm idt) : H i r shfield et al .

Rhabdoné lfop s is apophysata Cleve : Go ld and Mor ales ( 1 977)“ Fam i ly XYSTQNELL IDAE

Xys tonella sp . : H i r shfield et al .

Fam i ly UNDELL IDAEUndella hem ispher ica Laachmann : Go ld and MoralesP r op lectel la bior bicu lata H ada: Go ld and MoralesP r op lectella perpus i l la Kofo id and Campbe l l : Go ld and Morales

Fam i ly DICTYOCYSTIDAED ictyocys ta occiden tal is Kofo id and Campbel l : Go ld and Mor ales

Fam i ly TINTINN IDAEAmphor ella brandi i Jorgensen : Go ld and MoralesCan thar iella brevis Kofo id and Campbe l l : H i r shfield et al .

Dadayiella ganymedes Entz , Sr . : H i r shf ield et al .

Steens t r up iella gr aci lis (Jorgensen) Kofoid and Campbel l : Go ld and MoralesSteens tr up iella s teens t r up i i (Clapar ede and Lachmann) Kofo id and Campbel l : Go ld and MoralesT in t innus f r akno i i Daday : Go ld and Mor alesT in t innus lu sus undae var tenuis (Kofo id and Campbe l l) H ada: Go ld and MoralesT in t in nu s lu sus undae var tu rgescens : Go ld and Mor alesTin t in n us sp . : H i r shfield et ai .

Phy lum CN IDAR IAClass H YDROZOA

Or der SIPH ONOPH ORASubo rder CALYCOPH ORAFam i ly DIPH YIDAE

‘Abylop s is es chscho ltz u H ui dey : Sear s , 1 950 .

Abylop s is tetragona Otto : Sear s , 1 950 ; Ger ber ,’ 1 98 1 .

‘Agalma oken i Es chscho ltz : Sear s , 1 950 .

Bas s ia bas sens is Quoy and Gaimard: Sear s , 1 950 ; Ger ber , 1 98 1 .

Chelophyces con tor ta Lens and Van R iem sdijk : Sear s , 1 950 ; Ger ber , 1 98 1 ; Alvar i no (per sonal commun ication) .D iphyes cham is son is H uxley : Sear s , 1 950 ; -G er ber , 1 98 1 .

D iphyes dispar Cham i so and Eysenhardt : Sear s , 1 950 ; Ger ber , 1 98 1 .

‘Lens ia cono idea Kefer s tein and Ehler s : Sear s , 1 950 .

Lens ia s ubt i ll is Chum : Ger ber , 1 98 1 .

Lens ia subt i lo ides Lens and Van R iem sdijk : Ger ber , 1 98 1 .

Suborder PH YSOPH ORIDAFam i ly PH YSAL IIDAETPhysal ia ut r icu lus (Gmel in , 1 790)

Or der H YDROIDASubo rder LEPTOMEDUSAE (CALYPTOBLATINA)Fam i ly AEQUOREIDAETAequorea sp .

Subo rder CH ONDROPH ORAFam i ly VELELLIDAETVelella sp .

Fam i ly PORPITIDAETPorp ita pacifica Lesson , 1 826 .

Clas s SCYPH OZOAOr der SEMAEOSTOMAEFam i ly ULMARIDAETAu relia sp . of A . labiata Cham is so and Eysenhardt, 1 820 .

Fam i ly PELAG IDAE“

(Pelagic noct i luca (For sskal , 1 775)On ly recorded outs ide lagoon but in the immediate v icin i ty of Enewetak Ato ll .

1‘These were ident i fied in the ocean water s neighbo r ing B ikin i Ato ll (Ch iba et al . ,

(Th is table continued on next page. )

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208 GERBER

TABLE 1 (con t ’d)

Fam i ly SCOLECITRICH IDAEScolec ithr icella den tate (Giesbr echt) : Barn et t , 1 967 ; Ger ber , 1 98 1 .

Scolecithr icella marginata (Giesbr echt) : Mahnken , 1966 ; Bar nett , 1 967 .

Scolecithr icella tenu is ser ata (Giesbr echt) : Bar nett , 1 967 .

Scolecithr ix danae (Lubbock): Johnson , 1 954 ; G i lmar ti n , 1 958 ; Mahnken , 1966 ; Ger ber , 1981 .

Fam i ly TEMORIDAETemor a discaudata Giesbr echt : Mahnken , 1 966 ; Ger ber , 1981 .

Fam i ly PSEUDODIAPTOMIDAEPseudod iap tomus com u tus N icho l l : Barnett , 1967 .

Fam i ly METRIDIIDAEMetr idia sp . Boeck : H obson and Ches s , 1 978 .

P leu r omamma abdom inalis (Lubbock) : Bar nett , 1 967 .

Pleu r omamma gr aci lis (Claus ) : Barnett , 1 967 ; Ger ber , 1 98 1 .

Pleu r omamma x iph ias (Giesbr echt) : H obson and Ches s , 1 978 .

Fam i ly CENTROPAG IDAECen tr opages calan in us (Dana) : Gi lmart in , 1958 ; Mahnken , 1966 ; Bar nett , 1 967 ; Ger ber , 198 1 .

Cen tr opages elongatus Giesbr echt : Johnson , 1 954 ; Mahnken , 1 966 ; Bar nett , 1 967; Ger ber , 198 1 .

Cen tropages gr aci lis (Dana) : Barnett , 1 967; Ger ber , 1981 .

Cen trOpages or s in i i Giesbr echt : Mahnken , 1966 ; Barnett , 1967 ; Ger ber , 198 1 .

Fam i ly LUCICUTIIDAELucicu t ia flavicor n is (Claus ) : Mahnken , 1 966 ; Barnett , 1967 ; Ger ber , 198 1 .

Lucicu t ia ova lis Wo lfenden : Ger ber , 198 1 .

Fam i ly H ETERORH ABDIDAEH etero r habdus pap i ll iger (Claus ) : Barnett , 1 967 .

Fam i ly ARIETELL IDAEParam isophr ia sp . Scott : Barnett , 1967.

Fam i ly PSEUDOCYCLOPIDAEPseudocyclop s sp . B r ady : Bar nett , 1 967 .

Fam i ly CANDACIIDAECandacia aeth iop ica (Dana) : Gi lmar tin , 1 958 ; Mahnken , 1966 ; Bar nett , 1967; Gerber , 1 98 1 .

Candacia bisp inosa (Claus ): Bar nett , 1 967; Ger ber , 198 1 .

Candaela catu la (Giesbr echt) : Bar nett , 1 967 ; Ger ber , 1 98 1 .

Candacia discaudata A . Scott : Bar nett , 1967 .

Paracandacia s imp lex (Giesbrecht) : Bar nett , 1967 .

Par acandac ia t runcata (Dana) : Mahnken , 1966 ; Bar nett , 1 967 ; Ger ber , 198 1 .

Fam i ly PONTELL IDAECalanOp ia m inor A . Scott : Johnson , 1 954 ; Bar nett , 1 967 ; Ger ber , 1 98 1 .

Labidocer a laeviden tata (B rady) : Gi lmar tin , 1 958 ; Barnett , 1 967 ; Ger ber , 1 98 1 .

Pon tell ina p lumata (Dana) : Gi lmar t in , 1 958 ; Ger ber , 1 98 1Pon teIIOp s is sp . G . B rady : Gi lmar t in , 1 958 .

Fam i ly ACARTIIDAEAcar t ia fos sae Gur ney : Ger ber , 198 1 .

Acar t ia hamata Mor i : John son , 1 954; Mahnken ,1 966 ; Bar nett , 1 967 .

Acart ia neg l igen s Dana: Mahnken , 1966 ; Barnett 1967; Ger ber , 198 1 .

Acar t ia sp in icaudata Giesbr echt : Bar nett , 1 967 .

Fam i ly TORTANIDAETor tan us gr aci lis (Br ady) : Johnson , 1 954 ; Gi lmar t in , 1 958 ; Mahnken , 1 966 ; Barnett , 1967; Ger ber , 1 98 1 .

Tor tanus m u r r ayi A . Scott : Johnson , 1954 .

Order CYCLOPODAFam i ly OITH ON IDAE

Oi thona nana (Claus ) : Bar nett , 1967; Ger ber , 1981Oithona p lum ifera Ba i rd : Johnson , 1954 ; Mahnken , 1 966 ; Bar nett , 1 967; Ger ber , 198 1 .

Oi thona p seudof r ig ida (Giesbrecht) : Bar nett , 1 967; Ger ber , 1 98 1 .

O i thona r igida Giesbr echt : Bar nett , 1967; Ger ber , 1 98 1 .

O ithona tenuis Rosendor n : Bar nett , 1967 ; Ger ber , 1 98 1 .

Fam i ly CLAUSIDIIDAESaphi rella t rop ica Wo lfenden : Ger ber , 1 98 1 .

Fam i ly ONCAEIDAEOncaea media Giesbr echt : Barnett , 1 967; Ger ber , 1 98 1 .

Oncaea uen us ta Ph i l ippi : Johnson , 1 954 ; Barnett , 1967 ; Ger ber , 1 98 1 .

Lubbock ia squ i ll imana Claus : Johnson , 1 954 ; Ger ber , 1981 . (Th is table continued on next page .

LAGOON PLANKTON 209

TABLE 1 (con t'd)Fam i ly SAPPH IRIN IDAE

Sapphi r ina s tel lata Giesbr echt : Johnson , 1 954 ; Ger ber , 198 1 .

Sapph i r ina t rop ica Wo lfenden : Ger ber , 1 98 1 .

Cop i l ia m i rabi lis Dana: Johnson , 1954 ; Gi lmar t in , 1 958 ; Mahnken , 1 966 ; Barnett , 1967 ; Ger ber , 198 1 .

Cop i l ia quadrata Dana: Bar nett , 1 967 .

Fam i ly CORYCAEIDAECorycaeus agi l is Dana: Mahnken , 1966 ; Bar nett , 1967 ; Ger ber , 1 98 1 .

Corycaeus andrews i Far r an : Mahnken , 1 966 .

Corycaeus as iaticus F . Dahl : Barnett , 1 967 ; Ger ber , 1 98 1 .

Corycaeus catus F . Dahl : Bar nett , 1967; Ger ber , 1 98 1 .

Corycaeus crass iuscu lus Dana: Mahnken , 1 966 ; Bar nett , 1967 ; Ger ber , 1 98 1 .

Corycaeus flaccus Giesbr echt : Ger ber , 198 1 .

Corycaeus gr aci lis Dana: Mahnken , 1 966 .

Corycaeus lotus Dana: Ger ber , 1 98 1 .

Corycaeus lau tus Dana: Ger ber , 1 98 1 .

Corycaeus limbatus B r ady : Barnett , 1 967 ; Ger ber , 1 98 1 .

Corycaeus longis ty lis Dana: Barnett , 1967; Ger ber , 1981 .

Corycaeus medius Gur ney : Barnett , 1967 ; Ger ber , 1981 .

Corycaeus pum i lus M . Dahl : Mahnken , 1966 .

Corycaeus robus tus Giesbr echt : Mahnken , 1 966 ; Barnett , 1967 .

Corycaeus specios us Dana: Mahnken , 1 966 ; Barnett , 1 967; Ger ber , 1 981 .

Corycaeus tenuis Giesbrecht : Ger ber , 1 98 1 .

Corycaeus typ icus (Kroyer ) : Barn ett , 1 967; Ger ber , 1 981 .

Corycaeus vitreus Dana: Mahnken , 1966 ; Bar nett , 1967 ; Ger ber , 198 1 .

Far r an u la car inata (Giesbrecht) : Ger ber , 1981 .

Corycaeus car inatus Giesbrecht : Barnett , 1 967 .

F ar ran u la concinna (Dana) : Ger ber , 1981 .

Corycaeus concinnus Dana: Mahnken , 1966 ; Bar nett , 1967F ar r anu la gibbu la (Giesbrecht) : Ger ber , 1981 .

Corycaeu s gibbu lus Giesbr echt : Mahnken , 1 966 ; Barnett , 1 967 .

Order H ARPACTICOIDASubo rder OL IGOARTH RAFam i ly ECTINOSOMATIDAE

M icrosetella nor vegica (Boeck) : Barnett , 1967 .

M icrosetella rosea (Dana) : Bar nett , 1 967; Ger ber , 1 98 1 .

Fam i ly CLYTEMNESTRIDAEC lytemnes t ra ros tr ata B rady : Ger ber , 198 1 .

C lytemnes tr a scutel lata Dana: Ger ber , 198 1 .

Fam i ly MACROSETEL LIDAEMacrosetella gr aci lis (Dana) : Gi lmar t in , 1958 ; Barnett , 1967 ; Ger ber , 1 98 1 .

Fam i ly LONG IPEDIIDAELongipedia cor onata Claus : Ger ber , 1981 .

Longipedia weber i A . Scott : Ger ber , 198 1 .

Fam i ly H ARPACTICIDAEH arpact icus spp . M . Edwards : Ger ber , 1981 .

Fam i ly PELTIDIIDAEPelt idium spp . Ph i l ippi : Ger ber , 1 98 1 .

Fam i ly TEGASTIDAETeges tes sp . Norman : Ger ber , 1 98 1 .

Fam i ly TH ALESTRIDAEEudactylopus andrewi Sewel l : Ger ber , 1 98 1 .

Eudactylop u s anoma la Sewell : Ger ber , 1 98 1 .

Eudacty lop us fasc iatus Sewel l : Ger ber , 1 98 1 .

Dactylopoda sp . G . O . Sar s : Ger ber , 198 1 .

Fam i ly DIOSACCIDAEAmphias cus coral icola Sewell : Ger ber , 1 98 1 .

Amph ias cop s is cinctus (Claus ) : Ger ber , 1 98 1 .

Metamph iascop s is h ir sutus (Thompson and A . Scott): Ger ber , 198 1 .

Fam i ly METIDAEMetis sp . Phi l ippi : Ger ber , 198 1 .

(Th is table cont inued on next page . )

2 10 GERBER

TABLE 1 (con t’d)Fam i ly LAOPH ONTIDAE

LGOphon te cor nuta Ph i l ippi : Ger ber , 1 98 1 .

Order MONSTRILLOIDAFam i ly MONSTRILLIDAE

Mons tn'

lla sp . : Ba rnett , 1 967 ; Ger ber , 1 98 1 .

Subc lass MALACOSTRACAOrder MYSIDACEAFam i ly MYSIDAESubfam i ly SIRIELL INAESi r iella affin is H anson : Mur ano , 1983 .

Sir iella aequ i rem is H anson : Mur ano , 1 983 .

Sir iella sp . : H obsen and Chess , 1 978 .

Subfam i ly MYSINAEAn isomys is obes s i Murano : Murano , 1983 .

An isomys is cons tr icta Murano: Murano , 1 983 .

An isomys is enewetakens is Mur ano : Mur ano , 1 983 .

Mys inae sp . : H obsen and Chess , 1 978 .

Subfam i ly GASTROSACCINAEAnchial ina gr os sa H ansen : Ger ber , 1 98 1 .

Anchial ina typ ica (Krayer ): Ger ber , 1 98 1 .

Gas tr osaccus bengalens is H ansen : Mur ano , 1 983 .

Gas tr osaccus indicus H ansen : Ger ber , 198 1Gas tr osaccu s pacif icas H ansen : Ger ber , 1981 .

Gas tr osaccus par vus H ansen : Ger ber , 1 98 1 .

Gas tr osaccus sp . : Ger ber , 1 98 1 .

Metamblyop s is sp . : Ger ber , 198 1 .

Pseudanchial ina inerm is I ll ig : Ger ber , 1 98 1 ; Murano, 1983 .

Or der AMPH IPODASuborder H YPERIIDEAFam i ly H YPERIIDAE

Les tr igonus bengalen s is Gi les : Ger ber , 198 1 as H yper ia dys schis tus Stebbing .

Les tr igonus latis s ima (Bor allius ) : Ger ber , 198 1 as H yper ia hydr ocephal ia Vosseler .

Fam i ly SYNOPIIDAESynop ia u lt ramar ina Dana: Ger ber , 1 98 1 .

Synop ia var iabi lis Spandl : Bernard, 1 965 .

Order EUPH AUSIACEAFam i ly EUPH AUSIIDAE

Pseudeuphaus ia lat if r ons (G . O. Sar s ) : Ger ber , 1 98 1 .

Order DECAPODAFam i ly LUCIFERIDAE

Lucifer chacei Bowman , 1967

Phy lum CH AETOGNATH AClass SAG ITTOIDEA

Fam i ly SAG IT TIDAEPterosagi tta dr aco (Krohn ) : Alvar i no , per sonal commun ication : Ger ber , 198 1 .

Sagi tta bedfordi i : Alvar i no , per sonal commun icati on .

Sagi tta bipunctata Quoy and Ga imard : Alvar i no , per sonal commun ication ; Ger ber 198 1 .

Sagi tta enflata Gr ass i : Johnson , 1 954 ; G i lmar tin , 1958 ; Alvar i no, per sonal commun ication ; Ger ber 1 98 1 .

Sagi tta ferox Doncaster : Alvar i no , per sonal commun ication :Sagi tta neg lecta A ida: G i lmar t in , 1 958 ; Alvar i no, per sonal commun ica tion ; Ger ber , 1981 .

Sagi tta ocean ic : Alvar i no , per sonal commun ication .

‘Sagi tta pac ifica Tok ioka: Alvar i no , per sonal commun icati on .

Sagi tta regu lar is Aida: Gi lmar tin , 1958 ; Alvar i no , per sonal commun icat ion ; Ger ber , 1 981 .

Sag i tta robus ta Doncaster : Johnson , 1 954 ; Alvar i no, per sonal commun ication .

Sagi tta ser r atoden tata Kr ohn : Johnson , 1954 ; Ger ber , 198 1 .

Phy lum CH ORDATASubphy lum UROCH ORDATAClas s TH AL IACEA

Order DOL IOL IDA (CYCLOMYARIA)Fam i ly DOLI OLI DAE

Do lio lum sp . : Kinzi e, 1 976 .

On ly recorded outs ide lagoon but in the immediate v icin i ty of Enewetak Atol l .(Th is table continued on next page. )

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2 12 GERBER

CTENOPH ORA

Ten tacu lata

Loba taOne specimen pr eser ved in the MPML r efer ence co l lee

t ion has been ident i fied to genus and was col lected in thelagoon off Enewetak Is land December 28 , 1 975 . Ctenophares wer e co l lected fr om the lagoon in Apr i l and

Decem ber 1 955 (G i lmar t in , 1 958) and in the w in ter s of

1 972 , 1 974 , and summer 1974 by Ger ber ( 198 1 ) but notident i fied to species .

MOLLUSCA

Gas t r opodaTwo spec ies of the ho loplankton ic Thecosomata, collec

t ively known as pter opods , have been r epo r ted fr omEnewetak Lagoon . Johnson ( 1 949) and G i lmar t in ( 1958)each r epor ted one spec ies fr om Enewetak and near byB ik in i , Rongelap , and Ronger ik Lagoons . Pteropods wereco l lected fr om shal low s tat ion s at Enewetak and Majur oLagoons by H obson and Chess (1 973 , 1 978) but were not

iden t i fied fur ther . Ger ber (1 98 1 ) ident i fied two specieswh ich common ly occur red in lagoon samples fr om shal lowand deep s tat ions ; they wer e especial l y common in the

summer mon ths .

On ly one un iden t i fied species of the heter opod genusA t lan ta was recor ded by G i lmar t in (1 958) fr om EnewetakLagoon wher e i t is no t ver y abundan t . Member s of th isgenus have been r epo r ted in low abundance in near byocean ic water s (Ch iba and H i r akawa, The class ictaxonom ic wor ks on thi s and the above group have beendone by Tesch ( 1 946 , 1 949 ,

ARTH ROPODA

Cr u s tacea

Os tr acodaPlankton ic os tr acods have been r ecor ded fr om

Enewetak Lagoon (Ger ber and Mar shal l , 1 974 ; H obsonand Chess , 1 978 ; Ger ber , 1 98 1 ) and Majur o Lagoon (H obson and Ches s , Those recor ded by Ger ber ( 1 981 )and mos t of those r epor ted by H obson and Chess ( 1 973 )belong to the or der Myodocopa,

but fu r ther identi ficat ionwas not made. Though they occur fai r ly r egular l y in net

co l lect ions , os tracods ar e r epresen ted by on ly a few ind iv iduals per sample .

CopepodaAppr ox imately 58 species of calano id copepods have

been iden t i fied from Enewetak Lagoon . Johnson ( 1 954)ident i fied the 1 2 mos t common spec ies from Enewetak ,Rongelap , Ronger ik , and B ik in i Lagoons but un for tunatelydid not ind icate wh ich species came from each lagoon . H is

co l lect ions at Enewetak , Ronger ik , and Rongelap Lagoons

wer e made in June and at B ik in i Lagoon fr om Mar ch toMay in 1 946 . Twelve species of calano ids were ident i fiedby G i lmar t in ( 1 958 ) at Enewetak Lagoon fr om samples collected in Apr i l , Novem ber , and Decem ber 1 955; eight ofthese spec ies were no t r ecor ded by John son Two

plankton samples wer e co l lected at Enewetak Lagoon in

September 1 959 and 1 96 1 by Mahnken (1 966) in conjunc

t ion w i th a detai led zooplankton s tudy of RongelapLagoon . H e iden t i fied 25 species fr om Enewetak Lagoonand 45 spec ies fr om Rongelap Lagoon wh ich includedmos t of the spec ies prev ious l y r epor ted fr om Enewetak . InJanuar y and Febr uar y 1 966 , Bar nett ( 1 967) s tud ied thed i s tr ibution of copepods at sever al locat ions in EnewetakLagoon . H e r ecor ded about 46 spec ies of calano id

copepods , mos t of wh ich also occur r ed in RongelapLagoon . Ger ber ( 1 981 ) sampled Enewetak La goon dur ingJanuar y and Febr uar y 1 972 and 1 974 , respect ively , and inJune thr ough Augus t 1 974 . H e ident i fied 45 species fr oma m id- lagoon s tat ion ; mos t of those species had been prev iou s ly r ecor ded fr om th is lagoon . Even after these s tudiesat Enewetak , ther e ar e species of calano id copepods wh ichhave been r ecor ded fr om Rongelap that have not beeniden t i fied fr om Enewetak Lagoon . Those species are

Euchi r ella sp . , Labidocer a bataviae Scott , Pon tella fer a,

Pan tella tenu i rem is , Pon tell ina g lobosa, Pon tellops is kram

er i,Pon tellop s is macr onyx , and U ndeuchaeta p lumosa. At

Majur o Ato l l Lagoon , two spec ies of calanoid copepods(Candacia discaudata and Undin u la vu lgar is ) wh ich alsooccur at Enewetak Lagoon wer e r ecor ded fr om the

s tomach conten ts of fish ; a th i r d species co l lected there,Labidocer a acuta, has not been r ecor ded fr om the otherlagoons (H obson and Ches s , In a s tudy of the feeding hab i ts of var ious p lankt ivor ous fishes at Enewetak ,H obson and Chess (1 978) iden t i fied three of the lar ge s i zecopepods to species and four other s to the gener ic level ;these had been r epor ted prev ious ly . Calano id copepods inreduced abundance wer e r ecor ded by Johnson (1954) fr omouts ide B ik in i Lagoon , but these species wer e not

r epor ted . Ch iba et al . ( 1 955) co l lected zooplankton in the

open ocean near B ik in i Ato l l and throughout the Mar shal lIs lands fr om May to June 1954 . They pr esented 80species of calano id copepods , about half of wh ich havebeen r ecor ded from Enewetak Lagoon ; the r emain ingspecies con s is ted of ocean ic and deep water forms . Therear e many impor tan t wor ks deal ing w i th the taxonomy andident i ficat ion of calano id copepods wh ich occur at

Enewetak Lagoon . The mos t impor tan t wor ks ar e B radfor dDak in and Colefax Delsman Fr os t

and F lem inger G iesbrecht Gr iceMor i Scott Sewell Ver voo r t

and Wi lsonCyclopo id copepods wer e fi r s t r ecor ded fr om Enewetak

and near by ato l ls by Johnson (1 954) dur ing the sampl ingper iods ment ioned above. H e r ecor ded fou r species plustwo gener a cons is ting of several un identi fied species .

G i lmar t in ( 1 958) iden t i fied four species at EnewetakLagoon , and Mahnken (1 966) iden t i fied 1 2 species fr omEnewetak and five at B ik in i and Rongelap Lagoons . These

LAGOON PLANKTON 2 13

resear cher s used rather coar se mesh nets wh ich did not

adequatel y sample th is group cons i s t ing mos tly of smal lerforms ; th is per haps accounts for the few species r ecor ded .

About 27 species wer e r ecor ded at Enewetak Lagoon byBar nett (1 967) and included those species ident i f ied therepr ev ious ly ; Ger ber ( 1 98 1 ) ident i fied 28 spec ies whichincluded mos t of Bar nett ’s spec ies and sever al addi t ionalspec ies . On ly one cyclopo id copepod , Corycaeus cur tas ,r ecorded at Rongelap has not been found at EnewetakLagoon . In the sur round ing water s of B ik in i Ato l l , 29species wer e found by Ch i ba et al . but on ly 1 3 of

these occur w i th in the lagoon at Enewetak . Four cyclopo idgenera wer e recor ded at Majur o Ato l l Lagoon fr om the

s tomach con ten ts of plankton feed ing f ishes (H obson and

Chess , all of wh ich have been r ecor ded fromEnewetak and include Oncaea sp . , Corycaeus sp . ,

Sapphir ina sp . , and Saphi rella sp . For iden t i fi cat ion and

taxonomy of cyclopo id copepods at Enewetak , the fol lowing wor ks should be consulted : Dahl Dak in and

Colefax Far r an G ies brecht Mor iMotoda Scott and W i lson

Few harpact icoid copepods are s tr ictly ho loplankton ic ,and of these, five species have been found in the

Enewetak Lagoon . They include member s of the fi r s t thr eefami l ies pr esen ted in the checkl is t under thi s or der . G i lmart in (1958) ident i fied one of these species , Bar nett ( 1 967)ident i fied thr ee spec ies , and Ger ber ( 1 981 ) found four of

these spec ies . At a shallow s tat ion (2 m depth) beh ind awindwar d r eef area of the lagoon , Ger ber ( 1 981 ) collectedin net tows 1 6 species of harpact ico ids that are bel ieved tobe ep ibenthi c types . These were res tr icted to the shal lows tation and may have been inadver ten tly washed fromadjacent cor al r eefs . H arpact ico id copepods wer e also in

the lagoon plankton and in the gu t conten ts of var iousfishes Ger ber and Mar shal l , 1974 ; H obson and

Chess , but these wer e not iden t i fied . A thor oughstudy of the ep ibenth ic harpact ico id copepods has not

been made at any locat ion in the Mar shal l Is lands . The fivepelag ic species noted above were not ident i fied fr om net

tow samp les in the neighbor ing water s of B ik in i Ato l l(Ch iba et al . , Th i s sugges ts that they ar e probablyner i tic types , sur v iv ing bes t in shal lower protected ar eas

such as atol l lagoons .

Both Bar nett (1 967) and Ger ber (1 98 1 ) ident i fiedmember s of a s ingle genus belonging to the other Monst r i llo ida. Whether the same species wer e r epr esented in boths tudies is not known , because no ident i ficat ions were madeto the species level . Mons t r i llo id copepods occur r ed on l yr arel y in net col lect ions in both s tud ies , and no r efer enceindicates that they occur outs ide of ato l l lagoons in the

Mar shal l Is lands . In formation on the ident i fi cat ion and tax

onomy of harpact ico id and monst r i llo id copepods fr omEnewetak are scar ce, but the fo l low ing wor ks ar e helpful :Dak in and Co lefax Lang Scott and

Ver voo rt

M ys idaceaA detai led inves t igat ion of the mys ids at Enewetak

Atol l has not been car r ied ou t . A plankton ic mys id was

co llected in net tows by G i lmar tin (1 958) at EnewetakLagoon , but the specimen was not ident i fied . Swarms of

mys ids belonging to two gener a (Si r iella and Mys inae) wer eobser ved and co llected over lagoon patch r eefs at

Enewetak by H obson and Chess Ger ber (1 98 1 )iden t i fied eight species of mys ids fr om net tows at a

near by shal low s tation ; and Mur ano ( 1 983 ) iden t i fied sevenspec ies fr om net tows and emer gence tr aps at depthsbetween 2 and 1 0 m . Fu r ther sampl ings , espec ial ly at

n ight , would no doubt r eveal add i t ional specimens . Un iden

t if ied mys ids were r eco r ded in the plankton and in the gu t

contents of plankton feeding fishes at near by Majur o Ato l lLagoon by H obson and Chess Plankton tows in theocean ic water s near B ik in i Ato l l by Ch iba et al . (1 955)revealed two spec ies of m ys ids , Si r iella thompson i and

Si r iella sp . , and Tatter sal l (1 951 ) descr ibed two addi tionalspec ies of Si r iella fr om the Mar shall Is lands (8 . vulgar is r os

tr ata, S . anoma la) and Anchial ina typ ica and A . pen ici llata .

As noted ear l ier , the genus Si r iella was fi r s t r ecor dedins ide the lagoon at Enewetak by H obson and Chess

and later two other lagoon Si r iella were iden t i fiedto spec ies by Mu r ano Several excel len t wor ks byH ansen (1 91 0) and Tatter sal l (1 957) have pr oved useful inident i fy ing the mys ids from Enewetak .

Am ph ipoda

At leas t thr ee and poss ibly four spec ies of holop lank

ton ic amph ipods that belong to the subor der H yper i ideahave been r epor ted from the Enewetak Lagoon (Ger ber ,

Ear l ier plankton co l lect ions by G i lmar t in (1 958)r evealed one un iden t i fied spec ies , and H obson and Chess( 1978) r ecor ded but did not ident i fy several spec ies of

hyper i ids . Man y more nonp lankton ic species occur w i th inthe lagoon (Bar nar d , and these are mos t ly gammar ids wh ich occu r in the epiben th ic shal low water hab i tatd i scussed elsewher e in th is book . Bar nar d (1 965) r epor tedone gammar id spec ies , SynOp ia var iabi l is , wh ich was col

lected fr om Enewetak and I faluk Lagoons by bo th planktonhaul and dip-nett ing w i th a n ight l ight ; i t is included in thecheckl is t . H yper i id amph ipods have been recor ded at

Maju r o Ato l l (H obson and Ches s , 1 973) where one

species , Les t r igonus bengalens is , was iden t i fied from net

tows and made up an im po rtan t d ietar y componen t ofanther in id fi sh . In the ocean ic water s ar ound the Mar shal lIs lands , Ch iba et al . ( 1 955) co l lected n ine species of

plankton ic amph ipods , none of wh ich appear to occur inEnewetak Lagoon . These are Oxycepha lus por cel lus ,Ph r on ima pacif ica, Ph r on ima sp . , Lep tocot is tenu i r os t r is ,

Isopoda

The i sopods wh ich have been co l lected in plankton net

hauls from Enewetak Lagoon have not been ident i fied togenera G i lmar t in , 1 958 ; H obson and Chess , 1 978 ;and Ger ber , Th is gr oup is gener al ly scar ce in suchsampl ings ind icat ing that they are mos t l ikely of epibenthicor ig in rather than plankton ic . Plankton ic isopods fr omother atol l lagoons or in the ocean ic water s in the Mar shal lIs lands have not been col lected .

2 14 GERBER

Par ascelus edwards i , Phr on imop s is sp in ifer a,H yper io ides

s ibag in is , H yper ia sp . , and Anchylomer a blossevi ll i i . For the

taxonomy and iden t i fi cat ion of the amph ipods fr omEnewetak , see Bar nar d Bowman and Gr uner

and Dak in and Colefax

Euphaus iacea

A s ingle species of euphaus i id has been r epor ted fr omEnewetak Lagoon where i t occur s regular l y at bo th shal lowand deep s tat ions (Ger ber , Th is species was alsothe dom inan t euphaus i id r ecor ded in s ide the Gr eat Bar r ierReef Lagoon (Rus sel l , G i lmar t in (1 958) repor tedeuphaus i ids in hi s plankton hauls at Enewetak Lagoon butdid not iden t i fy them . In the sur round ing ocean ic water s of

Enewetak and B ik in i Ato l ls , seven species of typ i cal lyocean ic euphaus i ids wer e r epor ted by Ch iba et al .

and none of these occur s in the ato l l lagoons . They areStylochei r on car inatum , S. s ubm i i , Euphaus ia graci l is ,E . g ibba,

E . kr ohn i i , Thysanopoda obtus if r ons , and

T . tr icusp idata . U seful wo r ks concer n ing the iden t i ficat ionand taxonomy of euphaus i ids include Dak in and Colefax

Delsman H ansen Mauchl ine and

F isher and Mauchl ine

CH AETOGNATH AAt leas t n ine species of chaetognaths have been

repor ted and ident i fied fr om Enewetak Lagoon , in add i tionto seven species fr om outs ide the lagoon or in the near byocean water s . They are an impor tant plankton ic gr oup andappear in mos t net sampl ings . Johnson (1 949 , 1 954)repor ted thr ee species fr om net tows taken dur ing 1 946 at

Enewetak Lagoon (June), and the lagoons at Rongelap

(June), Ronger ik (June), and B ik in i (Mar ch to May) and

from outs ide B ik in i Atol l (Mar ch to May) . D is tr ibution of

chaetognaths w i th in B ik in i Ato l l was also s tud ied . A

t r ans lagoon s tudy of plankton at Enewetak fr om Apr i lthr ough December 1 955 by G i lmar t in ( 1958) r evea led fourspec ies , mos t of wh ich were abundan t in the net samples .

Ger ber ( 1 98 1 ) r epor ted at leas t s ix species from a m id

lagoon s tat ion in the w inter and summer (1 972, 1974)w i th incr eased abundances in summer . Co l lect ions made in1 946 and later r evealed s ix species fr om the lagoon and

sever al mor e fr om outs ide, eas t of the lagoon (A lvar ino,

per sonal commun i cat ion) . In the neighbo r ing water s ar oundB ik in i Ato l l , extens ive plankton co l lect ions were made in

May and June 1954 by Ch iba et al . (1 955) and later byTsur uta These r esear cher s iden t i fied at leas t 1 6species ; the eight not found wi th in the lagoons includeSag i tta decip iens , S . hexap ter a, S . lyra, S . m in ima, S. p u l

ch r e, S . pseudoser r atoden tata, Kr ohn i tta pacifica, and

K . subt i l is . T hr ee species of chaetognaths were repor tedfr om the gu t con ten ts of lagoon fish at Majuro Ato ll (H obson and Chess , 1 973 ) and at Enewetak Ato l l (Ger ber andMar shal l , Impor tan t sour ces for the iden t i fi cation of

chaetognaths include A lvar ino , Sund and

Thomson

CH ORDATA

Thal iacea

Th i s gr oup i s much les s abundan t than the Lar vacea inthe Enewetak Lagoon and from near by ato l ls . They werefi r s t r epor ted fr om Enewetak by G i lmar t in ( 1 958) whoident i fied two species from Apr i l thr ough December in hist rans lagoon plankton s tudy of 1955 . An addi t ional spec ieshas been co l lected from the lagoon in 12 m of water and

iden t i fied by R . A . K in z ie (MPML reference col lect ion) .

CONCLUSIONSThere appear s to be l i tt le di fference of the holoplank

ton ic zooplankton spec ies compos i t ion between Enewetakand near by ato l l lagoons such as Rongelap . Th is observat ion i s largel y based on the copepod data wh ich have beenmos t extens ivel y s tudied at these two lagoons . Clear ly , thi sfaunal s im i lar i ty reflects the r ather s im i lar habi tats of the

two lagoons and thei r s im i lar locat ions wi th r espect to thepr evai l ing ocean cur r en ts .

The in format ion avai lable compar ing the copepodspec ies assemblege ins ide the lagoon w i th that of the sur

r ound ing ocean outs ide the lagoon demons tr ates that theyar e d i fferent . The lagoon zooplankton populations are par

t ially composed of ocean ic species known to occur in the

neighbor ing water s outs ide the ato l l , in addi t ion to ner i tics pec ies wh ich occur on ly in shal low pr otected env i ron

ments . It has also been found that the dens i ty of zooplank

Appendi cu lar ia (Lar vacea )

Th i s is an abundan t zooplankton gr oup in the ato lllagoons and sur r ound ing water s of the Mar shal l Is lands . At

leas t seven species have been repor ted fr om EnewetakLagoon , five of wh ich have been co llected fr om outs ide theatol l . Johnson (1 954) r epor ted dens i t ies of appendicu lar i

ans wh ich wer e cons iderably h igher for Enewetak and

near by ato ll lagoon s than in the near by open ocean ; he didnot ident i fy genera or species . At Enewetak Lagoon , G i l

mar t in ( 1958) also repor ted h igh dens i t ies of th i s gr oupespec iall y dur ing Apr i l and in ear ly November and ident i

f ied them as Oikop leu r a sp . Ger ber s im i lar l y (1981 ) foun dh igh dens i ties of lar vaceans in the lagoon especiall y in thesummer ; at leas t s ix species ar e r ecor ded belong ing to twofam i l ies . In the water s near B ik in i Atol l , Ch i ba et al .

(1 955) iden t i f ied 1 6 spec ies , five of wh ich are common toEnewetak Lagoon . These add i t ional spec ies includeFr i t i l lar ia bo real is forma sargass i , F . form ica, F . ha

_p

_

Ias toma, F . pellucida, F . venus ta, O ikop leu ra cophocer ca,

O ikop leu r a sp . , Kowa levskaia tenu is , Megalocercus hux leyi ,Stegosoma magnum, and Pelagop leu r a sp . Appendiculari ahave been repor ted at Majur o Atoll La goon fr om the gu t

contents of plankton feeding fishes (H obson and Chess ,the species wer e not iden ti fied . The ident ificat ion

and taxonomy of the Thal iacea and La r vacea are includedin the wor ks of Bjo r nberg and For ner is Delsman

and F raser

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2 16 GERBER

Johannes , R . E . , and R . P . Ger ber , 1974 , Impor t and Expor t ofNet P lankton by an Enewetak Coral Reef Commun i ty , in

Pr oceedings of the Second In ter nat ional Coral Reef Sympas ium , Austral ia, 1973 : pp . 97

_

- 104 .

S. L . Co les , and N . T . Kuenzel , 1970 , The Ro le of Zoo

plankton in the Nutr it ion of Some Scleract in ian Co rals ,L im nol . Oceanogr .

, 15: 579-586 .

Johnson , M . W . , 1 949, Zooplankton as an Index of WaterExchange Between B ik in i Lagoon and the Open Sea, Trans .

Am . Geophys . Un ion , 30 : 238-244 .

1 954 , P lankton of the Nor ther n Mar shal l Is lands , B ik in i andNear by Ato l ls , U . S. Geo l . Sur v. Profes s ional Paper 260-F ,

30 1 -3 1 4 .

Kr amer , A . , 1 897, Ueber den Bau der Koral lenr iffe und die

Planktonver theibeng an den Samoan ischen Kus ten nebs t ver

g leichenden Bemerkungen , K iel and Leipzig .

Kr amp , P . L . , 1 96 1 , Synops is of the Medusae of the Wor ld,J . Mar . B io l . Assoc . U . K . , 40 : 7-469 .

Kofo id, C . A . , and A . S. Campbe l l , 1939, Repor ts on the Scien

t i f ic Resu lts of the Expedi tion to the Eastern T r opica l Pacific ,in Charge of A . Agass i z , by the U . S. Fish . Comm . SteamerA lbat r os s fr om Oct . 1 904— Mar . 1 905, The Ci l iata:Tin ti n no inea, Bu l l . Mus . Comp . Zoo l H ar var d Un iver s ity, 84

1 4 73 .

Lang , K . , 1948 , Monogr aph ie der H arpact iciden , H . Ohlssons ,

Lund, Sweden .

Lebour , M . , 1 925 , D inoflagellates of the Nor ther n Seas , Mar .

B io l . As soc . U . K . , P lymouth .Mahnken , C . , 1 966 , D is tr ibut ions of Copepoda at Rongelap A to l l,Mar shall Is lands , M . S. thes is , Un iver s i ty of Wash ington .

Mauchl ine, J. , 1 980 , The B io logy of Mys ids , Euphaus i ids , Adv .

Mar . B io l . , 1 8: 1 -68 1 .

and L . R . Fisher , 1 969, The B io logy of Euphau snds , Adv .

Mar . B io l . , 7: 1 -454 .

Mayer , A . G . , 1910 , Medusae of the Wor ld, H ydromedusae and

Scyphomedusae, Car negie In st . Wash . , Publ . 1 09, 3 vo ls .

M ichel , A . , 1969 , P lancton des Lagoon et des Abo rds Exter ieur sde L

’

Atoll de Murur oa , Cah . Pac . , 1 3: 8 1 -1 32 .

C . Co l in , R . Desr os ieres , and C. Oudot , 1971 , Observationssu r L

’

H ydrolog ie et le P lancton des Abords et de la Zone desPasses de L ’

A toll de Rangi roa (Arch ipel des Tuamotu, OceanPacifique Centr al), Cah . ORSTOM , Ser . Oceanogr . , 9:

375-402 .

Mor i , T . , 1 937 , The Pe lagic Copepoda f rom the Neighbo r ingWater s of Japan , Soyo Co . lnc . , Tokyo , 1 964 .

Motoda, S. ; 1938 , Quant itat ive Studies on the Macroplankton off

Coral Reefs of Palao Por t , Tr ans . Sappo ro Nat . Soc . , 1 5:

242-246 .

1 963 , Corycaeu s and F ar r an u la (Copepoda, Cyclopo ida) in

H awai ian Water s , Seto Mar . B io l . Lab . , 1 1 : 209-262 .

Murano , M . , 1 983 , Mys idacea Fauna fr om Enewetak Lagoon ,

M icrones ia, Bu l l . P lankton Soc . Jpn . , 30 : 81 -90 .

Odum , H . T . , and E . P . Odum , 1955 , T rophic Structur e and Pr o

duct iv i ty of a W indward Co r al Reef Commun i ty on EnewetakAto l l , Eco l . Monogr . , 25: 291 -320 .

Par ke, M . , and P . S. Dixon , 1 976 , Check l i st of Br i ti sh Mar ineAlgae

— Th i rd Rev is ion , J . Mar . B io l . Assoc . U . K . , 56 :

527-594 .

Russel l , F . S. , 1934 , The Zoop lankton , III , A Compar i son of the

Abundance of Zooplankton in the Great Bar r ier Reef Lagoonwith that of Some Reg ions in Nor thern European Water s ,Gr eat Bar r ier Reef Expedit ion , 1 928 — 1 929, Sci . Rept . , 2

159-20 1 .

Sch i l ler , J. , 1937 , Dr . L . Rabenhor s t’

s Kryptogamen-Flora von

Deutschland , Os ter r eich und der Schwei z , Flagellate,Di noflagettae, Zehnter Band, Dr i tte Abtei lung .

Scott , A ., 1 909 , The Copepoda of the Siboga Expedition ,

Part l . ,F reeswimm ing , L i ttor al , and Sem iparas i t ic Copepoda, Siboga

Exp . Monogr . , 29: 5-320 .

Sear s , M . , 1 950 , Notes on Siphonophores ,I , Siphonophores from

the Mar shal l Is lands , J . Mar . Res . , 9: 1 - 1 6 .

Sewel l , R . B . , 1929, The Copepoda of Indian Seas , Calanoidae,

Mem . Indian Mus . , 10: 1 -221 .

Sund , P . N . , 1 959 , A Key to the Chaetognatha of the TropicalEas tern Paci fic Ocean , Pac . Sci . , 13: 269-285 .

Tatter sa l l , O . S. , 1957 , Mys idacea , D iscover y Rep . , 28 : 3- 1 89 .

Tatter sal l , W . M . , 1 951 , A Rev iew of the Mys idacea of the

Un i ted States National Museum , Bu l l . U . S. Nat l . M us ., 201 :

1 -29 1 .

Tesch , J . J. , 1 946 , The Thecosomatous Pterpods , I , The At lant ic , Dana Rep . , 28 : 1 -82 .

1 949 , H eter opoda, Dana Rep ., 34: 1 -54 .

1950 , The Gymnosomata, II , Dana Rep . , 36: 1 -55.

Thom son , J . M . , 1 947 , The Chaetognatha of South-Eas tem Aus

t ral ia, Commonwealth of Aus tralia, Counci l for Scien tif ic andIndus t r ial Resear ch (D iv . of Fis her ies — Rep . No . Bul l . No .

222 .

Totton , A . K . , and H . E. Bargmann , 1965 ,A Synop s is of the

Siphonophor a, B r . Mu s . (Nat . London .

T r anter , D . J. , and J . George, 1 972 , Zooplankton Abundance at

Kavaratti and Kalpen i Ato lls in the Laccadives , Symp . Cor alsand Coral Reefs , Pr oc . 1 969 (Mar . B io l . As soc . India) : pp .239-256 .

Tsuruta, A . , 1 963 , D istr ibution of P lankton and Its Char acter i s

t ics in the Ocean ic F ish ing Ground , wi th Special Reference toThei r Relation to Fi sh ing , J . Sh imonoseki Co ll . Fish , 1 2:

13-2 1 4 .

Ver voo r t , W . , 1 964 , Fr ee-L iv ing Copepoda fr om Ifaluk Ato l l inthe Car o l ine Is lands . U . S . Nat . M us . Bu l l . , 236 : 1 -43 1 .

W i lson , C . B . , 1 942 , The Copepods of the P lankton GatheredDur ing the Las t Cru ise of the Car negie, B io logy-I, Car negieIns t . Wash . Pub . , 536 : 1 -237 .

Chapter 2 1

Os tracoda (M yodocop i na) of Enewetak A to ll

LOUIS S. KORN ICKER

Depar tmen t of Inver tebrate Zoo logy, Nationa l M useumof Natur al H is tor y, Sm ithson ian Ins titution ,

Wash ington , D . C . 20560

INTRODUCTIONOs tr acods in the superor der Myodocopa, excluding theplankton ic subor der H alocyp r idina, wh ich is outs ide the

scope of the pr esent r epor t , have not pr ev ious ly beenrepor ted from Micr ones ia. A compr ehen s ive checkl is t ofOs tracoda from southeas t As ia (H anai et al . , 1980) is con

cerned w i th the area between 25°N and 1 1 °S lat i tude and

14O°E and SS

°E longi tude. The ar ea includes the wes tern

edge of M icrones ia, but a br ief per usal of the check lis tr evealed no l is t ing of specimens fr om M icr ones ia.

A small co llect ion of Enewetak os tr acods in the

subor der Myodocop ina at the National Museum of NaturalH istor y , Sm i thson ian Ins t i tut ion , contains member s of all

five fam i l ies that compr ise the subor der (TableAlthough the species in the co l lect ion are capable of swimming, and do so occas ional ly , they spend mos t of thei rexis tence cr awling on , or bur r ow ing in , the subs trate. An

except ion to th is i s member s of the main ly pelag ic genusCyp r idina, wh ich is represen ted in the co l lect ion by twospecies . Some of the Enewetak specimens are i l lus trated inF ig . 1 .

The feeding habi ts of the fami l ies di ffer : the Cylindraleber ididae i s composed of fi l ter feeders , the Sar s iell idaeand Rut ider rnat idae of car n ivores , the Phi lomedidae of

detr i tus feeder s , and the Cypr id in idae of s cavenger s ,detr i tu s feeder s , and also car n ivores (Korn icker ,

B IOGEOGRAPH IC COMPARISONSAT GENERIC LEVELThe genera Skogsbergia, H arbans us , Ru t iderma,

Sar s iella, Paras terope, and Cylindroleber is ar e wor ldw ide indis tr ibut ion ; on ly Par as terope is presen t in Antar ct ic water s(Korn icker , and none has been r epor ted fr om theAr ctic Ocean .

TABLE 1

Checkl i s t of Enewetak A tol l Os t r acoda(Myodocop i na)

New Micrones ian r ecord .1‘ New Enewetak r ecord .

The pelag ic genus Cyp r idina is pr imar i l y lndo Paci fic .

Ancohen ia is pr ev ious ly known fr om Maunalua Bay,Oahu Is land , H awai i (Kor n icker , 1 976z7) and Vuna Po in t ,Taveun i , Fi j i (Kor n icker ,

Anscott iella has pr ev ious ly been co l lected off Ceylonand Thai land (Poulsen , 1965z65; Kor n icker , 1975z607 ;Kor n icker and McKenz ie,

Par avargu la has pr ev ious ly been col lected off the t ip of

South Afr ica and from the Eas t- Ind ian re'

g ion (Poulsen ,

Kor n icker , 1 975z3) .

Phy lum ARTH ROPODASubphy lum CRUSTACEAClass OSTRACODASuperorder MYODOCOPAOrder MYODOCOPIDASuborder MYODOCOPINASuper fam i ly CYPRIDINOIDEAFam i ly CYPR IDIN IDAE"(Cypr idina spp .

‘TParavargu la sp .

‘TSkogsbergia sp .

Fam i ly PH ILOMED IDAE‘TH ar bans us sp .

Fam i ly RUT IDERMATIDAE‘TRu t iderma sp .

Fam i ly SARSIELL IDAE‘TAncohen ia sp .

‘TAnscott iella sp .

‘TSar s iella sp .

Fam i ly CYLINDROLEBERID IDAE‘TParas terope sp .

‘TCyl indr oleber is sp .

KORNICKER

EF ig . l a, H ar bansua species . female. length mm . USNM 158313 ; b. R ut i derma specia . female. length USNM

158309; c, Ansoot riel la speci es . male, length mm . USNM 158313 ; ( I , Anaoott iel la species. female, length USNM

15831 7; e, Sar s iel la speci es , female. length mm . USNM 158310: f . Ancohen ia species. male. length mm . USNM 1583 12.

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Chapter 22

Crus tacea D ecapoda (Penaeidea , Stenopod idea ,

Caridea, and Pa l i n u r a) of Enewetak A to ll

DENN IS M . DEVAN EY‘(deceased) and A . J . BRUCET

‘Ber n ice P . B is hop Museum

, H ono lu lu , H awa i i 968 1 7;“

( Nor ther n Ter r itor y Museum of A r ts and Sc iences ,GPO Box 4646 , Darwin , N . T . 5794 , Aus tr a l ia

INTRODUCTIONB iolog ical sur veys made dur ing 1 946 and 1947 in the

nor ther n Mar shall Is lands included faunal co l lect ions thatr esulted in a publ icat ion by Chace ( 1 955) on severalgr oups of shr imps . In th is wor k , s ix species of ser ges t id,

pas iphaeid, p rocess id, and alpheid shr imps were r ecor dedfr om Enewetak . Since then the number of taxa fr om the

atol l has increased unt i l now near l y 150 decapod shr impand lobs ter spec ies ar e r ecogn i zed . Pon ton i ines and

alpheids accoun t for 7396 ( 1 06 spec ies ) of the total

number . There ar e sever al new r ecords not on ly forEnewetak but also for the Mar shal ls , and in some cases ,for the Paci fic . A taxonomic checkl is t (Table 1 ) and a sec

t ion on co l lect ion data for new r ecor ds are pr ov ided .

TABLE 1

Check l i s t of Enewetak Penaeidea , Stenopod idea , Car i dea, and Pal i nu r a‘

Subphy lum CRUSTACEAClas s MALACOSTRACAOrder DECAPODASuborder DENDROBRANCH IATAInfr aorder PENAEIDEA

Super fam i ly PENAEOIDEAFam i ly PENAEIDAE

“

l'Metapenaeop s is bo r radai lei (De Man ,

TMetapena p s is sp .

Fam i ly SICYONIIDAE‘l’

Sicyonella maldivens is Bo r r adai le, 1 910 .

Super fam i ly SERGESTOIDEAFam i ly SERGESTIDAE

Lucifer chacei Bowman , 1967 : Bowman , 1 967 .

Lucifer sp . : Chace, 1 955 .

‘The higher class i ficat ion used here fo l lows that of Bowman and AbeleTNew records for Enewetak .

(Th i s table cont inued on next page.)

PEN AEIDEA

Two penaeid species in the genus Metapenaeop s is , one

a w idel y d is tr ibuted Indo-Wes t Paci fic form and the otheras yet un iden t i fied , ar e now r ecor ded for the fi r s t t imefrom Enewetak . Among the s icyon id shr imps , the w idelyd i s tr ibuted Indo-Wes t Pac i fic species , Sicyonella mal

divens is , has been co l lected at Enewetak on the sur faceus ing a l ight at n ight . One plankton ic ser ges t id, an undeter

m ined species of L ucifer , was repor ted by Chace ( 1 955)and was co l lected at Enewetak in the same manner . In

1967 , Bowman r ecor ded L . chacei as a new species fr oma num ber of Indo-Wes t Paci fic locat ions includ ing Enewetakand Rongelap Ato l ls in the Mar shal ls .

222 DEVANEY AND BRUCE

TABLE 1 (con t’d)Subo rder PLEOCYEMATAInfr aorder STENOPODIDEA

Fam i ly STENOPODIDAE‘

l'M icr op r os thema p lum icom e (R ichter s ,1

’

M icrop ros thema scabr icaudatum (R ichter s ,1

’

Odon tozona scu lp t icaudata H olthu is , 1 947 .

‘

l’

Odon tozona sp . nov .

‘l’ Stenopus hisp idus (Ol iv ier ,l Stenopus tenu ir os tr is De Man , 1 888 .

TStenopu s zanz ibar icus B ruce, 1976 .

Infraorder CAR IDEASuper fam i ly PASIPH AEOIDEAFam i ly PASIPH AEIDAE

Lep tochela i r robu s ta Chace, 1 976 : Chace, 1976 .

Lep tochela robus ta Stimpson , 1 860 : Chace, 1955 .

Superfam i ly RH YNCH OCINETOIDEAFam i ly RH YNCH OCINETIDAE

Rhynchocinetes hiatt i H olthu i s and H ayash i , 1 967 : H olthu i s and H ayash i , 1 967 .

Rhynchoc inetes hender son i Kemp, 1 925 : H olthu is and H ayash i , 1967 .

Rhynchoc inetes mar shallens is Edmondson , 1 952 : Edmondson , 1952 .

‘

l‘

Rhynchoc inetes aff . R . r igens Gordon , 1936 .

‘

t ynchoc inetes sp .

Super fam i ly PALAEMONOIDEAFam i ly PALAEMON IDAESubfam i ly PALAEMON INAE

‘l‘

B rachycarpus biungu icu latu s (Lucas ,‘

l Leandr i tes cyr tor hynchu s Fuj i no and M iyake, 1969 .

Subfam i ly PONTON IINAEl Anchis tus aus tr a lis B ruce, 1 977 .

Anchis tus m ier s i (De Man , Fankboner , 1972 .

1'

Ar aiopon ton ia odon tor hyncha Fuj i no and M iyake, 1970 .

l'

Conchodytes biungu icu latus (Paulson ,

‘l'

Conchodytes me leagr inae Peter s , 1 852 .

“

l'

Conchodytes tr idacnae Peter s , 1 852 .

l Corall iocar is brevi ros t r is Bor radai le, 1 898 .

l Corall iocar is gr am inea (Dana,

l Corall iocar is nudi r os t r is (H el ler ,TCorall iocar is super ba (Dana,

TCor all iocar is venus ta Kemp, 1 922 .

Ctenopon ton ia cyphas t reophi la Bruce, 1979: B ruce, 1979b .

‘

l'F enner a chacei H o lthu is , 1 951 .

TH arp i l iops is beaup r es i i (Audou in ,

‘

l’

H arp i l iop s is depressa (Stimpson ,

1’

H arp i l iop s is sp in igera (Or tmann ,

1‘Jocas te japon ica (Or tmann ,

“

l’

Jocas te lucina (Nobi l i ,‘

l‘Onycocar is quadr atophthalma (Balss ,Palaemonella po tts i (Bo r radai le, Bruce, 1 970 .

Palaemonella r o tumana (Bo r radai le, Bruce, 1 970 .

TPalaemoneIla tenuipes Dana, 1 852 .

TPalaemonella sp .

Par anch is tus armatus (H . M i lne Edwards , Bruce, 1 975 ; l 979a .

Paranchis tus biungu icu latus (Bo r r adai le, Rosewater , 1965 .

Par atyp ton s ieben r ocki Bals s , 1 914 : B ruce, 1969 .

“

l'Per icl imenaeu s sp .

l Per iclimenes agag Kemp , 1 922 .

T¢Per icl imenes (De Man ,

“

l’

Per icl imenes bayer i H olthu is , 1981 .

Per icl imenes br evicarpal is (Schenkel , B ruce, 1979a.

TNew r ecords for Enewetak .t The status of P . ambo inen s is (De Man ) is not clear . It may be synonymous wi th P . com utas Bor r adai le.

(Th i s table cont inued next page. )

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224 DEVANEY AND BRUCE

TABLE 1 (con t ’d)

Fam i ly ALPH EIDAE (cont ’d)A lpheus s trenuus s trenuus Dana, 1 852.

A lpheus s t renu us Dana, 1852 : Banner and Banner , 1968 .

A lpheus su lcatus Kings ley , 1 878 .

A lpheus macrochi r us R ichter s , 1 880 : Banner and Banner , 1968 .

A thanas areteform is Cou t ier e, 1 903 : Banner and Banner , 1968 .

A thanas djibou tens is Cou t ier e, 1 897 : Chace, 1955 ; Banner and Banner , 1960 , 1968 .

Athanas dor salis (Stimpson , Banner and Banner , 1960 , 1 968 .

A thanas esaki i Kubo , 1940 : Banner and Banner , 1 968 .

A thanas indicus (Cou t ié re, Banner and Banner , 1960 , 1 968 .

A thanas mar shallens is Chace, 1 955: Chace, 1 955 ; Banner and Banner , 1960 , 1 968 .

Arbanas r hothionas tes Banner and Banner , 1960 : Banner and Banner , 1 960 , 1968 .

A thanas ver r ucosus Banner and Banner , 1960 : Banner and Banner , 1 960 , 1968 .

Automate dolichognatha De Man , 1 888 .

Automate gardiner i Cou t ier e, 1902 : Banner and Banner , 1 968 .

Automate johnson i Chace, 1 955: Chace, 1 955 .

Metalpheus par agr aci l is (Cou t iere,

A lpheus par agr aci l is Cou t iere, 1 897 : Banner and Banner , 1968 .

Metalpheus r os tr at ipes (Pocock ,A lpheus ros t rat ipes Pocock , 1 890 : Banner and Banner , 1968 .

Salmoneus ser r at idig i tus (Cou t iere,

Salmoneus s ibogae (De Man , Banner and Banner , 1 968 .

Salmoneus tr icr is tatus Banner , 1 959 : Banner and Banner , 1968 .

Synalpheus car inatus (De Man , Banner and Banner , 1968 .

Synalpheus choron (H el ler , Banner and Banner , 1968 .

Synalpheus near cou t ierei Banner , 1953 : H ighsm i th , 198 1 .

Synalpheus deman i Bor r adai le, 1900 : Banner and Banner , 1968 .

Synalpheus has t i l icrassu s Goutiere. 1 905 : Banner and Banner , 1 968 .

Synalpheus beran i Goutier e,

Synalpheus paraneomer is Cou t iere, 1 905 : Banner and Banner , 1968 .

Synalpheus s timpson i i (De Man , Banner and Banner , 1968 .

Synalpheus tum idomanus (Paulson , Banner and Banner , 1 968 ; H ighsm ith , 1 98 1 .

Fam i ly H IPPOLYT IDAE"l'H ippo lyte H . M i lne-Edwards , 1 837 .

l H ippo lyte sp .

'

I'

L igu r uveae (Bor radai le,

‘

l'Lysmata vi ttata (Stimpson ,

l Lysmata sp .

‘l Saron marmoratu s (Ol iv ier ,l Saron neg lectus De Man , 1 902 .

‘l'Thor ambo inens is De Man , 1 888 .

“

I'Thor maldivens is Bor radai le, 1 915 .

TThor paschal is (H el ler ,Fam i ly PROCESSIDAE

N iko ides s tein i i (Edmondson ,

N iko ides nanus Chace, 1 955 : Chace, 1955 .

‘l'N iko ides m u lt isp inatus H ayash i , 198 1 .

‘l'P rocessa (De H aan ,

A

'

l‘Pr oces sa mo lar is Chace, 1955 .

‘

TPr oces sa neg lecta H ayashi , 1 975 .

Super fam i ly PANDALOIDEAFam i ly PANDAL IDAE

M icr Opandalus hardingi Bruce, 1 983 : Bruce, 1 983 .

Fam i ly TH ALASSOCARID IDAE‘l

’

Thalassocar is cr inata (Dana,

Super fam i ly CRANGONOIDEAFam i ly CRANGON IDAEl Pon tophi lus sp . aff . P . sabsechota Kemp, 1 91 1 .

‘

l'Ver co ia gibbosa Baker , 1904 .

“

( New r ecords for Enewetak . (Th is table continued on next page. )

CRUSTACEA DECAPODA 225

TABLE 1 (con t'd)Infr aorder PAL INURA

Super fam i ly PAL INUROIDEAFam i ly SYNAXIDAE

‘l

’

Pal inur ellus wienecki i (De Man ,

Fam i ly PAL INUR IDAE“

I'Pan u l i r us longipes fem or is tr iga (v

'

on Mar tens ,Panu l i r us penc illatus (Ol iv ier , Ford et al . , 1 979 ; CookeMacDonald, 198 1 ; Eber t and Ford, 1986 .

Fam i ly SCYLLAR IDAEl Par r ibacus an tar ct icas (Lund,

TNew r ecords for Enewetak .

STENOPOD IDEA

Ident i fied s tenopodid shr imps determined fromEnewetak include the pantrop ical banded coral shr imp ,Stenopus h isp idus ; S . tenu i r os t r is , from the wes ter n Ind ianOcean ,

Ph i l ipp ines , and the wes ter n Paci fic ; 8 . zan z ibaricus , fr om the wes ter n Indian Ocean and centr al Pac i fic(Canton Is land) ; Odon tozona scu lp t icaudata, known fromMalayan and nor theas ter n Austr al ian water s ; and two

spec ies of M icr op r os thema . One of these, M . scabr icauda

tum , has been r eco r ded fr om the wes ter n Indian Ocean ,

New Gu inea, and the Ryukyu ’s , wher eas M . p lum icom e

has prev ious l y been recorded on l y from Maur i t ius in the

wester n Ind ian Ocean . All of these s tenopod id shr imps are

appar en tly new r ecor ds fr om the Mar shal l Is lands .

J . W . Goy (per sonal commun icat ion) has also iden t i fied8 . h isp idus fr om Jalu i t Atol l (USNM spec imens ) . Recen tly ,an undescr ibed species of Odon tozona was found and

awai ts descr ipt ion .

CARIDEA

One w idely dis tr ibuted Indo-Wes t Paci fic pas iphaeidshr imp , a spec ies of Lep tochela, is known fr om Enewetakand several other Mar shal l Is lands (Chace, 1 955 ; Itwas said to have been the mos t common decapod founddur ing the 1 946 and 1 947 sur vey and was co l lected at

n ight on the sur face us ing a l ight .Four spec ies of r hynchocinet id shr imps ar e r ecor ded :

Rhynchoc inetes h iatt i occu r s at Enewetak as wel l as severalother Pac i fic is land ar eas ; R . hender son i , r epor ted as a

new species (R . mar shallens is) from Enewetak by Edmondson is a w ide-

spread Indo-Wes t Paci fic form (H olthu is and H ayash i , a species appr oach ing R . r igens

is newly r ecor ded from the ato l l ; a four th species i s as yet

undeterm ined and awai ts descr ipt ion (H olthu is , per sonalcommun icat ion) .Palaemon id shr imps known fr om Enewetak are

r epr esented by 43 ident i fied and 1 0 un ident i fied species in1 8 genera. All but two species are member s of the sub

fa-mi ly Pon ton i inae. Rosewater (1 965) repor ted a species ofParanchis tus and Fankboner ( 1 972) a species of Anchis tus

as symbion ts fr om t r idacn id clam s . Anchis tus aus tr a lis and

thr ee spec ies of Conchodytes can now be added to the lis tof s ym bion ts assoc iated wi th t r idacn id and os t reid bivalvesat Enewetak . Par atyp ton s ieben rocki , a spec ies forminggal ls in Acr opor a cor al , was r ecor ded fr om Enewetak byB r uce Two w idely d is tr ibuted Indo-Wes t Paci ficspec ies of Palaemonella (P . potts i and P . rotumana) werer epor ted fr om Enewetak by Br uce the former a

commensal on the cr inoid Coman thu s bennetti but mor er ecently found on two add i t ional cr ino id species (TablePer icl imenes soror , a symbion t on man y tr opi cal as ter o ids ,was fi r s t recor ded from Enewetak by Br uceLaborator y sponsored projects in 1 974 led to the co l leet ion and descr iption of eight ponton i ine shr imps , one beingnew and five being w idel y d is tr ibuted tr opi cal species prev iou s ly un r ecor ded fr om the Mar shal l Is lands (Br uce,

l 979a) . Fou r of these species wer e assoc iated w i th sea

anemones ; one each was found on a sea s tar , sponge, and

t r idacn id clam ; and one is appar ent ly a fr ee-l iv ing form . A

r eef coral wor kshop held at the M id-Paci fic Resear chLaborator y (MPRL ) in 1 976 led to the collect ion and

descr ipt ion of a new genus and species of ponton i ine

shr im p associated w i th a s tony coral (Br uce, 1979h) . Studies on the commensals and par as i tes assoc iated w i th comatul id cr ino ids (feather -s tar s ) by Ann Field ing and Denn is

Devaney in 1 976 r esulted in find ing seven ponton i ine and

three alpheid shr imps as well as a galatheid and a br achyur an crab (Table One of the pon ton i ines , Per icl imenes

p i l ipes , is a spec ies descr ibed as new from Enewetak(B r uce and Zmar zly, A s tudy detai l ing the biolog icalr elat ionsh ips between Enewetak cr ino ids , thei r symbiot iccr us tacean s , and other inver tebr ates has r ecent ly been published (Zmar z ly , Two species of H arp i l iops is and

one species of Fenner a ar e s ymbionts on poci llipor id

cor als . Sim i lar ly , the five species of Corall iocar is and two

species of Ph i lar ius , herein r ecor ded fr om Enewetak for thefi r s t t ime , ar e also symbion ts of Acropo ra corals in otherpar ts of the Indo Wes t Paci fic (B r uce, Th i r ty of the48 ident i fied ponton i ine shr imps r ecor ded fr om Enewetakar e also known fr om H eron Is land , Aus tr al ia (B r uce,

The r emainder are, w i th few except ions , w idespr ead trop ical Indo-Wes t Pac i fic or pan tr op ica l spec ies . All

but thr ee (Anch is tus deman i , Onycocar is s tenolep is , and

Pon ton io hur i i) of the 20 ponton i ines r ecor ded by H olthu is

226 DEVANEY AND BRUCE

TABLE 2

Sym b iot i c Decapod Cr u s tacea Assoc iatedw i t h Cr i no ids at Enewetak A tol l

Sym bion t‘ H os t cr i no idi‘No. sym bion ts /N o. hosts

Pa laemon idae

A r aiopon ton ia odon tor hyncha

Palaemonella pott s i

Coman thus ben nettiComan thus par vicir r us

Comas ter gr aci lisPer icl imenes ? ambo inens is Coman thus bennett i

Coman thus parv ici r r us

Per icl imenes commen sal is

Per icl imenes tenuisPer icl imenes p i l ipes

Coman thus bennett iComan thus ben nettiComan thus par vici r r us

Coman thina schegel i

Pon ton iop s is coman thi Coman thus bennett iComan thus par v icir r u s

Alpheidae

Comas ter gr aci lisComan thina schegel i

Athanas sp .

Synalpheus car inatusComan thus par v icir r u s

Synalpheus s t imp son i Coman thus bennettiComan thus par vici r r us

Comas ter gr aci lisGalatheidae

Coman thina schegeliAllogalathea e legan sComan thus bennett i

Par thenop idae

H ar r ovia elegan s Comas ter g raci lis‘Determ inations or ver ifi cat ions of palaemon ids made by AJE ; alpheids

by A . H . Banner ; galatheid by K . Baba; par thenop id by Ann Fielding .

TDeterm inat ions of Coman thina schegeli and Coman thus par vic i r r u s

resulted fr om work by Debbie Zmar z ly, Scr ipps Insti tution of Oceanogr aphy ,dur ing her more extens ive research on cr ino id eco logy and symbionts in

1 980 .

( 1 953 ) fr om other atol ls in the Mar shal l Is lands have beenfound at Enewetak .

The gnathophyll id shr imp , H ymenocer a p icta, has beenobser ved at Kwajalein and Enewetak atol ls . At Enewetak ,a pai r was seen at n ight wel l back in a cave (S. Johnson ,

per sonal comm un ication) and a spec imen photog raphed inthe Enewetak quar r y . An ov iger ous specimen of

G nathophyllum amer icanum was co l lected in the Enewetaklagoon . Bo th gnathophyllids are w idely d is tr ibutedthr oughout the Indo-Wes t Paci fic reg ion ; the latt er speciesis also found in the tropica l eas ter n Paci fic and Ca r ibbean .

Thr ee species of alpheid shr imps wer e r ecor ded at

Enewetak by Chace (1 955) based on col lect ions fr om the

nor ther n Mar shal l Is lands . Mor e intens ive s tudy on

Enewetak alpheids was made by A . H . Banner in Febr u

ary and Mar ch 1 957 . The r esults of th is latter wor k(Banner and Banner , 1 960 ; 1 968) incr eased the knownnum ber of alpheids fr om Enewetak to 49 spec ies includ ingtwo new spec ies of A thanas (three spec ies of A thanas

l isted on l y fr om Mar shal l ls lands”in these two paper s

‘In 1 96 1 a fi r e destr oyed many of the specimens upon wh ich

these two paper s were based (Banner and Banner , For

tunately, r epresentatives of most of the species were depos i ted inthe MPRL r efer ence co l lection and ar e st i l l avai lable.

include specimens fr om Enewetak) . The 1 968 paper is

especial l y impor tan t in pr ov id ing s tat ion data w i th co l leet ion local i ty , depth , and habi tat .‘Beg inn ing in 1 976 , examinat ion of symbion ts associated w i th comatul id cr ino ids at

Enewetak r evealed two species of Synalpheus (S. car inatusand S . s t imp son i) and an undeterm ined species of A thanas

(Table One or mor e spec ies of alpheids have also beenobser ved together w i th gobi id fishes in bu r r ows made in

Enewetak Lagoon sed imen t . Seven alpheid spec ies werefound by H ighsm i th ( 1 98 1 ) as as soc iates of s ix d i ffer entcor als . On l y two , A lpheus acu tofemoratus and A . par

vi ros t r is , wer e found in mor e than two cor al heads of the

same species , and the former alpheid was found in all butone of the cor als s tud ied . Altogether , 52 iden t i fied alpheidspec ies in seven genera are now known fr om Enewetak of

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228 DEVANEY AND BRUCE

Pac i fic water s includ ing the Mar shal l Is lands (Chace,

Two cr angon id shr imps wer e taken in the EnewetakLagoon . One i s poss ibly a new species of Pon toph i lus ,

whereas the other , Ver co ia g ibbosa, was pr ev ious ly knownon l y fr om south Aus tr al ia.

PALIN URA

At leas t four palinu ran lobs ter s ar e r epr esented at

Enewetak . Panu l i r us pen ici llatu s is a common inhabi tant ofthe w indwar d r eef flat and was r ecor ded from Ar no Ato l lin the southern Mar shal l Is lands (H olthu is , Th isspiny lobs ter was the focus of populat ion and other l i fe hi stor y s tud ies at Enewetak dur ing 1 978 and 1979 (For d etal . , 1 979 ; Eber t and For d , Juven i le (puer u lus and

pos t-puer u lus ) s tages of th i s w ide-spr ead Indo-Wes t Paci ficrock lobs ter have been r ecor ded by Cooke and McDonald

(198 1 ) fr om Enewetak . Three spec imens of P . long ipesfemor is t r iga were iden t i fied fr om a co l lect ion made in a

surge channel near the outer reef . A l though not prev ious l yrepor ted fr om the Mar shal l Is lands , th is subspecies is

r epo r ted w idely from Japan , the Mo luccas , New Gu inea,

eas ter n Aus tr al ia and Po l ynes ia (George and H olthu is ,Panu l i r us ver s icolor , r ecor ded by H o lthu is (1 953)

fr om Ar no , has not been repor ted at Enewetak . Par r ibacus

an tar ct icus and Pal inu rellus wienecki i are now known fromEnewetak . Although each of the las t two species has a

broad Indo-Wes t Pac i fic d i s tr ibut ion , on ly P . an tarct icus

had prev ious ly been r ecor ded fr om the Mar shal l Is lands(Ar no) by H olthu is (op . The latter has also beenr ecogn i zed in water s off Kwajelein Ato l l (S. Johnson , per

sonal comm un icat ion) .

COLLECTION RECORDS

Taxa ar e l i s ted in the same or der as in the checkl is tunder thei r r espect ive fam i l ies . H igher categor ies may befound in the checkl is t . These recor ds ar e not known to

have been pr ev ious ly publ ished . When known , the is let ,month , year , habi tat , depth , co l lector , ident i fier , and

number of spec imens are given . Many of the spec imensco l lected between 1 956 and 1971 and determined byL . B . H olthu is (LEH ) are depos i ted at the Allan H ancockFoundat ion . Other specimens have been depos i ted in

sever al ins t i tut ion s , main ly MPRL , EPEM (H ono lulu),RMNH (Leiden ) , and USNM . Ident i fier s AJB and DMD

r efer to the author s ; other ident i fier s are named in full .

Fam i l y PENAEIDAEMetapenaeops is bor r adai lei (De Man )Enewetak Is let : Sept . 1 980 , lagoon , nor th end of is let ,in sandy subs trate us ing ai r l i ft , 1 2 m , DMD and P . Ocr

l in , id. LEH ; 1 spec . (juv . F) . Medr en Is let : Aug . 1 956 ,

lagoon s ide , l ight at n ight , id. LBH ; 1 spec .Metapenaeops is sp .

Aomon , B iji le, Lojwa Is lets : July 1 959,lagoon s ide,

sand bo ttom , to m , F . C . Z iesenhenne, id.

LEH ; 1 spec . Enewetak Is let : Feb. 1 982 , on sand at

n ight (“ bur y ing 5 m , S . Johnson , id. LBH ; 1

spec . (M ) . Medr en Is let : July 1 959 , lagoon s ide, sandbottom , to m

, J . S. Gar th , id. LBH ; 1 spec .Fam i ly SICYONIIDAESicyonella maldiven s is Bor r adai le

Medren Is let : Aug . 1956 , lagoon =s ide, l ight at n ight , id.

LBH ; 9 spec . (3 lots ) .Fam i ly STENOPODIDAEM icrop r os thema p lum icom e (R ichter s)Enewetak Is let : Dec . 198 1 , lagoon , under deadAcropor a, 4 m , S. Johnson , id. P . Gal loway and DMD ;1 spec . , ov ig . ; BPBM .

M icrop r os thema scabr icaudatum (R ichter s )Alembel Is let : July 1 959 , lagoon s ide, Acropo r a cor al ,J . S. Gar th , id. LBH ; 1 spec . ; BPBM .

Odon tozona scu lp t icaudata H olthu is

Medren Is let : Feb . 1 982 , lagoon s ide,under dead coral ,

5 m , S. Johnson , id. DMD ; 1 spec . , ov ig . ; BPBM .

Odon tozona sp .

Lagoon : Sept . 1 982 , in submerged par t of cement sh ipat n ight , 5 m , S. Johnson ; 1 spec . ; Apr i l 1 983 , exposedon r ather s i lty bo ttom next to bulkhead of submer gedpar t of cemen t sh ip , at n ight (ca . 2030 h), 5 m , S. Johnson ; 2 spec .

Stenop us hisp idus (Ol iv ier )B iken Is let : July 1 975 , oceans ide, rotenone s tat ion , 9 m ,

P . B . Lam ber son , id. DMD ; 1 spec . ; BPBM . Reefer 8P innacle: June 1 982 , in cave, 8 m , S. Johnson , id.

DMD ; 2 spec . (1 ov ig . , 1 male) ; BPBM .

Stenop us tenu i r os t r is De Man

Enewetak Ato ll : May 1983 , lagoon , under sheet ofalum inum debr is , 3 m , S. Johnson ; id. DMD ; 1 spec . ;BPBM .

Stenop us zanz ibar icus B r uceEnewetak Is let : May 1983 , lagoon , under sheet of alum inum debr i s , 3 m , S. Johnson , id. DMD ; 2 spec . ; BPBM .

Fam i ly RHYNCH OCINET IDAERhynchoc inetes hiatt i H olthu is and H ayash iEnewetak Ato l l : May 1 978 , algal r idge, recover ed fr omfish po ison (r otenone) s tation , id. AJE ; 5 spec .

Rhynchoc inetes r igens Gor donJaptan Is let : Feb. 1 982 , lagoon , in submerged metallocker , 1 2 m , S. Johnson , id. LEH ; 2 spec .

Rhynchocinetes sp . aff . R . r igens

Japtan Is let : May 1978 , lagoon s ide, 1 00 yds . off p ier atbase of coral head , id. LEH ; 1 spec . Lagoon Moor ingBuoy P innacle: Feb. 1 982, on ledge, n ight , 1 0 m

,

S. Johnson , id. LBH ; 1 spec ; BPBM .

Fam i l y PALAEMON IDAEAnch is tus aus t r a lis B r uceAnan ij Is let : Aug . 1968 , in Tr idacna squamosa (Lam ) ,J . W . Knudsen , id. AJE ; 1 spec . Jinderol Is let :Mar ch 1 96 1 , p r obably in Tr idacna noae (Rbding)[Tr idacna max ima R . Palumbo , id. LEH ; 2

spec . , ov ig . Medr en Is let : July 1 959 , r eef between southend of is let and wreck on r eef, under large pancakecoral cemented to reef , F . Ziesenhenne, id. LBH ; 1

spec . , ov ig .

CRUSTACEA DECAPODA

Anchis tus m ier s i (De Man )Alembel Is let : July 1 959 , nor th s ide, Acropo r a cor al ,J . S. Gar th , id. LEH ; 3 spec . Bokoluo Is let : Feb . 1968 ,

in Tr idacna sp . , J . W . Knudsen , id. AJB ; 2 spec .Jinedrol Is let : Mar ch 196 1 , lagoon s ide, probably inH ippop us hippopus , shal low water , R . Palumbo , id.

LEH ; spec .A r iopon ton ia odon tor hyncha Fuji no and M i yakeMedren Is let : July 1 975 , ocean s ide, on cr ino idComan thus bennett i , 1 2 m , Lamber son , id. AJE ; 1 spec .Medr en Is let : Feb. 1 976 , lagoon , ca . mi le off is let , oncr inoid C . bennett i , 6 to 9 m , D . M . Devaney and

A . Field ing , id . AJB ; 2 spec . ( 1 male, 1 ov ig . female) ;BPBM .

B r achycarp us biungu icu latus (L ucas)B iken Is let : May 1 946 , 2 m i les south of is let , leewar ds ide of r eef , l ight at n ight , L . P . Schultz , id. LEH ; 1 9

spec .Conchodytes biungu icu latus (Pau lson)Pole P innacle: Nov . 1 977 , lagoon , m , coll . and id.

Ann Field ing ; spec .Conchodytes meleag r inae Peter sEnewetak Is let : Apr i l 1 96 1 , lagoo n s ide, in P inctada?galtsoff i, 6 m , E. S. Reese, id. J . Yaldwyn ; 2 spec . (1male, 1 female) .

Conchodytes t r idacnae Peter sLojwa Is let : Feb . 1971 , in s ide Tr idacna gigas , J . Shoup ,id. LBH ; 4 spec . (2 July 1 976 , ins ide Tr idacnamax ima, id. Ann Field ing ; spec .

Co rall iocar is br evi ros t r is Bo r r adai le

Enewetak Is let : July 1967 , in Acr opora cor ymbosa, jus tbeh ind algal r idge, J . W . Knudsen , id AJE ; 1 spec . ,ov ig .

Cor alliocar is gr am inea (Dana)Alembel Is let : July 1 959, lagoon , nort h s ide, Acropor a,

J . S. Gar th , id. LBH ; 2 spec . Anan ij Is let : Feb . 1 957 ,

lagoon s ide, dead Acr opora, m , A . H . Banner , id.

LBH ; 3 spec . Anan ij Is let : Jan . 1 960 , lagoon s ide, nor thend, Acr opor a, E . S. Reese, id. LEH ; 2 spec . Anan ijIs let : Jan . 1 960 , lagoon s ide, fr om Ser iatOpo ra, E. S.

Reese, id. LBH ; 2 spec . B iken Is let : Feb. 1 957 , oceans ide, inner edge reef flat , on Acr opo r a and Poc i llopo r a,

m , A . H . Banner , id. LEH ; 2 spec . , ( 1 male, 1

lkur en Is let : July 1 959 , ocean s ide, Acropor a,

J . S. Gar th , id. LBH ; 2 spec . Medr en Is let : July 1 959,

lagoon s ide,l iv ing Acropor a, J . Coatswor th , id. LEH ; 1

spec .Corall iocar is nudi r os t r is (H eller )Boken Is let : July 1 957 , southeas t s ide, l iv ing coral ,J . S. Gar th , id. LEH ; 1 spec . (female) . Enewetak Is let :Apr i l 1 961 , in Acr opor a hyacin thus , E . S. Reese, id.

LEH ; 2 spec . Medr en Is let : Jul y 1 959 , lagoon s ide,

coral , to m , J . Coatswor th , id. LBH ; 2 spec .Cor al liocar is super ba (Dana)B iken Is let : Aug . 1 956 , ocean s ide in cor al head on r eefflat , common , found among cor al branches , id. LBH ; 1

spec . Boken Is let : July 1 957, southeas t s ide,l iv ing

cor al , J . S. Gar th , id. LEH ; 1 spec . Bokoluo Is let :

229

Mar ch 1 968 , in Acr opo r a digi t ifera, on ocean r eef flat ,J . W . Knudsen , id. AJB ; 1 spec . , (male) . Enewetak Is let :Apr i l 1 96 1 , in Tu r binar ia? danae, E . S. Reese, id. LEH ;

1 damaged spec . , ident i ty no t fully cer tain . Enjebi Is let :July 1 959, shore, F . C . Ziesenhenne, id. LBH ; 2 spec .Japtan Is let : Jan . 1960 , lagoon s ide, nor th end,

Acropo r a, E. S. Reese, id. LBH ; 2 spec . Jin im i Is let :July and Aug . 196 1 , lagoon s ide, m , J . Shoup , id.

LBH ; 2 spec .Cor all iocar is venus ta KempEnewetak Is let : Lagoon cor al mound , m i le off is let ,fr om l ive Acropo ra, 1 0 m , co l l . S. Swer dloff , id. AJB ; 1

spec . Enewetak Is let : Aug . 1 969, lagoon , in

Acropo r a sp . , 10 m , J . W . Knudsen , id. AJB ; 1 spec .

Fenner a chacei H o lthu is

Enewetak Is let : Feb. 1 976 , ocean s ide, co l l . wi th cr ino idComan thus schelegi , 23 m , P . Lam ber son , id. AJE ; l

spec .H arp i l iop s is beaup res i i (Andou in)B iken Is let : July and Aug . 1 961 , lagoon s ide, m ,

J . Shoup , id. LBH ; 5 spec . Boken Is let : July 1 957,

southeas t s ide, Poc i llopo ra coral , J . S. Gar th , id. LBH ;

5 spec . Enjebi Is let : July 1 959, shore, F . C . Ziesen

henne, id. LEH ; 2 spec . Japtan Is let : Jan . 1 960 , lagoons ide, Poci llopo r a cor al , 3 m , E . S. Reese, id. LEH ; 2

spec . Medr en Is let : July 1957 , lagoon s ide, cor al heads ,3 to m , J . S. Gar th , id. LBH ; 1 spec . Medr en Is let :July 1 959 , lagoon s ide, fr om Poci llopo ra dam icor n is ,

to m , J . Coatswor th , id. LBH ; 2 spec .H arp i l iops is dep r es sa (St impson)Anan ij Is let : Feb . 1 957 , lagoon s ide, Poc i llopor a cor al ,m

, A . H . Banner , id. LEH ; 2 spec . B iken Is let : Julyto Aug . 1 96 1 , lagoon s ide, m , J . Shoup , id. LBH ;

10 spec . Enewetak Is let : Apr i l 1 96 1 , in Ser iatopo r a?hys tr ix , E . S. Reese, id. LBH ; 2 spec . Enewetak Is let :July and Aug . 196 1 , lagoon s ide, and 1 2 m ,

J . Shoup , id. LBH ; 8 spec . (2 lots ) . lku ren Is let : July1 959 ,cor al , J . S . Gar th , id. LBH ; 3 spec .

H arp i l iop s is sp in igera (Or tmann)Lojwa Is let : July 1 967 , beh ind r eef on south s ide of

is let , dead and over gr own cor al w i th smal l amount ofl ive cor al , in Stylophor a p is t i llata (mordax) , 2 m , J . W .

Knudsen ,id. AJB ; 1 spec .

Jocas te japon ica (Or tmann)B iken Is let : July and Aug . 1 96 1 , lagoon s ide, m ,

J . Shoup , id. LEH ; several spec .Jocas te lucina (Nobi l i )Alembel Is let : July 1 959, nor t h s ide, l iv ing Acropo r aco r al , J . S. Gar th , id. LEH ; 1 spec . Anan ij Is let : Feb.

1 957 ,lagoon s ide,

dead Acr opo r a cor al , m , A . H .

Banner , id. LBH ; 2 spec . Anan ij Is let : Jan . 1 960 , nor thend, lagoon s ide

,Acropor a coral , E . S. Reese,

id. LEH ;

2 spec . (2 lots ) . B iken Is let : Aug . 1956 , ocean s ide, for

mal in wash ings fr om cor al , D . Reish , id. LEH ; 4 spec .B iken Is let : Feb . 1 957 , ocean s ide, inner edge r eef flat ,fr om cor al , A . H . Banner , id. LBH ; 1 spec . B iken Is let :July and Aug . 1 96 1 , lagoon s ide, m , J . Shoup , id.

230 DEVANEY AND BRUCE

LEH ; 3 spec . Boken Is let : July 1957 , southeas t s ide, l iv

ing coral , J . S. Gar th , id. LEH ; 5 spec . Enjebi Is let : July1 959 , shore, F . C . Ziesenhenne, id. LEH ; 2 spec .lkur en Is let : July 1 959, ocean s ide, liv ing Acropo racor al , J . S. Gar th , id. LEH ; 1 spec . Japtan Is let : Jan .

1 960 , lagoon s ide, nor th end, Acropo ra coral , E . S.

Reese, id. LEH ; 2 spec . (2 lots ) . Jin im i Is let : July andAug . 196 1 , lagoon s ide, m , J . Shoup , id. LEH ; 1 0

spec . Medr en Is let : July 1 959, lagoon s ide, l iv ingAcropor a, J . Coatswor th , id. LEH ; 3 spec .

Leandr ites cyr tor hynchus Fuj ino and M i yakeJaptan Is let : Feb . 1 982 , lagoon off Japtan pier , in metalshr imp locker , 1 2 m , S. Johnson , id. LBH ; 6 spec . (1

Onycocar is quadr atOphthalma (Balss )lku ren Is let : Aug . 1956 , ocean s ide, sponges fr om undersur faces of r ocks , D . J . Reish, id. LEH ; 5 spec .

Palaemonella po tts i (Bo r r adai le)Enewetak Is let : Feb. 1 976 , ocean s ide, on cr ino idComan thus par vici r r u s , 30 m , P . Lam ber son , id. AJE ; 1

spec . Enewetak Is let : Feb . 1 976 , lagoon (M in i PowerPlan t P innacle) , on 3 cr ino ids Comas ter gr aci lis , 6 to 12

m, D . M . Devaney , id. AJB : 6 spec . (3 lots : 1 ov ig . and

1 juv . ; 1 male;Palaemonella r o tumana (Bor radai le)B iken Is let : Feb . 1 957 , ocean s ide, inner edge of reefflat , fr om cor al , m , A . H . Banner , id. LBH ; 1 spec .Bokonbako Is let : July 1957 , ocean s ide, old cor al heads ,m , D . J . Rei sh , id. LEH ; 1 spec .

Palaemonella tenuipes DanaEnewetak Is let : July 1 957 , ocean s ide, r eef flat , J . S.

Gar th , id. LBH ; 4 spec . Medr en Is let : Feb . 1957 ,lagoon

s ide, m ixed coral , m , A . H . Banner , id. LBH ; 1

spec . Medr en Is let : July 1 957, lagoon s ide, cor al , 3 to

m , J . S. Gar th , id. LEH ; 2 spec .Palaemonella sp .

Anan ij Is let : Feb . 1 957, lagoon s ide, Acr opo r a coral ,m , A .

’

H . Banner , id. LBH ; 1 spec .Par anch is tus armatus (H . M i lne Edwar ds)A lembel Is let : Sept . 1 956 , lagoon s ide, from Tr idacnagi gas , id. LEH ; 1 spec . Jinedrol Is let : Mar ch 1 96 1 ,lagoon s ide, pr obably in Tr idacna g igas , shal low water ,R . Palumbo , id. LBH ; 1 spec . ov ig .

Per icl imenaeus sp .

Medr en Is let : Feb . 1 957 , lagoon , south end,patch reef

about 600 ft o ffshor e, su r round ing bo ttom sand , thr eedead , overgr own coral heads (Poc i llopor a, Acr opo r a,

Por ites) br oken up , m , A . H . Banner , id. LBH ; 1

spec .Per icl imenes agag KempEnewetak Lagoon : Jan . 1 982, under dead co r al , 5 m ,

S. Johnson , AJE ; 1 spec .Per iclimenes ?am bo inen s is (De Man)Enewetak Is let : Feb . 1 976 , ocean s ide, on cr ino idComan thus par vic i r r us , 3 1 m , P . Lamber son , id. AJB ; 1

spec . Medr en Is let : Feb . 1 976 , lagoon , ca . m i leoff is let , on 3 cr ino ids Coman thu s bennetti , 6 to 9 m ,

DMD and A . F ield ing , id. AJB ; 6 spec . (3 lots , eachw i th male and ov ig . female) .

Per icl imenes bayer l H o lthu i sEnewetak Is let : Apr i l 196 1 , fr om Poc i llopo r a ver r ucosaor P . e legans coral , E . S. Reese, id. LBH ; 3 spec .Enewetak Is let : July and Aug . 1 96 1 , ocean s ide,

Poc i llopor a? e legans cor al , 1 5 m , J . Shoup , id. LBH ; 2

spec . Enewetak Is let : July and Aug . 1 96 1 , lagoon s ide,

m , J . Shoup , id. LEH ; 2 spec .Per iclimenes commen sal is Bor r adai le

Enewetak Is let : Feb . 1976 , lagoon p innacle ca . V2 to 1

m i le off is let , on cr ino id Coman thus bennetti , 5 m , DMD

and A . Field ing , id. AJE ; spec . Medren Is let : Feb .

1 976 , lagoon , ca . m i le off is let , on 4 cr inoidsComan thus bennett i , 6 to 9 m , DMD and A . Field ing , id.

AJE ; 54 spec . (4 lots : 1 6 ; 4 p lus pos t- lar va: 22 including 1 ov ig . ; 9 including 2

Per icl imenes cr is t imanu s Br uceEnewetak Is let : Aug . 1 968 , r ock quar r y , on sea ur ch inEch inoth r ix calamar is , J . W . Knudsen , id. AJE ; 6 spec .(2 lots , one w i th 5 including ov ig . , another w i th 1

Per icl imenes den t icu latus Nobi l iB i llae and Run i t Is let : July 1 967 , acr opor id cor al andalgae in lagoon dr edge hauls between the two is lets , 29to 32 m , J . W . Knudsen (Stn . id. AJE ; 2 spec .

Per icl imenes elegans (Paulson )Alembel Is let : Feb . 1 957 , reef flat , coral , m , A . H .

Banner , id. LBH ; 1 spec . (d is car ded— mo ldy) . Anan ijIs let : July 1 957 , ocean s ide, r eef flat , J . S. Gar th , id.

LEH ; 1 spec . B iken Is let : July 1959, southeas t s idebetween ocean and lagoon , rock w i th sand subs tr ate,

dead coral , J . S. Gar th , id. LEH ; many spec . BokoluoIs let : Aug . 1 968 , ocean reef, fr om Por i tes sp . , J . W .

Knudsen , id. AJE ; 1 spec . , av ig . Bokonbako Is let : July1 957 , lagoon s ide, old cor al heads , D . Rei sh , id. LEH ; 1

spec . Enewetak Is let : July 1 957, ocean s ide, r eef flat ,J . S. Gar th , id . LEH ; 5 spec . Enewetak Is let : Aug .

1968 , J . W . Knudsen ,id. AJB ; 4 spec . (1 Enjebi

Is let : July 1 957 , ocean s ide, r eef flat , J . S. Gar th , id.

LBH ; 1 spec . Japtan Is let : Aug . 1 968 , inner r eef flatpoo l , J . W . Knudsen , id. AJB ; 7 spec . (3 Japtan

Is let : Aug . 1 968 , lagoon t ide poo ls , J . W . Knudsen , id.

AJE ; 13 spec . (5 male, 8 female w i th 7 of theseJaptan Is let : Aug . 1 968 , r eef flat poo l , po i son s tat ion , J .W . Knudsen , id. AJB ; 7 spec . , (2 Lojwa Is letJuly 1 967 , in Stylophor a p is t i llata (mordax) m , J . W .

Knudsen , id. AJB ; l spec . Lojwa Is let : Ocean r eef pool ,J . W . Knudsen , id. AJB ; 1 spec . (male) .Per icl imenes ens if rons (Dana)Lujor Is let : June 1 946 , outs ide lagoon , in ter t idal , M . W .

Johnson , id. LBH ; 1 spec .Per icl imenes gr andis (St impson)Kedrol Is let : May 1 946 , inter t idal , in pot holes , M . W .

Johnson , id . LEH ; 1 spec .Per iclimenes holthu is i B r uceEnewetak Is let : Mar ch 1 982 , lagoon s ide,

found in

exposed tentacles of por i t id coral (AlveOpo r a or

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232 DEVANEY Al

bottom sand , 3 dead , overgr own coral heads ,Poc i llopora, Acropor a, Por ites , br oken up , m , A . H .

Banner , id. LBH ; 2 spec . Medr en Is let : July 1 957,

ocean s ide, o ld cor al heads , m , D . J . Reish, id.

LBH ; 2 spec . Medr en Is let : July 1957 , coral heads and

dead cor al , 3 to 5 m , T . Goreau and R . Nesh ida, id.

LEH ; 1 spec .Fam i ly PROCESSIDAEN ikoides s tein i i (Edmondson)Boken Is let : Sept . 1 956 , ocean s ide, oppos i te channelen tr ance,

cemen ted coar se cor al sand r ock , D . J . Reish,id. LBH ; 1 spec .

N iko ides m u lt isp inatus H ayash iEnewetak Is let : Jan . 1 982, lagoon at n ight , 5 m , underdead Acropo ra coral , S. Johnson , id. K . I . H ayash i , 2spec .Pr ocessa ?japon ica (De Man)Medr en Is let : Lagoon s ide, dr edging oppos i te m iddle of

is land , sand bo ttom , 2 to 3 frn s . F . C . Ziesenhenne, id .

LBH ; 1 spec .Pr ocessa mo lar is ChaceB iken Is let : Feb . 1 957 , ocean s ide,

inner edge of outerr eef flat , fr om coral , m , A . H . Banner , id. LBH ; 1

spec . Enjebi Is let : July 1959, shor e, m tide,

F . C . Ziesenhenne, id. LBH ; 1 spec . Medren Is let : July1957 ,

lagoon s ide,coral , 3 to m , J . S. Gar th , id.

LEH ; 1 spec .P roces sa neg lecta H ayash iMedr en Is let : June 1 957 , lagoon s ide , sand and wormtubes , 3 m , D . J . Rei sh , id. LEH ; 2 spec .Fam i ly PANDAL IDAEM i r opandalu s harding i B r uceR ibewon Is let : June 1 982 , on br anch of antipathar ian(black cor al) in smal l cave, 23 m , S. Johnson , id. AJB ; 1

spec (M) .Fam i ly TH ALASSOCARID IDAEThalas socar is cr inata (Dana)Lagoon Aug . 1 968 , dr edge, 60

m , J . Knudsen , id. AJB ; 2 spec .Fam i ly CRANGON IDAEPon tOphi lus sp . aff . P . sabsechota KempMedr en Is let : July 1 959 , lagoon s ide, dredg ing oppos i tem iddle of i s land , sand bottom , to 6 m , F . C .

Ziesenhenne, id. LBH ; 1 spec .Verco ia g ibbo sa BakerMedr en Is let : 1 982 , smal l lagoon p innacle at n ight ,L . Boucher , id. AJE ; 1 spec .Fam i ly SYNAX IDAEPal inu rella wienecki i (De Man )Enewetak P innacle: June 1 98 1 , in smal l cave, 1 0 m ,

co l l . and id. S. Johnson ; 1 spec . (car apace of mo lt) .Fam i l y PAL INUR IDAEPanu l i r us long ipes femor is t r iga (von Mar tens )Enewetak Is let : Oct . 1969

, ocean s ide su rge channel ,

EA DECAPODA 233

1 976 , Shr imps of the Pas iphaeid Genus Lep tochela wi thDescr ipt ions of Thr ee New Species (Crustacea : Decapoda:Car idea) , Sm ithson . Con t r ib . Zoo l . , 222: 1 -51 .

Cooke, W . J. , and C . D . MacDonald, 1 98 1 , The Puer u lu s and

Pos t-Puer u lus of the H awai ian Spiny Lobster Panu l ir us marginatus , Pr oc . B io l . Soc . Wash . , 94 : 1 226- 1 232 .

Ebert , T . A . , and R . F . Ford , 1 986 , Population Eco logy and

Fishery Potential of the Spiny Lobster Pan u l ir us penci llatus at

Enewetak Ato l l , Mar shal l Is lands , Bu l l . Mar . Sci . , 38 : 56-67 .

Edmondson , C . H . , 1 952 , Addi tional Centr al Pacific Crustaceans ,Occas . Pap . Ber n ice P . B ishop M us . , 21 : 67-86 .

Fankboner , P . V . , 1 972, On the Association Between the Pon

ton i id Shr imp Ach is tus m ier s i De Man (Decapod , Palaemon idae) and Giant Clams (Lamel l ibranch ia, T r idacn idae) , Wasman

J . B io l . , 30 : 35-42 .

Ford , R . E . , T . A . Eber t , M . McCollum and D . Bar t ley , 1979,

Population Character istics and Fishery Potential of Spiny Lobs ter s at Enewetak Ato ll , MPRL Ann . Rept . F Y 1 978 — 79.

George, R . W . and L . B . H olthu is , 1965 , A Rev i s ion of the IndoWest Paci fi c Spiny Lobster s of the Panu li r u s japon icus Group,Zoo l . Ver hand, No . 72 , pp . 1 -36 .

H ighsm ith , R . C. , 1 98 1 , Coral B ioeros ion at Enewetak : Agentsand Dynam ics , In t . Revue Gesamen . H ydrobiol . 66 : 335

-375 .

H olthu i s , L . B .

, 1 953 , Enumeration of the Decapod and Stomatopod Crustacean from Pacific Coral Is lands , A to l l Res . Bu l l .No . 24 , pp . 1 -66 .

and K . I . H ayash i , 1 967, A New Species of Shr impRhynchoc inetes h iatt i (Crus tacea, Decapoda), Annot . Zoo log .

Japon . , 40 : 1 6 1 -1 70 .

Rosewater , J. , 1 965 , The Fam i ly Tr idacn idae in the lndo—Pacifi c,Indo -Pacific Mo ll , 1 : 347-393 .

Zmar z ly, D . L . , 1 984 , Di str ibution and Eco logy of Shal low-WaterCr ino ids at Enewetak Ato l l , Mar shal l Is lands , wi th an Annotated Check l is t of Thei r Symbionts , Pac . Sci . , 38 : 1 05-1 22 .

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236 GARTH , HAlG , AND KNUDSEN

fam i l y Coenobi t idae, genus Coenobita, and the coconutcr ab , B i rgus latr o . B rachyur an cr abs found in the ter r es tr ialenv i r onm en t ar e grapso id cr abs of the fam i l ies Gecar cin idae, Ocypodidae, and a few Graps idae .

The spr ay zone on rocky shor e is inhabi ted by theSally L ight foo t cr ab , her e Grapsus tenu icr u s tatus r atherthan Gr apsus gr apsus L innaeus , 1 758 . The h ighes t elevat ions of sandy beaches ar e inhabi ted by the graps id cr absCyclogr ap s us and Pseudograp s us ; the in termed iate elevat ions (w i th bu r r ows extend ing below water level) ar e ln

habi ted by the ghos t crab , Ocypode cer atophthalma. The

mo le cr ab , H ippo pacif ica, an anom ur an , l ives in the sur fzone,bur ying i tself qu ickly and emerg ing sudden l y to gr asp

i ts food w i th i t s fi r s t pai r of legs . Analogous s i tuat ions on

muddy beaches (r ar e at Enewetak) ar e inhabi ted by theocypodid cr abs Macr ophthalm us and Uca. Where so i l accumu lates , as at the bases of coconut palms , the land cr abGecar cin us , fam i l y Gecar c in idae, bur r ows . The col lectorwho tu r n s in land w i l l encounter the gr aps ids Geogr ap sus

cr in ipes and G . gr ayi among leaf l i tter and the

Metasesarma and Sesarma among r oo ts and low branches .

It was no acciden t that the co l lector r espons ible formos t of the r ecor ds of land crabs in the Pac i fic ScienceBoard ’

s Cor al A to l l Su r vey (H olthu is , 1 953 ) was F . R . Fos

ber g , a ter r es tr ial botan is t , or that the fi r s t cr abs to reachthe sen ior author soon after ar r iv ing at Enewetak wer ebrought to him by Edwar d H eld and Ralph Palumbo ,

mem ber s of the U n iver s i ty of Wash ingt on gr oup s tudy ingthe effects of r ad iat ion on ter res tr ial or gan isms . Thus ever yhabi tat avai lable to them , both ter res tr ial and mar ine, has

been co lon i zed by these ubiqu i tous ar thr opods , the

decapod cr us tacean s .

CRABS AS SYM B IONTSThe relationsh ips between cr abs and the corals in

wh ich they may be found var y w idel y , fr om par as i ti sm and

commensal ism (o r mutual i sm) to facultat ive s ymbios i s . In

paras i t ism , the crab bur r ows in to the coral or the cor al

gr ows around the crab , in bo th cases enclos ing i t . In com

mensal ism , the crab , although appar en tly fr ee- liv ing ,

invar iably selects a l iv ing coral of a par t icular k ind as i tshos t . Facultat ive symbios i s i s a r elationsh ip in wh ich thecrab , whi le often found in l iv ing cor al , is also found in deadcor al , in cor al r ubble, or even on a noncor al , r ubblysubs tr ate .

The tr ue paras i tes , long thought to include on ly thecoral gal l crabs and cor al-bur r owing crabs of the fam i lyH apalocar cin idae, ar e now known to include some of the

appar en t ly fr ee- l iv ing cr abs of the fami ly Xanthidae as

wel l — in par t icular , the gener a Tr apez ia and Tetr al ia, the

fo rmer found on poci llopor id, the latter on acropor id

corals . These wer e shown by Knudsen ( 1967) to feed oncor al po l yps , wh ich they macerate wi th thei r speciallyadapted dactyls before inges t ing . The genera Domecia and

Cyma, d i ffer ent species of wh ich occur on poci llopor id and

acr opo r id cor als , appar entl y have s im i lar feed ing habi ts ,

CRABS AS SUBJECTS FOR RESEARCHThe Anomura and B r achyur a of Enewetak A tol l have

pr oven valuable as subjects for r esear ch . The terr es tr ialherm i t cr abs , Coenobita, wer e used by Un iver s i ty of Was hington Schoo l of Fi sher ies per sonnel in determ in ing re

s idual activ i ty fr om tes ts conducted by the Atomic EnergyCommis s ion in the late 1 9405 and ear ly 19505 (H eld ,

Reese ( 1968a) used the coconut crab , B irgus lat r o ,

in li fe h is tor y s tudies show ing use of a shel l by the g laucothoe lar va. H erm i t crabs , Coenobita, and the ghos t cr ab ,Ocypode ceratophthalma, were used by Reese in mot ionpicture s tudies of locomotion . Exper iments conducted by

although on l y those of Domec ia have been inves t igated(Patton ,

Ot her crabs found in coral appar entl y use i t only forshelter and pr otect ion fr om pr edator s . These include not

on l y xan th id cr abs l ike L iomera (=Carp i lodes ) and

Pseudol iomera— some spec ies of wh ich. rarel y , i f ever ,

occu r elsewhere— but also a hos t of gener a such as

Chlorod iella,Phymod ius , and P i lod ius , wh ich ar e found

abundan t ly in dead coral and coral r ubble. These are

jo ined by cr ab genera of o ther fam i l ies : the smaller swimming cr abs of the genus Thalam i ta,

fami ly Por tun idae; anumber of spider cr abs , includ ing Per inea and

Tylocar c in us , fam i ly Maj idae; herm i t cr abs of the genusCalcinus , fam i l y D iogen idae; Cor all iogalathea and Calathea,

fam i l y Galatheidae; and por celain cr abs of the gener a

Pachycheles and Pet rol is thes , fam i ly Por cel lan idae.

Smal l xanth id cr abs of the genus Actum n us have beenobser ved to car r y a smal l piece of l ive coral as they moveabout the ocean bo ttom , much as a herm i t crab car r ies a

gas tr opod shel l . Actum nus an telm i War d , the species or igi

nal ly descr ibed as hav ing th is habi t , occu r s at Enewetak ,but i t s coral-car r y ing pr opens i ty was not obser ved (Lamber ts and Gar th ,Cr abs that u se not a coral po lyp but an act in ian

coelenterate, wh ich they car r y in each claw to fend off

attacker s , are mem ber s of the genera Lybia and

Polydectus . Several species of diogen id herm i t cr abs of the

genus Dardanus car r y anemones on thei r shells . In bothcases , the crab benefi ts from the s t inging nematocys ts of

the coelenter ate, and the anemone ach ieves mobi li ty andprobably food as wel l . The po r celain cr ab Neopetr olis thes ,fam i l y Por cel lan idae, inhabi ts several spec ies of largeanemones .

Cr abs assoc iated wi th l iv ing mo llusks include the p in

nother id Xan thas ia mu r igera, found in the man tle cavi ty of

the g iant clam , Tr idacna gigas . Cr abs as soc iated wi th liv ingech inoderms include the par thenopid H ar r ovia elegans ,

found on the cr ino id Coman thus , and the galatheids

A llogalathea e legans and Calathea ambo inens is , also foundon cr ino ids . The por t un ids L is socar cin u s or bicu lar is and

L . ho lothu r ico la found in the respi rator y tr ee or cloaca of

the sea cucum ber s H o lothur ia at r a Jaeger and Act inopyga

maur it iana (Quoy and Gaimard) proved imposs ible to part it ion between thei r ho lothur ian hos ts .

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 237

Knudsen ( 1967) showed that the cor al-inhabi t ing xan th idcr abs , Tr apez ia and Tetralia, ate cor al po lyps aft er fi r s tmacerat ing them and, hence, wer e tr ue par as i tes ratherthan commensals . The coral bur r ow ing cr abs , fam i l yH apalocar cin idae (Fi ze and Serene, wer e used byKnudsen in unpubl ished s tud ies . The por tun id cr ab ,Thalam i ta in tegr a , was used by Pomer oy and Kuenz ler

(1 967) in s tud ies of phosphor us tur nover by co r al r eefan imals . H ighsmi th (1 98 1 ) involved the xanth id crabs ,Tetr o lic and Maldivia, in s tud ies of cor al er os ion by invertebr ates and fishes . In ter t idal crabs of the fam i l y Xanthidaewer e ut i l i zed by H avens ( 1974) in s tud ies of compet i t iveexclus ion (the par t i t ion ing of food r esou r ces) . Wenner(1 977) and Wenner and Fusar o ( 1 979) conducted s tud iesof populat ion s tr uctur e and dynam ics us ing the Paci ficmo le cr ab , H ippo pacif ica . The xanth idcr abs — Dacryop ilumnus erem ita, E r iphio sebana,

Phymadius ungu lotus , P i lum nus long icom is , and Trapez iaspeciosa— were shown by Dan for th ( 1967 , 1 970) to hos tepicar id i sopod paras i tes of h i ther to undescr ibed species .

The hermi t cr abs Dardanus wer e shown by H umes (1 971 )to be hos ts of harpact ico id copepods . The herm i t cr absCa lcinas and D iogenes wer e used by Or ians and K ing(1964) in s tudies on shel l select ion and invas ion r ates .

Sys temat ic s tudies have been published on the genusPet rolis thes , fam i ly Por cel lan idae, by H aig on

coral- inhabi t ing cr abs by Gar th and on sw imm ingcr abs , fami l y Por tun idae, by Stephenson and Rees

wi th addi t ional s tud ies by Gar th , H aig , and Knudsen in

prog r ess .

PO ISONOUS CRABS

Du r ing the 1 9705 Gar th and Alca la (1 977) showedbeyond a reasonable doubt that numer ous reef- inhabi t ingcrabs of the Indo-Wes t Paci fic ar e harmful when eatenbecause they ar e po isonous . Included among these are

several common Enewetak spec ies : Doldorf ia ho r r ida (L innaeus ), A tergot is f lor idus (L innaeus ), E r iphio sebana (Shawand Nodder ) , and Zozymus oeneus (L innaeus) . The fi r s t isan elbow cr ab , fam i ly Par thenop idae; the remain ing thr eeare mem ber s of the fam i ly Xanthidae, as ar e mos t of thecrabs found to be tox ic to man and to domes t ic an imals .

The po i son , a saxi tox in , i s chem i cal l y ind is t ingu i shablefrom that pr oduced by cer tain mol lusks . It causes vom i ting ,fo l lowed by locomotor y and neur olog i cal s ymptoms wh ich ,i f not treated , result in paral ys is and death .

Cr ab-caused fatal i t ies have been documented , and thecr abs were ident i fied by competen t taxonom i s ts in the

Ryukyu Is lands , the Phi l ippines , and Palau . Nat ive popu lati ons of many South Sea i s land groups have tr ad i t ions of

k i l ler crabs and ver nacular names for the po isonousspecies . A lthough no po i son ings fr om cr abs are known to

have occur r ed at Enewetak Ato l l , cr abs known to be po i

sonous elsewhere ar e common on the reef at Enewetak .

Caution i s ur ged in the handl ing of these crabs (a per sonwho has handled such a crab m ight exper ience numbnes s

after touch ing his tongue to hi s hand) . Abs t inence fr omthei r cul inar y u se i s also adv ised .

FOSSIL CRUSTACEANSFoss i l anomur an and brachyur an decapod species

obtained by U . S. Geological Sur vey dr i l l ings at Enewetakand r epor ted by Rober t s (1 964) include Coll ichi r us armatus(A . M i lne Edwar ds) , Coll ichi r us ar t icu lotus (Rathbun) ,Actaeodes hi r s u t is s im us (R i ippell), and Et is us laevimanus

(Randal l) . A lthough known from elsewhere in the

lndo-Wes t Paci fic , these four species have not been foundl iv ing at Enewetak . Th is could mean ei ther that subtlechanges have occur r ed in the r eef env i r onmen t that r enderEnewetak no longer a su i table habi tat or , as seems mor el ikel y , that the su i te of spec ies inhabi t ing Enewetak ischanging cons tantly as new species ar e in troduced , becomees tabl i shed , ar e el im inated by compet i tor s , and becomelocal ly ext inct un t i l r ein troduced in another cycle. It is alsoposs ible that fur ther and mor e d i l igen t sear ch ing may yetuncover these four species at Enewetak in the l iv ing s tate.

‘Th is secti on was pr ov ided by Jens Knudsen .

COLLECT ING DECAPODCRUSTACEAN S‘

Enewetak Atol l ascends abr uptly to the sur face wher ethe Nor th Equator ial Cur r ent , the pr evai l ing trade w inds ,and the ocean ic waves s tr ike the atoll . Waves refr act allthe way ar ound the atol l reef and penetrate the lagoonthr ough channels or over the algal r idge . Therefor e, we

fel t that ever y compass po int , fr om w indwar d to leewar d ,would have un ique phys ica l factor s that influenced dis tr ibut ion of both cor al species (p lus morphotypes of coralspec ies) and decapod crus taceans . Th is proved usefu l .Since sever al other exper ts wer e to receive cr us taceanspecimens beyond ou r interes t , much t ime was spent hun ting new cor als , cr ino ids , algae, and sed iments that harbo red decapods .

The inter t idal zone i s shal low (based on Kwajaleininformat ion), yet wh i le zones ar e compr essed , a cen t imeterelevat ion on the reef flat wou ld usual ly y ield some d i ffer en tbr ach yur an . We also bel ieved zones d i r ectly below and

above the t idal range wer e relat ivel y shal low but becamethicker (or deeper ) down '

into the lagoon depth , or to the

height of land and i t s vegetat ion on i s lands .

Th is , in theor y , kept us close to the inter t idal zonewher e we thought the greates t species dens i ty of decapodswas to be, and appar ently is , found . We collected on all

but thr ee of the 34 named is lands . The nor thwes t chainfr om Bogallua Is land to Bogon Is land was cons ider ed toodangerous for camp ing tr ips as opposed to 1 -day M

-boatr uns . Ou r r ad io car r ied les s than 5 m i les and the Equator ial Cur r en t flowed wes twar d . Later , the Gar th — Knudsenteam s found the res tr icted access l im i ted the wor k on thenor thwes t chain as compar ed to that done on the mor eacces s ible i s lands .

238 GARTH , HAIG , AND KNUDSEN

The windwar d algal r idge r eceives the lar ges t waveswh ich super saturate incom ing water w i th oxygen . Algae

gr ow her e in a pr ofus ion of species and mass . In i tial l y weused p ry bar s to loosen s labs fr om the algal r idge. Thesewer e placed in plas t ic bags or buckets wh i le another teammember attempted to secur e all fr ee decapods . Slabs werecar r ied to the near es t i s land for crack ing and co l lect ing of

decapods . Subsequently , a r ig id , heavy-welded pipe cracking tab le w i th a car -deck ing top was des igned and bu i lt forthe purpose of cr ack ing coral or s labs at any s i te includ ingthe algal r idge (F ig . Buckets , hammer s , and otherequ ipment wer e secured to the table . A mar ker al lowednote tak ing (even under water ) , so s tat ion number s werei s sued and r ecor ded as needed . Natur al l y someone heldthe table when ver y large waves wer e r unn ing . The y ieldof species was dr amati ca l l y incr eased by the use of the

cr ack ing table.

I llum inated by a Co leman lanter n , the r eef-flat and algal

r idge wer e also co llected at n ight w i th excel len t succes s .

The algal r idge off Enewetak Is land was also co l lected du ring waveless do ldr um s

— un t i l two lar ge tsunamic wavescame sho r ewar d , throw ing Knudsen 30 feet back onto ther eef flat .

Smal l amounts of formaldehyde wer e appl ied fulls tr engt h to the r eef flat at low — low t ide . The pr eservativedi luted in patches of water and enter ed worm bur rows ,ev ict ing numerous decapods wh ich otherwise would havebeen imposs ible to col lect . Beh ind the r eef flat , for rnal

dehyde was in jected 6 inches deep in cor al r ubble. Soonafterwar ds , decapods that r esembled ch ips of coral wor kedup to the sur face and wer e captur ed w i th a guppy net .

On ou r fi r s t tr ip , we snor keled ever y day and averagedabout 5 to 1 0 m i les of sw imm ing per day in local ar eas .

A lways new corals or new formations , new wave patterns ,r eef dr ainage cur r en ts , etc . , pr ov ided new recor ds . Plas ticbags wer e placed over cor als o r cr inoids , and the cor aland/or cr ino id was r emoved w i th decapod species in tact .The author was towed by our s low mov ing outboar d sk i ffto sur vey mi les of lagoon mar g in . A hand s ignal mean tnew cor al fo rmat ion —

or pos s ible shar k attack . We

snor keled down to about 60 feet in ques t of some cr ino idsand corals .

Co l lect ing was success fu lly attempted w i th a dr edgebu i lt and out f i tted at Pac i fic L uther an Un iver s i ty (PLU ) .We used the dr edge, powered by a sk i ff , in shal low water(to 30 feet) . In deeper lagoon water , we used an M-boat inr ever se, wi th the dr edge r ope and buoy r eady to go over

boar d when fouled . Mar ker s placed and recor ded to allowwor k to continue the next day were never found again .

H owever , many r are and some new r ecor ds of cr abs werewor th the effor t . A cable and w inch , as opposed to pu llingthe cable by hand , would have greatl y faci l i tated lagoons tud ies .

Is lands pos sessed man y species of decapods on landand even in tr ees . Since is lands ar e s cat ter ed ar ound muchof the reef, and r efr acting waves s tr ike is lands di fferen tly ,

F i g . l Cr ack i ng table on the algal r i dge at Enewetak , abou t 1000 ft fr om land.

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240 GARTH , HAIG , AND KNUDSEN

Calcinas elegan s (H . M i lne Edwards , Reese, 1969 .

Ca lcinas gaimordi i (H . M i lne Edwards , Reese, 1969 .

.

1'

Colcinus guamens is Wooster , 1984 .

IlCalcin us sp . indet . #2 .

"I'

Colcin us imper ialis Whi telegge, 1901 .

Calcinas laevimanus (Randall , Reese, 1 962; Reese, Reese, 1969 .

Calcinas latens (Randall , Provenzano , 1 963 ; Or ians and King , 1964 ; Reese, 1969 ; H umes , 1971 .

Calcinus seurat i Forest , 1 951 : Reese, 1 969.

°

l Colcin us sp . , aff . sp icatus Fores t , 1951 .

°

1Cl ibanar iu s sp . , aff . boschmai Bu i tendijk , 1 937 .

Cl ibanar iu s coroll in ir s (H . Mi lne Edwards , Reese, 1 969 .

‘Cl ibanar ius eu rys ter nus (H i lgendor f,

°

1 Cl ibonor ius zebra rhobdodoctylus Forest, 1953 .

1lCl ibonar ius zebra var . r hobdodoctylus .

‘l’

Cl ibanar ius sp .

‘l‘Dardonus crass iman us (H Mi lne Edwards ,Dardanus deform is (H . Mi lne Edwards ,

‘TDardanus gemmatus (H . Mi lne Edwards ,Dar danus gu ttatus (O l iv ier , H umes , 1971 .

Dardanus lagopodes (For sskal , H umes , 1971 .

Dar danus sanguino len tas (Quoy and Gaimard, Pr ovenzano, 1 963 .

IlDardanu s sangu ino len tus

Dar danus meg is tosDar danus scu tellatus (H . Mi lne Edwards , Pr ovenzano , 1 963 ; 0 r ians and King, 1964 ; Gar th , 1 964 ; H umes , 1971 .

Dardanus woodmoson i (A lcock , Gar th , 1964 .

D iogenes gardiner i A lcock , 1905: Provenzano, 1 963 ; Or ians and King, 1964 .

IlD iogenes sp .

‘TD iogenes pallescens Whi telegge, 1 897.

‘TPagu r is tes sp .

‘TTr iz opagu r u s s tr igotus (H er bst,Fam i ly PAGURIDAE.

1'Cotapagu r u s sp .

”(Pagu r ixus anceps (Forest , McLaughlin and H aig , 1 984 .

l lPagu r us sp . (in par t).IlPogu r us (Pagu r ixus) sp . 1 .

Pagu r ix us bon inens is (Melin , McLaughlin and H aig, 1 984 .

Pagur ixus moor us (Nobi l i , McLaughl in and H aig, 1984 .

1lPagu r us sp . (in par t) .llPagu r us (Pagu r ixus) sp . 2 .

°'

l New genus , sp .

Superfam i ly GALATH EOIDEAFam i ly GALATH EIDAEA llogalathea elegan s (Adams and Wh i te, Baba, 1 977 ; Baba, 1 979 ; Bruce and Zmar zly, 1 983 .

llGalothea elegans .

°

TCoroll iogolotheo hum i lis (Nobi l i ,llGalotheo t r iden t ir os tr is M iyake, 1953 .

‘TGolathea aegypt iaca Paulson , 1 875.

‘TGalathea aff in is Or tmann , 1 892 .

IlGoIotheo sp inosor os t r is Dana, 1 852 (in par t) .IlGalotheo sp . , aff . aus tr al iens is Stimpson , 1 858 .

”l Galatheo ambo inen s is De Man , 1888 .

°

1’

Calathea sp . , aff . tonegashimae Baba, 1 969 .

l lG olathea sp inosor os tr is Dana, 1852 (In par t) .°

TPhylladior hynchus ser r i r os tr is (Mel in ,

IlGolatheo ser r i ros tr i s .

Fam i ly PORCELLAN IDAE°

Ne0 pet rol is thes maculatus (H . M i lne Edwards ,'Pochycheles johnson i H aig , 1965 .

“Pachycheles scu lptas (H . M i lne Edwards ,‘New Enewetak record .

"( New Mar shal l Is lands r ecord .

l lJ. S. Gar th manuscr ipt li sts .

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA)

‘TPachycheles p iso ides (H el ler ,‘i Pachycheies spin ipes (A. M i lne Edwards ,

‘TPetr olis thes as latlcus (Leach,‘1'Petr olisthes bisp inos us Bor radai le, 1900.

Pet r olis thes bo r r adailei Kropp, 1984 : Kr opp, 1984.

llPetrol is thes lamarcki i (Leach , 1820) (in par t) .‘1'Petrol is thes coccineus (Owen ,

°

TPetrolls thes decacan thus Or tmann , 1897 .

Petrolis thes elegans Haig, 1981 : Haig, 1981 .

Petrolisthes fimbr iatus Borradai le. 1898 : H ighsmith, 1981 .

Petrolisthes lamarcki i (Leach,"l"Petrolis thes mosok'll Miyake, 1943 .

‘1‘Petrolis thes pen ici llatus (Heller ,"l’

Petr olis thes pubescens Stimpson, 1858 .

"I‘P

'

etrolls thé s [undescr ibed sp . 1 , R. K. Kropp MS].‘i P

'

etr ollsthes [undescr ibed sp. 2, J . Haig and R . K . Kropp MS].ilPetfollsthes deeaeanthus Ortmann, 1897 (in par t).

Infraorder BRACH YURASection DROMIACEASuper fam i ly DROMIOIDEAFam i ly DROMIIDAE‘TCryp todrom ia canaliculata Stimpson, 1858 .

‘TCryp todramla sp.

Fami ly DYNOMENIDAEDynamene hispida Desmarest , 1825: H ighsmith, 1981 .

‘TDynomene p ilumnoides Alcock , 1899.

‘Dynornene

'

p raedatar A. Mi lne Edwards , 1879.

‘Dynamene spinosa Rathbun, 191 1 .

Section OXYSTOMATASuperfami lyCALAPPOIDEAFami ly LEUCOSIIDAE‘1'Cryp tocnemus haddon l Calman, 1900 .

‘1'Ebalia woodmason i Alcock , 1986.

Ebollop s is erosa (A. Mi lne Edwards , Gar th, 1964 .

Heter olithadla sp.

"l'Heleronuela venusta Nobi l i . 1906 .

°

1Merocryptus durandl Serene, 1955.

”rMyrafugax coalito H i lgendorf , 1878.

"l'Nucio ingens (Rathbun,

‘l'Nucio speciosa Dana, 1852.

°'l‘Or eophor us (Oreotlos) lotus Bor r adai le, 1903 .

‘1‘Species incer tae sedis .

Fami ly CALAPPIDAE'l'Calappa calappa (L innaeus ,Calappa gal lus (H erbst,

‘Calappa hepatica (L innaeus , Rober ts , 1964 .

Section OXYRH YNCH ASuperfam i ly MAJOIDEAFam i ly MAJ IDAE°‘l'

Campascio retusa Latrei l le, 1825 .

°

1 Camposcio sp .

°

Cyclax suborbicular is (Stimpson ,‘l'

H uen ia brevifrons Ward, 1 941 .

H uen ia pr oteus De H aan , 1839: Gar th , 1964 .

°

1 H yas tenus irom i (Laur ie,

°

1 H yos tenus unc ifer Ca lman , 1900 .

”l H yas tenus ver r ucos ipes (Adams and Wh ite,

°

1 H yos tenus sp .

‘New Enewetak r ecord.~

(New Marshall Is lands record .t Foss i l record.1lJ. S. Gart h manuscr ipt l ists .

24 1

242 GARTH , HAIG , AND KNUDSEN

Menaethius monocer os (Latrei l le, Gar t h , 1 964 .

°

1 M icippa morgar i t ifer o H ender son , 1 893 .

M icippa ph i lyra (H er bst , Gar th , 1 964 .

°

1 M ic ippa p latypes R i ippell , 1 830 .

°

1 M icippa thalia (H er bst ,°

TNax iodes sp in iger a Bor r adai le, 1 903 .

"I Par atymolus bi tuber cu lotu s M ier s , 1882 .

Paratymolus sexsp inos us Mier s , 1 884 : Gar th , 1 964 .

°'

l‘Por ozewa bocki Baiss , 1 938 .

Per inea tum ida Dana, 1 851 : Gar th , 1964 .

Schiz ophrys aspera (H . M i lne Edwards , Gar th , 1 964 .

°

1 Tr igono th i r obtu s i ros tr is Mier s , 1 879 .

"I Tylocar c inus ?gr aci lis M ier s , 1879.

Tylocar c in u s s tyx (H er bst , Gar th , 1 964 .

Super fam i ly PARTH ENOPOIDEAFam i ly PARTH ENOPIDAE"l Actaeomorpha sp . , nr . erosa (M ier s ,"l“ Cr yptopod ia ?pan Laur ie, 1905 .

'Doldo rf io hor r ida (L innaeus ,

"l Doldorfio (or Par thenope) sp .

H ar r ov io e legans De Man , 1 888 : Gar th , 1 964 .

°

1 H eter ocryp to? sp .

‘l'Par thenope (Au locolombr us) cu r visp in is (M ier s ,

°

l Por thenope (Au locolombr us) hop lonotu s (Adam s and Wh ite,

°

TPar thenope (Au locolombr us) sp .

'1

’ Par thenope (Pseudo/ombr us) sp .

Par thenope sp . : Gar th , 1 964 .

°

TThyro lombr us er os us (M ier s ,.

1'

Thyrolombr us sp .

Section CANCRIDEASuper fam i ly CORYSTOIDEAFam i ly ATELECYCL IDAE‘TKr ous s ia in tegr o Rathbun , 1 906 .

‘1

’

Kr ous s ia sp . , cf . mar quesa Serene, 1 972 .

°

TKr ouss ia n i t ida St impson , 1 858 .

o

l Krous s ia ros t r ipes F . M il lier , 1 857 .

°

l Krous s ia rugu losa (Kr auss ,

Section BRACH YRH YNCH ASuper fam i ly PORTUNOIDEAFam i ly PORTUN IDAE

‘Corupo tenuipes Dana, 1 851 .

llCar upa laeviuscu la H el ler , 1 86 1 .

‘TCotop t r us inaequal is (Rathbun ,

‘Catop t r us n it idus A . M i lne Edwards , 1870 .

‘TCotop t r u s rathbunoe Serene, 1 965 .

”r Cotop t r us ? t r uncot ifr ons De Man , 1887 .

‘Charybd is (G on iosup r aden s) erythrodoctylus (Lamarck ,

"I Coelocarc inus fo l iotus Edmondson , 1930 .

"I“ L ibis tes v i l losus Rathbun , 1 924 .

‘TL is socar c in us ho lo thu r icola Streets , 1 877 .

‘TL is socar c in us or bicu lar is Dana, 1 852 .

Por tunus (Ache lous) gr on u lotus (H . M i lne Edwards ,Por tunus gr onu lotus : Gar th , 1 964 .

Por tunus (Achelous) sp . , nr . or bicu lar is (R ichter s ,Por tunus or bicu lar is (R ichter s ) : Gar t h , 1 964 .

Po r tun us (H ellenu s) long isp inos us Stephenson and Campbel l , 1 959 .

Por tunus longi sp inos us (Dana, Gar t h , 1964 .

Thalam ita admete (H er bst , Gar th , 1 964 ; Stephenson and Rees , 1 967 .

Thalam i ta bouvier i Nobi l i , 1 906 : Stephenson and Rees , 1 967 .

‘New Enewetak r ecord .

‘

l'New Mar shal l Is lands record .

1lJ. S . Gar t h manuscr ipt l ists .

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244 GARTH , HAIG , AND KNUDSEN

Cymo dep lanatus A . M i lne Edwards , 1 873 : Gar th , 1 964 .

Cymo melanodactylus De H aan , 1 835: Garth . 1964 .

'Cymo quadr i lobatus Mier s , 1 884 .

Dacryop i lumnus erem ita Nobi l i , 1 906 : Danfor th, 1970.

‘TDacryop i lum n us rathbunoe Baiss , 1932 .

‘Dai ra per loto (H er bs t ,Domecia g labr a A lcock , 1 899 : Gart h , 1964 .

Domecia h isp ida Eydoux and Sou leyet , 1 842 : Gar th , 1 964.

‘E r iphio scabr icu lo Dana, 1 852 .

E r iphia sebana (Shaw and Nodder , Reese, 1 969; Danfor th , 1970 .

IlE r iphia laevimono Guer in , 1 838 .

‘I'

Et is us bifr on tal is (Edmondson ,

°'

l'

Et is us deman i Odhner , 1925 .

°

I Et is us sp . , nr . deman i Odhner , 1 925 .

‘Et is us den tatus (H er bst ,Et is us elect ra (H er bst , Gart h , 1964 .

‘1

’

Et is us f ron tolis Dana, 1 852 .

TE t is us laevimanus (Randal l , Rober ts , 1964.

°

TEt is us molokoiens is (Rathbun ,

"l'Et is us sp lendidu s Rathbun , 190621: Rober ts , 1964.

°

1 Et is us sp . 1 .

'l Et is us sp . 2 .

"I Et is us sp . 3 .

Euxan thus exscu lp tus (H er bst , Gar th , 1964 .

°'

l'Eux an thus (or H ypocolp us ) sp .

G oi llordiellu s r ueppell i i (Krauss ,Actaea reppell i i [s ic]: Gar th , 1 964 .

Goi llordiellus superci liar is (Odhner ,Actaea super ci liar i s : Gar th , 1 964 .

_

‘G lobop i lumnus g lobosus (Dana,

‘TH eter op i lumnus sp . , cf . long ipes (Stimpson ,

‘Lachnopodus ponopens is (Rathbun ,

IlParaxan thias haematos t ictus Ward, 1 930 .

‘1‘Lachnopodus s ubocutus (Stimpson ,

‘Lachnopodus tahitens is De Man , 1 889.

'Lep tod ius exarotus (H . Mi lne Edwards ,

°

Lep todius gr aci lis (Dana,

°

1 Lep tod ius dovaoens is Ward, 1941 .

°

Lep tod iu s nudipes (Dana,

llXan tho danae Odhner , 1 925 .

°

Lep tod ius sangu ineus (H . M i lne Edwards ,°

TLep tod ius woioluonus Rathbun , 1 906 .

L iocorp i lodes arm iger pacif icus Balss , 1 938 : Gar t h , 1964 ; H ighsm ith , 1981 .

L iocorp i lodes biungu is (Rathbun , H ighsm i th , 1981 .

Zozymodes biungu is : Gar th , 1 964 .

IlXan thod ius biungu is .

L iocorp i lodes in teger r imus (Dana, Gar th , 1964 .

L iocorp i lodes pum i lus (Jacquinot , Gar th , 1964 .

llZozymodes pum i lus .

“Zozymodes cr is tatus (Bo r r adai le,L iomera be lla (Dana,

Carp i lodes be l lus : Gar th , 1964 ; H ighsm i th , 1 98 1 .

L iomera coe lata (Odhner ,Carp i lodes coelatu s : Gar th , 1964 .

‘TL iomero loevis (A . Mi lne Edwards ,1ICarp i lodes loevis .

‘TL iomera mon t icu losa (A . M i lne Edwards ,llCorp i lodes mon t icu losus .

‘New Enewetak record.TNew Mar shal l Is lands record .

t Foss i l reco r d.1IJ. S. Gar th manuscr ipt l i sts .

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA)

°

L iomer a pal lida (Bor r adai le,

llCarp iIodes pal lidus .

‘TL iomera rugata (H . M i lne Edwards ,

“Carp i lodes r ugatus .

”( L iomera s t impson i (A . M i lne Edwards ,

“Carp i lodes s t imp son i .‘L iomera tr is tis (Dana,

“Carp ilodes tn‘

s ti s .

"( L iomera sp .

llCarp i lodes sp .

‘l Lophozozymus dodone (H er bst ,

°l Lophozozym us incisus (H . Mi lne Edwards ,°

Lophozozymus pu lchel lus A . Mi lne Edwards , 1867 .

”I“

Lybia caes t ifera (A lcock ,"l“Lybia tesse llata (Latr ei l le,

‘Lydia onnu l ipes (H . Mi lne Edwards ,Macromedaeus nudipes (A . M i lne Edwards ,l lXan tho nudipes .

Maldivia palmyrens is Rathbun , 1 923 : H ighsm i th , 198 1 .

Maldivia tr iungu icu lata (Bor r adai le, H ighsm i th , 198 1 .

"l’

Medaeus elegans A . M i lne Edwards , 1867.

°

TMedaeus or natus Dana, 1 852 .

°

1'Neoxan th ias impr es sus (Lamarck ,l Xan tho impressus .

°

TParactaea retusa (Nobi l i ,“Actoea retusa .

Paractaea r ufopunctata (H . M i lne Edwards ,Actaea r ufopunctata : Gar th , 1 964 .

"I'Paractaea r ufopunctata f . p lumosa Guinot , 1 969 .

‘TParactaea tum u losa (Odhner ,llActaea tumu losa .

°

TParamedaeus s imp lex (A . M i lne Edwards ,l lMedaeus s imp lex .

°

TParap i lumnus corall iophi lus Takeda and Miyake, 1968 .

Parap i lum nus ver r ucos ipes (Stimpson) : Gar th , 1964 .

°

TPorap i lumnus ?incer tas Takeda and Miyake, 1 969.

ll H eterop i lumn us sp . , nr . quadr isp inosus (Zehntner ,Paraxan thias notaras (Dana, Gar th , 1 964 ; H ighsm ith , 1 98 1 .

‘TParaxan thias pachydactylus (A . Mi lne Edwards ,‘Phymod ius ?gr anu latu s (Targioni -Tozzett i ,

°

Phymodius laysan i Rathbun , 1 906 .

”l Phymod ius mon t icu losus (Dana,

Phymodius n it idus (Dana, Gar th , 1 964 .

Phymod ius ungu latus (H . M i lne Edwards , Gar th , 1 964 ; Danfor th , 1 970 .

P i lodiu s areolatus (H . Mi lne Edwards , H ighsm i th , 1 98 1 .

Chlorodops is ar eo lata: Gar th , 1964 .

Pi lodius flavu s Rathbun , 1906 : Gar t h , 1 964 .

‘1

‘P i lodius melanodactylus (A . M i lne Edwards ,l lChlorodops is melanodactylus .

P i lodius p i lumno ides (Wh ite,Gar th , 1964 .

llChlorodops is p i lumnoides .

Pi lodius pugi l Dana, 1 852: Gar th , 1 964 .

t Chlorodops is pugi l : Rober ts , 1964 .

°

TP i lod ius scabr icu lus Dana, 1852 .

Pi lodiu s sp in ipes H el ler , 1 861 : Gar th , 1964 .

l lChlorodops is sp in ipes .

”( Pi lumnus ander som

‘

De Man , 1 887 .

‘Pi lumnus oaer u lescens A . Mi lne Edwards , 1 873 .

‘Neyv Enewetak r ecord.?New Mar shal l Is lands r eco rd .t Foss i l record .i lJ. S. Gar th manuscr ipt li sts .

246 GARTH . HAIG , AND KNUDSEN

‘1‘Pi lumnus ?elegans De Man , 1 888 .

Pi lum nus longicom is H i lgendor f , 1 878 : Gar th , 1964.

Pi lumnus sp . : Danfor th , 1 970 .

‘TP i lum nus r onsan i For est and Guinot , 196 1°

TP i lum nus r otumanus Bor r adai le, 1900 .

'Pi lum nus tahiten s is De Man , 1 890 .

‘Pi lum nus vesper t i lio (Fabr icius ,‘1'Pi lum nus sp .

P lanop i lumnus verm icu latus (A . Mi lne Edwards , Gar th , 1 974 .

Polydectus cupu l ij'er (Latrei l le, Gar th , 1964 .

‘1'Pseudol iomera gr anos imanus (A. Mi lne Edwards ,"l'Pseudol iomera he l ler i (A . Mi lne Edwards ,TlActaea hel ler i .

°

TPseudol iomera sp . , nr . hel ler i (A . Mi lne Edwards ,l lActaea sp . , nr . he l ler i .

'Pseudo l iomera lata (Bor r adai le,

l lActaea lata.

°

TPseudo l iomera sp . , nr . lata (Bo r r adai le,

l lActaea sp . , nr . lata.

°

TPseudol iomera r ueppell ioides (Odhner ,l lActaea r ueppell io ides .

Pseudol iomera speciosa (Dana,

Actaea speciosa: Gar th , 1964 .

°

Pseudoz ius cays tr us (Adam s and Wh ite,

°

Pseudoz ius pacificus Balss , 1 938 .

"l Ralum ia dah li Balss , 1933 .

Tetr al ia g laber r ima (H er bst , Gar th , 1964 ; Knudsen , 1 967 .

°

Tet ro lic glaber r ima r ubr idactylu s Patton , 1 966 .

Tet r alo ides n igr ifr ons (Dana,

Tet r o lic heterodacytyla H el ler , 1 86 1 : Gar th , 1 964 ; Knudsen , 1 967 .

llTetr al ia heterodacytyla fusca Serene, 1 959 .

l lTet ral ia ?n igr ifrons Dana, 1 852 .

Tr apez ia cymodoce (H er bs t , Gar th , 1964 ; Knudsen , 1 967 .

"I“

Trapez ia den tata Macleay , 1 838 .

Tr apez ia digi talis Latrei l le, 1825: Gar th , 1964 ; Knudsen , 1 967 .

"l“ Trapez ia digi tal is be lla Dana, 1 852 .

Trapez ia sp . , dig i talis group: Gar th , 1964 .

Trapez ia fer ruginea Latrei l le, 1 825 : Gar th , 1964 ; Knudsen , 1967‘TTrapez ia gu ttata R ii ppell , 1 830 .

Tr apez ia r ufop unctata (H er bs t , Gar th , 1 964 ; Knudsen , 1967 .

"l'Tr apez ia r ufopunctata flavop unctata Eydoux and Souleyet , 1 841 .

‘1

’

Tr apez ia r ufopunctata macu lata Macleay , 1838 .

Trapez ia speciosa Dana, 1852 : Gar th , 1 964 ; Danfor th , 1970 .

Trapez ia t igr ina Eydoux and Sou leyet , 1 842 .

Trapez ia danai Ward , 1939 : Gar th , 1964 ; Knudsen , 1 967 .

Tr apez ia wardi Serene, 1 970 .

:l Tr apez ia sp . 1 .

Trapez ia sp . 2 .

fl Xan th ias canal icu latus Rathbun , 1 906 .

l Xan th ias gi lber tens is Balss , 1 938 .

Xan thias lamar ck i (H . M i lne Edwards , H ighsm i th , 1981 .

.

l Xan th ias lividus Lamar ck , 1 808 .

.

l Xan thias p unctatus (H . M i lne Edwards ,c

l Xan tho sp .

.

l Zozymodes cavipes (Dana,

1'

Zozym us actaeo ides (A . M i lne Edwards ,l lP latypod ia actaeo ides .

Zozym us sp . , n r . actaeo ides (A . Mi lne Edwards ,llPlatypod ia sp . , nr . actaeo ides .

‘New Enewetak reco rd .

TNew Mar shal l Is lands record .

i lJ. S. Gar th manuscr ipt l i sts .

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248 GARTH , HAIG , AND KNUDSEN

SUMMARYThe Anom ura pr esen tl y known fr om Enewetak Atol l

compr ise 76 species , r epr esen t ing 29 gener a and 1 0 famil ies . Of th i s num ber , 48 spec ies are new to Enewetak , and43 ar e new to the Mar shal l Is lands as wel l . The fam i lyD iogen idae i s bes t r epr esen ted , w i th 27 spec ies in seven

gener a; the fam i ly Por cel lan idae has 1 7 species in thr eegener a . Expressed in per cen tages , of the 76 species l i s ted ,

are new to Enewetak , and ar e new to the

Mar shal l Is lands . The D iogen idae con tain of the

species and of the gener a; the Po r cellan idae con

tain of the species and of the gener a.

The Br achyur a pr esen tl y known from Enewetak Ato l lcompr i se 291 species , repr esent ing 1 1 5 gener a and 1 6

fam i l ies . Of th i s number , 21 8 species ar e new to

Enewetak , and 1 70 are new to the Mar shal l Is lands as

wel l . The fam i ly Xan th idae is bes t r epr esen ted , w i th 1 55species in 49 gener a; the fam i l y Por tun idae by 36 speciesin n ine gener a; the fam i ly Maj idae by 23 species in 1 3 gener a; and the fam i ly Gr aps idae by 21 species in 1 0 genera.

Expres sed in per cen tages , of the 291 species ar e

new to Enewetak and are new to the Mar shal lIs lands as wel l . The Xan thidae contain of the spe

cies and of the genera ; the Por tun idae containof the species but on ly of the genera; the

Maj idae contains of the species and of the

gener a; and the Gr aps idae contain of the speciesand of the gener a r epor ted . (The num ber of speciesand genera new to Enewetak and the Mar shal l Is landswould be even gr eater had not many been repor ted in publ icat ions in wh ich the cr abs , often iden t i fied by the fi r s twr i ter , wer e not the pr imar y in teres t of the inves t igatorwho r epo r ted them but inciden tal as the hos ts of i sopod orcopepod par as i tes or as agen ts of bioeros ion of cor als . )

Because so large a number of brachyur an crabs r emainidenti fied .to genus (35) or even to fami ly (3) on ly , no

mean ingful com par ison w i th other cr ab faunas is poss ibleat th is t ime . Th is shor tcoming w i l l be r ect i fied when the

Enewetak crabs are elabo rated a fami l y at a time and the

new or obscur e species descr ibed and i l lus trated . The deficiency is par t icular l y appar en t in the fami ly Par thenop idae,

of wh ich no comprehens ive r ev iew has been made s incethe Siboga r epor t (F l ipse,

ACKNOWLEDGMEN TSThe generous ass is tance in the ident i ficat ion of r ar e or

obscure spec ies of B rachyur a by special i s ts who v is i ted theH ancock Foundat ion wh i le the fi r s t author ’

s s tudies wer ein prog r ess , or to whom specimens wer e later sent , isgratefull y acknowledged . These include R . S. K . Bar nes ,the genus Macrophthalm us ; I . Gor don , the s ing le pinnother id; D . J . G . Gr i ff in , the genus Cyclog r ap s us and the fam

i ly Maj idae; M . Takeda, the P i lumn inae; T . Sakai , theOxys tomata; R . . Serene, the T rapez i inae; and W . Stephenson , the Po r tun idae. The second author ’

s s tud ies on the

Anomur a wer e fac i l i tated by Susan L . B r unenmeis ter , who

pr ov ided in format ion on her Enewetak co l lect ions ; PatsyA . McLaughl in , who sent a por t ion of th i s co l lect ion forthe ver i fi cat ion of a few identi ficat ion s ; and Ph i lip H elfr ich ,who pr ov ided a copy of the Br unenmeis ter r epor t (Br unenmeis ter , For the thalass ino ids , M ichele de SaintLaur en t and L . B . H olthu is helped w i th ident i ficat ions andR . B . Mann ing w i th in format ion . The author s are mos tgrateful to the late Denn is M . Devaney and to Ann F ieldingfor pr ov id ing informat ion concer n ing Enewetak specimensin B i shop Museum co l lections , and mos t of all , to the manyco llecto r s whose recor ds have been used w i th thei r permiss ion in cases wher e i t was pos s ible to contact them .

REF ERENCES

Baba , K . , 1 977, B io logi ca l Res u lts of the Snell ius Expedi tionXXVIII , The Galatheid Crus tacea of the Snell ius Expedi tion ,

Zoo l . Meded . (Leiden) , 50 : 243 -259 .

1979 , Expedi t ion Rumphius II Crustacés Paras ites ,Commensaux , etc . , (Th . Monod et R . Serene, VII ,

Galatheid Crustaceans (Decapoda, Anomur a), Bu l l . Mus . Nat l .H is t . Nat . (Par is ) , (4) 1 (A ) : 643 -657 .

Balss , H . , 1 938 , Die Dekapoda Brachyu ra von Dr . Sixten BocksPazi fik-Expedi tion 1 91 7— 1918 , K . Vet . O . Vitterh. Sam h . ,H andl . , 8 , 18 5.

Bruce, A . J. , and D . L . Zmar z ly, 1983 , Per iclimenes p i l ipes , New

Species , a Cr ino id Associate fr om Enewetak Ato ll , Mar shal lIs lands (Crus tacea : Decapoda: Ponton i inae), J . Crus t . B io l . , 3 :644-654 .

Br u nenmeis ter , S. L ., 1975 , Effects of Behav ioral In teractions and

H abi tat Complex i ty on N iche Breadths and Over laps of L i t

tor al H erm i t Cr abs on Enewetak , M id-Pac . Mar . Lab. AnnualRepor t 1 975 : 22 -23 .

Danfor th , C. G . , 1 967, Nor thern Pacific G igan tione (Isopoda),B io l . Bu ll . , 132 : 1 47-155.

1 970 , Ep icar ids (Isopoda) of Eniwetok Atol l , Pac . Sci . , 24 :

462-471 .

Fi ze, A . , and R . Serene, 1 957, Les H apaloca r cinide's , du

V iet-Nam , Mem . Ins t . Océanogr . N hatr ang , 1 0 : 1-202 .

F l ipse, H . J. , 1930 , Die Decapoda Br achyur a der Siboga

Expedition , VI , Oxyr hyncha: Par thenop idae, Siboga-Exped. ,

Monogr . , 39cz: 19 6 .

Forest , J. , 1984 ,Rev is ion du Genr e An iculus : Decapoda Ologeni

dae, Crustacea , Cr u s taceana (Leiden) Supp l , No . 8 ,pp . Ix

91 , frontispiece.

Fusar o , C. , 1 978 , Food Avai labi l i ty and Egg Pr oducti on : a FieldExper iment wi th H ippo pacifica Dana (Decapoda; H ippidae),Pac . Sci . , 32 : 1 7—23 .

Gal i l , B . , 1 985, Tetralo ides , A New Genus of Coral-Inhabi tingCrabs , Cr u s taceana (Leiden) , 50 : 68-77 .

and C . Lew insohn,1984 , On the Taxonom ic Status of .

T r apez ia tigr ina Eydoux and Sou leyet , 1 842 (Decapoda,

B rachyura) , Cr us taceana (Leiden ), 46 : 1 66- 1 75 .

Gar th , J . S. , 1 964 , The Crus tacea Decapoda (B rachyur a and

Anomura) of En iwetok Ato ll , Mar shal l Is lands , wi th SpecialReference to the Obligate Commensals of B ranchi ng Co rals ,M icrones ica, 1 : 137- 1 44 .

and A . Alcala, 1977, Poisonous Cr abs of Indo-Wes t Paci fi cCor al Reefs , wi th Special Reference to the Genus Deman iaLaur ie, Proceedings of the Th ir d In ter national Coral Reef Sympos ium , M iam i , F lor ida, 1977, 1 : 645

-651 .

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA)

and T . S. H opkins , 1968 , Pseudocryp tochir u s crescen tus(Edmondson), a Second Crab of the Corallicolous Fami lyH apalocar cin idae (Cru stacea, Decapoda) from the EasternPaci fi c wi th Remarks on Phragrnos is , H ost Specifi ci ty , and

Distr ibution, Bull. Seuth. Calif . Acad. Sci ., 67 : 40 48 .

Gross , W., 1964. Water Balance in Anomuran Land Cr abs on a

Dry Ato ll , Bio l. Bul l ., 1 26 : 54-68 .

H aig , J. , 1981 , Three New Species of Petrolis thes (Decapoda,

Anomur a, Porcel lan idae) from the Indo-West Pacific , J. Crus t .B io l 1 ; 265-271 .

H avens , A. D.,1974, The Ecology of Eight Species of Inter ti dal

Crabs of the Fami ly Xan thidae in the Mar shal l Is lands , Ph.D .

disser tation. Un iver sity of Southern Ca li forn ia, Los Angeles .

H eld, E. E. , 1 960 , Land Crabs and Fiss ion Pr oducts at En iwetokAto l l , Pac . Sci . , 14: 1 8 27 .

H ighsm ith , R . C . , 198 1 , Coral B ioer os ion at Enewetak : Agentsand Dynam ics , In t . Revue Gesam ten . H ydrobiol . , 66 : 335

—375.

H olthu is , L . B . , 1 953 , Enumerat ionfl

of the Decapod and Stomatopod Crustacea fr om Paci fic Coral Is lands , A to l l Res . Bu l l . No .

24 .

1983 , Notes on the Genus Enop lometop us , with Descr ipt ionsof a New Subgenus and Two New Species (CrustaceaDecapoda, Ax i idae) , Zoo l . Meded . (Leiden) , 56 : 281 -298 .

H umes , A . G . , 1 971 , Sunar istes (Copepoda, H arpact icoida) Associated wi th H erm it Crabs at Eniwetok Ato ll , Pac . Sci ., 25:

529-532.

Knudsen , J . W . , 1 967 , Tr apez ia and Tetral ia (Decapoda,

Brachyur a, Xanthidae) as Obl igate Ectoparas ites of Poc i llo

par id and Acr opor id Corals , Pac . Sc i 21 : 51 -57 .

Kropp , R . K . , 1 984, _Three New Species of Porcel lanidae (Crus

tacca; Anomura) fr om the Mar iana Is lands and a D iscuss ion of

Bor radai le’

s Petr o l is thes lamarcki i Complex, Micrones ica, 1991 - 106 .

Lamber ts , A . E . , and J . S. Gar th , 1977, Coral— Crab Commensali sm in Xanthids , Pac . Sci . , 31 : 245

—247 .

Lawr ence, J . M . , 1976 , Or gan ic Compos ition and Energy Contentof the H epatopancreas of H erm i t Crabs (Coenobito) fromEn iwetok Atol l , Mar shal l Is lands (Decapoda, Pagur idea),Cr u s taceana, 31 : 1 13-1 1 8 .

McLaughlin , P . A . , and J . H aig , 1984, A Review of Pagu r ixus

(Decapoda, Anomura, Pagur idae) and Descr iptions of New

Species , Cr u s taceana (Leiden), 47: 121 -148 .

Miyake, S. , 1939 , Notes on Crustacea Brachyura Co l lected byProfessor Tei sa Esaki ’s M icrones ia Expeditions of 1937- 1 938

Together wi th a Check L i st of Micr ones ian Brachyura, Rec.

Oceanogr . Works Jpn . , 10 : 168-247.

1943 , Studies on the Cr ab Shaped Anomura of N ippon and

Adjacent Water s , J. Dep . Agne. Kyushu Imp . Un iv . , 7:

49-158 .

Or ians , G . H . , and C. E. King , 1964 , Shell Selection and Invas ionRates of Some Pacific H erm i t Cr abs , Pac . Sc i . , 18 : 297-306 .

Page, H . M . , and S. W . Wi llason , 1982, Di str i bution Patterns ofTer restr ial H erm it Crabs at Enewetak Ato ll , Mar shal l Is lands ,Pac . Sci . , 36 : 107

- 1 17 .

Patton , W . K .

, 1967 , Studies on Domecia acan thophora, a Com

mensal Crab fr om Puer to R ico , wi th Par ticular Reference toModi fi cati ons of the Coral H os t and Feeding H abits , B io l .Bu l l . , 132 : 56-67 .

Pomeroy, R . , and E. T . Kuens ler , 1967 , Phosphorus Tur noverby Coral Reef Animal s , Sympos ium on Radioeco logy, D . J .Nelson -

and F . C. Evans (Eds ) , Ann Ar bor , M ich igan , pp .474 482.

Provenzano, A.J. , Jr . , 1 963 , The G laucothoes of Petr ochi r u s

Appendix

COLLECTION RECORDS

Fam i ly CALL IANASSIDAECal lianas sa sp .

Enewetak : 1980 , collected from the lagoonP . Col in and D . M . Devaney .

Call iax sp . , aff . novaebr i tann iae (Bo r radai le)Enewetak : 1 980 , col lected from the lagoonP . Co l in and D . M . Devaney .

Coll ichi r us vigi lax (De Man)Enewetak : 1 980 , co l lected from the lagoonP . Co l in and D . M . Devaney .

diogenes (L . ) and Two Species of Dar danus (Decapoda:D iogen idae), Bul l . Mar . Sc i . Gu ll Car ibb . , 13 : 242-261 .

Reese, E. S. , 1 962, Subm iss ive Posture as an Adaptation to

Aggress ive Behav ior in H erm it Cr abs , Z. Tierp sychol . , 1 9 :645-651 .

1 968a, Shell U se: an Adaptation for Em igrati on fr om the

Sea by the Coconut Crab, Science, 1 61 : 385 386 .

1968b, Annual Breeding Seasons of Thr ee Sympatr i cSpecies of T ropical Inter t idal H erm it Cr abs , wi th a Di scuss ionof Factor s Contr o l l ing B reeding , J . Exp . Mar . B io l . Eco l , 2:308-3 18 .

1 969, Behav ior al Adaptations of Intert idal H erm i t Cr abs ,Am . Zoo l . , 9: 343 -355 .

and R . A . K inzie I II , 1 968 , The Lar val Development of theCoconut or Robber Crab B i rgu s lotro (L ) in the Labor atory(Anomur a, Pagur idea) . Cr u s taceana, Suppl . 2: 1 1 7-144 .

Rober ts , H . B ., 1 964 , Foss i l Decapod Crustaceans fr om the

Mar shal l Is lands , Geo l . Sur v. Pr of . Pap . 260-H H : 1 127-1 1 3 1 .

Stephenson , W . , and M . Rees , 1967 , Some Por tun id Cr abs fromthe Pacific and Indian Oceans in the Co llections of the Sm i thsan ian Inst itution , Proc . U . S. Nat l . Mus .

, 120 : 1 - 1 14 .

Wenner , A . M . , 1977, Food Supply , Feeding H abi ts , and EggProduction in Paci fi c Mo le Crabs (H ippo pacifico Dana), Pac .

Sci . , 31 : 39-47 .

and C. Fusar o , 1979, An Analys is of Population Str uctur e inPaci fic Mo le Cr abs (H ippo pacifica Dana), B io l . Bu l l . (Woods

H o le) , 157 : 205-229 .

and S. R . H aley , 1 98 1 , On the Ques tion of Sex Rever sal inMale Crabs (Cru stacea , H ippidae), J. Crus t . B io l . , 1 : 506-517 .

Wooster , D . S. , 1984 , The Genus Calcinus (Pagur idea , Diogeni

dae) from the Mar iana Is lands Including Thr ee New Species ,M icrones ica, 18 : 1 21 1 62 .

Anomu r an Cm s tacea

Taxa ar e li s ted in the same or der as in the checklis tunder thei r respect ive fam i l ies . H igher catego r ies may befound in the checkl is t . For mos t species r epor ted below ,

the r ecor ds are abbr ev iated , giv ing on ly the is let , year , andcol lector ; fo l low ing these recor ds is a br ief s tatement onhabi tat . Where on ly one or two r ecor ds ar e avai lable for aspecies , mor e detai led info rmat ion is included for each . A

summar y of co l lector s , year s of co llect ion , iden ti fier s , anddepos i t ion of mater ial is g iven in Table 1 .

250 GARTH , HAIG , AND KNUDSEN

Thomos s in io sp .

Enewetak : 1 980 , col lected from the lagoon by ai r l i ft ,P . Co l in and D . M . Devaney

Fam i l y CALL IANIDEIDAEColl ion ideo [undes cr i bed sp .]Engebi : 1959 , shore co llect ing ,

-foo t t ide, fr omcoral , F . C. Ziesenhenne . Enyu : 1 959 , shor e co l lect ingat no r th end of is land , -foot t ide, in or under coral ,F . C. Z iesenhenne .

New genus [undes cr ibed sp . 1]Off R ig i l i : 1 957, ocean r eef flat , inner edge about 1 00feet ou t , fr om Acr opora or Poc i llopor a in feet ofwater , A . H . Banner .

New genus [undescr ibed sp . 2]Engebi : 1 959, shor e co l lect ing ,

-foot t ide, fr omcoral , F . C . Ziesenhenne.

Fam i l y AXIIDAEEnop lometop us (Enop lometop us) sp .

Enewetak : 1 969 , in sur ge channel , 0 to 3 feet , C . E .

Dawson .

'

Enewetak : 1 969,in sur ge channel , 2 to 1 0

feet , C . A . Ch i ld .

Enop lometopus (H o lometop us) holthu is i Gor donEnewetak : 1 98 1 , pinnacle, 1 0 m in smal l cave, fr agmen ts of bo th claws , S. Johnson .

EnOp lometopu s sp .

Off Enewetak : 1 946 ,juven i le in E . longi ros t r is s tage,

L . P . Schult z . R igi li : 1 946 , 2 m i les south , leewar d s ideof r eef , l ight at n ight , juven i le in E . long i ros t r is s tage,

L . P . Schultz .New genus [?undescr ibed sp .]Aaraanbi r u : 1 959 , share co l lect ing ,

-foot t ide, fromcor al , F . C. Z iesenhenne.

Fam i l y H IPPIDAEH ippo odoctylo Fabr iciusEnewetak : 1 975 , lagoon s ide southwes t end of is let , 1 0to 20 m nor theas t car go p ier , in sand at water ’

s edge,

C. Fusar o .

H ippo pacif ico (Dana)Bogallua: 1 968 , J . W . Knudsen . Enewetak : 1968 ,

A . H aven s . Igu r in : 1 967, C . V . MacCoy . R igi l i : 1956 ,

s tudent co l lector . On sandy beach , and in coar se sandand gravel on r eef .Fam i ly ALBUNEIDAEPar albunea dayr i t i (Serene and Umal i )Enewetak : 1 966 , dr edged in lagoon at about

1 62°20 ’E in 45 m of water , J . W . Knudsen .

Enewetak : 1 980 , co l lected fr om the lagoon by ai r l i ft ,P . Co l in and D . M . Devaney .Fam i ly COENOBITIDAEB i rgus lat ro (L innaeus )Igu r in : 1 960 , E . 5 . Reese and R . A . Booloo t ian . Mu i :

1957 , L . Donaldson and E. H eld . R ig i l i : 1 957,J . S.

Gar th . On land .

Coenobi to br evim on us Dana

Aaraanbi r u : 1957, J . S. Gar th . Igu r in : 1 960 , E . S.

Reese and R . A . Boo loo t ian . Mu i : 1957, L . Donaldson

and E . H eld . R ig i l i : 1 957 , J . S. Gar th . On land.

Coenobito cavipes St impsonIgur in : 1 960 , E . S. Reese and R . A . Boo loo t ian . Mu i :

1 957 , L . Donaldson and E . H eld . Mu i : 1 966 , J . W .

Knudsen . R ig i li : 1 960 , B . Sather and R . Stevenson .

In land and on beach .Coenobi to per latus H . M i lne Edwar dsAaman , B iji le, and Rojoa : 1 959 , F . C . Z iesenhenne.

Ch in im i : 1 965 ,G . Bakus . Igur in : 1956 , s tuden t collec

tor . Igu r in : 1 959, A . Sm i th and J . Coatswor th . Igur in :1 960 , E . S. Reese and R . A . Boo loo t ian . Medr en : 1 960 ,

E . S. Reese and R . A . Boo loo t ian . Medr en : 1 965 ,

G . Bakus . Muz in , Ki r in ian , and Bokonaar appu : 1 959 ,

F . C . Ziesenhenne. R ig i l i : 1 956 , s tuden t col lector . R igi li

1957, J . S. Gar th . R igi l i : 1 965 , G . Bakus . In land and on

beach ; on sandy bo ttom , outer r eef flat ; dr edged inlagoon , to m .

Coenobi to r ugosus H . M i lne Edwar dsAar aanbi r u : 1 959, F . C . Z iesenhenne. An iyaan i i : 1956 ,

s tuden t co l lector . Aomon ,B iji le, and Rojoa: 1 959, F . C .

Ziesenhenne. Igur in : 1 959, A . Sm i th and J . Coatswor th .

Igur in : 1 960 , E . S. Reese and R . A . Booloo t ian .

Medr en : 1959 , F . C. Z iesenhenne. Medren : 1960 , E . S.

Reese. Muz in , K i r in ian , and Bokonaarappu : 1959, F . C.

Z iesenhenne . R igi l i : 1 956 , s tuden t co l lector s . In land andon beach ; lagoon and seawar d s ides ; on sandy bot tomto m .

Fam i ly DIOGENIDAEAn icu lus an icu lus (Fabr ic ius )Aaraanbi r u : 1 959, F . C. Z iesenhenne. Enewetak : 1967,J . W . Knudsen . Between Enewetak and Bokandr etok :

1966 , J . W . Knudsen . Igur in : 1960 , E . S. Reese and

R . A . Boo loo t ian . On r eef flat and algal r idge.

An icu lus sp .

Off Bokandr etok : 1 974 , to 3 m on sandy bo ttom ,

S. L . B r u nenmei ster .

Calcin u s e legans (H . M i lne Edwar ds )An iyaan i i : 1 957 , J . S. Gar th . Bogen : 1957 , J . S. Gar th .Enewetak : 1 96 1 , E . S. Reese . Enewetak : 1 965 , 1 967,

J . W . Knudsen . Between Enewetak and Bokandr etok :

1 966 , J . W . Knudsen . Engebi : 1957 , J . S. Gar th .

Engebi : 1 959 , F . C . Z iesenhenne. Igu r in : 1957 , J . S.

Gar th . Medr en : 1 959, J . S. Gar th . Medr en : 1960 , E . S.

Reese. Medren : 1 974 or 1 975, S. L . Br u nenmeis ter .

R ig i l i : 1 957 , J . S. Gar th . Run i t : 1 959 , J . S.

Gar th . Enewetak Ato l l , no fur ther local i ty data : 1 946 ,

F . C. Z iesenhenne. Lagoon s ide of r eef; seawar d r eefflat and algal r idge. Under rocks and dead cor al , on

dead cor al heads , on l ive Acr opora and Poc i llopora.

Calcinus gaimardi i (H . M i lne Edwards )Enewetak : 1 968 , J . W . Knudsen . Igur in : 1 957 , J . S.

Gar th . Medr en : 1959, J . Rober ts and/or F . C. Ziesen

henne . R igi li : 1956 , s tuden t co llector . On r eef and on

sandy beach ; under and ar ound Por i tes co lon ies .

Calcin us guamens is Woos terEnewetak : 1 966 , 1 967 , J . W . Knudsen . BetweenEnewetak and Bokandretok : 1 966 , J . W . Knudsen . On

reef , fr om Po r i tes , F avia,l ive Acr opora, and live and

dead Poc i llopor a .

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252 GARTH , HAIG , AND KNUDSEN

Dardanus scu tellatus (H . M i lne Edwar ds )Aaman , B iji le, and Rojoa : 1959 , F . C. Ziesenhenne. Off

Bokandr etok : 1974 , S. L . B r u nenmeis ter . Chin ieero :

1 960 , E . S. Reese. Medren : 1 957 , 1 959, J . S. Gar th .

Medr en : 1 960 , E . S. Reese. Muti : 1956 , s tuden t collecto r . In lagoon ,

l i ttoral to 1 2 m , on sandy bottomsar ound and on cor al heads and among coral r ubble.

Dardanus woodmason i (A lcock)Aaman , B iji le, and Rojoa: 1959, to m on sandybottom , lagoon s ide, F . C . Ziesenhenne.

D iogenes gardiner i A lcockAomon , B iji le, and Rojoa: 1 959, F . C . Ziesenhenne.

Enewetak : 1 966 , 1967, J . W . Knudsen . Medr en : 1959 ,

J . S. Gar th . Between Pi i raai and Run i t : 1 967 , J . W .

Knudsen . On sandy bo ttom s in lagoon , to 24 m .

D iogenes pallescens Whi telegge

Enewetak Ato l l : 1 966 ,in lagoon 100 yds wes t of

mar ker , ran 250 yds due south , shel l— sponge bottomat 60 m ,

J . W . Knudsen . Enewetak Atol l : 1968 , in

lagoon at 1 1°26’N , dr edging near coral

mound , bo ttom of shel l and some algae at 60 m , J . W .

Knudsen .

Pagu r is tes sp .

Medren : 1 959, J . Coatswor th , J . S. Gar th . Rojoa : 1967,J . W . Knudsen . Enewetak Ato l l , 1 1 °26 ’N , 1 62

°1 7 .5’E

1 968 , J . W . Knudsen . Beh ind r eef on coral , m ; inlagoon , to 45 m .

Tr iz opagu r us s tr igotus (H er bs t)Enewetak : 1974 or 1 975 , in in ter is land channel atnor ther n end of is land , low t ide at about m depth ,S. L . Br unenmei s ter . Medr en : 1 959 , r eef between is landand L . C. T . wreck , inter t idal at +o.8-foo t t ide, J . Rober ts and/or F . C . Z iesenhenne.

Fam i l y PAGUR IDAECotopagu r us sp .

Enewetak Ato l l : 1 966 , dr edging in lagoon on spongeand sand bo ttom , depth not ind icated , J . W . Knudsen .

Enewetak Atol l : 1 968 , dredging in lagoon at 1 1°26’N ,

in 30 m depth on bo tt om of br oken cor aland shell , J . W . Knudsen .

Pagu r ix us anceps (Fores t)Bogallua: 1968 , J . W . Knudsen . Bokandr etok : 1 967 ,

J . W . Knudsen . Enewetak : 1975 , S. L . B r unenmeister .

Between Enewetak and Bokandretok : 1 966 ,

J . W . Knudsen . Igur in : 1959, J . S. Gar th . Medr en :

1957 , T . Gor eau and R . Nesh ida. Rojoa: 1967 ,

J . W . Knudsen . In ter t idal under rocks and to 15 ft ,lagoon s ide; more fr equently on reef flat at low water ,fr om Pori tes , Acropo r a, and Favia, some l ive.

Pagu r ix us bon inens is (Melin )Medr en : 1 957 , about 8 ft , from Poc i llopo r a and

Acropo ra, A . H . Banner .

Pagu r ix u s moor us (Nob i l i )An iyaan i i : 1966 , in mos tl y dead Acropo ra,

J . W . Knudsen . Between P i i r aai and Run i t : 1 967 ,

from 95 to 1 05

ft in lagoon , J . W . Knudsen . Rojoa: 1966 , nor thwes ts ide an outer reef channel , fr om dead Acropor a,

J . W .

Knudsen . Enewetak Ato l l : 1 968 , dr edging in lagoon at

1 1 °26'N ,1 00 ft , broken coral and shell

w i th much l ive coral , J . W . Knudsen .

New genus , sp .

Mut i : 1966 , col lected at pier from dead and heav i lyencr us ted poci llopor id cor al , J . W . Knudsen .

Fam i ly GALATH EIDAEA llogalathea e legan s (Adams and Wh i te)Near Jier or v , at about 1 1 °26 ’N , 1 62

°21 ’E: 1967, J . W .

Knudsen . Off Medr en : 1 968 , 1971 , J . W . Knudsen.

Enewetak Ato l l , 1 1 °29'N , 162°19'E: J . W. Knudsen.

Enewetak Ato l l : 1 976 or 1978, A . Fielding . In lagoon ,

to 9 m ; all specimens fr om cr ino ids .

Corall iogalathea hum i l is (Nobi li )Bogombogo : 1957 , D . Reish . Enewetak : 1967

J . W . Knudsen . Enewetak : 1967, S. Swerdloff . Rigi li1965,

J . W . Knudsen . Enewetak Atol l , 1 1°25’N ,

162°1 3 ’E: 1967, J . W . Knudsen . On reef fr om deadcorals and l ive Acr opor a; dr edged in lagoon in 45 m on

bo ttom of sponge, dead shell , coral , and algae.

Galathea aegyptiaca PaulsonAn iyaan i i : 1 960 , nor t h lagoon s ide, fr om coral head,E . S. Reese.

Galathea offin is Or tmannAn iyaan i i : 1957, A . H . Banner . An iyaani i : 1 966, 1 967

J . W . Knudsen . Between B i l lee and Bokonaarappu

1967 , J . W . Knudsen . Bogallua: 1968 , J . W . Knudsen.

Bogombogo: 1 957, D . Rei sh . Bokandretok: 1965, J. W .

Knudsen . Enewetak : 1965, 1967, 1968, J . W . Knudsen . Between Enewetak and Bokandr etok : 1966 , J . W .

Knudsen . Engebi : 1 967 , J . W . Knudsen . Igur in : 1 965,J . W . Knudsen . Between K i r in ian and Muzin : 1965,J . W . Knudsen . Medr en : 1 957, T . Goreau and

R . Nesh ida . Medr en : 1959, J . Coatswor th . Rigi li : 1965,

1968 , J . W . Knudsen . Rojoa: 1966 , 1967, J . W . Knudsen . On reef to m in l ive and dead corals , includingAcr opora and Por ites ; dredged in lagoon at 30 m .

Calathea ambo inens is De Man

Enewetak Ato l l : 1976 or 1 978 , fr om cr inoids , A . Fieldi ng .

Galathea sp . , aff . tonegashimae BabaEnewetak : 1967 , S. Swer dloff . Jieror v : 1967, 1968,J . W . Knudsen . Medr en : 1957 , A . H . Banner . Rojoa1 966 , J . W . Knudsen . On reef and in lagoon , to 9 mdepth ; on corals including live and dead Acropora, and

fr om cr ino ids .

Phylladior hynchu s ser r i ros t r is (Mel in )Enewetak : 1967, fr om coral head in lagoon about 300yds . fr om labo rator y s i te, J . W . Knudsen . Medren :

1 957 , d iv ing in 3 to m , coral heads and dead coral ,T . Gar eau and R . Nesh ida.

Fam i ly PORCELLAN IDAEN eopetr ol is thes maculatus (H . M i lne Edwar ds )Bokandretok : 1 968 , J . W . Knudsen . Medren : 1971

J . W . Knudsen . Between Medren and Enewetak : 1968 ,

J . W . Knudsen . R ig i li :_

1968 , J . W . Knudsen . To 3 mdepth in channels and lagoon , from anemones .

Pachycheles johnson i H aigB iji le: 1 968 , A . H avens . Bokandretok : 1 967, J . W .

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA)

Knudsen . Bokandr etok : 1 968 , A . H avens . Enewetak :1 965 to 1 967 , J . W . Knudsen . Enewetak : 1 968 , A . H a

vens . Enewetak : 1969 , C. A . Ch i ld . Between Enewetakand Bokandretok : 1966 , J . W . Knudsen . Engebi : 1 959 ,

F . C . Z iesenhenne. Engebi : 1 967 , J . W . Knudsen . Mu i :

1 967 , J . W . Knudsen . Muti : 1 966 , 1 968 , J . W . Knudsen . R ig i l i : 1957 , A . H . Banner . R ig i l i : 1966 , J . W .

Knudsen . On reef , in ter t idal or shal low subt idal , fr equently in or under l ive and dead Por ites , Poci llopor a,and Acr opor a; occas ional l y under r ocks .

Pachycheles p iso ides (H el ler )Enewetak : 1969, surge channel and blow ho le about 20ft back from outer r eef edge, C. A . Ch i ld . BetweenEnewetak and Bokandr etok : 1966 , on reef ins ide algalr idge, from dead encr us ted algae or fr om Poc i lIOporo,

J . W . Knudsen .

Pachycheles sp in ipes (A . M i lne Edwar ds )Chin ieero : 1 965 , J . W . Knudsen . Enewetak : 1 965 ,

1 967 , J . W . Knudsen . Between Enewetak and Bokandr etok : 1 966 , J . W . Knudsen . On r eef, from corals .

Petrol is thes as iat icus (Leach)Bogen : 1 957, J . S. Gar th . Bokandretok : 1 967 ,

J . W .

Knudsen . Bokandr etok : 1968 , A . H avens . Enewetak :1 965, J . W . Knudsen . Enewetak : 1 968 , A . H avens .

Enewetak or Gugegwe [Kwajalein Atol l]: 1 946 ,F . C.

Z iesenhenne. Engebi : 1 957 , J . S. Gar th . Igur in : 1 959,

J . S. Gar th . Jier u r o : 1 946 , M . W . Johnson . Medr en :

1957, J . S. Gar th . Medr en : 1 965 , J . W . Knudsen . Muti :1 957, J . S. Gar th . Muti : 1 965 , G . Bakus . Mut i : 1 967

,

1968 , J . W . Knudsen . R igi li : 1 956 , s tudent co l lector :R ig i li : 1 959 , J . S. Gar th . Run i t : 1 959, J . S. Gar th .

Run i t : 1 965, J . W . Knudsen . Run i t : 1968 , A . H avens .

On reef , usual ly under r ocks and cor al r ubble .

Petrolis thes bisp inos us Bor r adai le

Bokandretok : 1 967, J . W . Knudsen . Enewetak : 1 965 ,

1967 , J . W . Knudsen . Enewetak : 1 969, C . E . Dawson ,

C. A . Ch i ld . Between Enewetak and Bokandretok :

1 966 , J . W . Knudsen . Mu i : 1 967, T . Smi th . Mut i1 968 , J . W . Knudsen . On seawar d r eef flat and algalr idge, in corals including Poc iIIOpor a.

Petr ol is thes bo r radai lei Kr oppAn iyaan i i : 1 956 , s tudent col lector . Bogen : 1957, J . SGar th . Bokandretok : 1 967 , J . W . Knudsen . Enewetak :1 965 , 1 967, J . W . Knudsen . Engebi : 1 957 , J . S. Gar thIgur in : 1 959, J . S. Gar th . Mut i : 1 965 , 1 968 , J . WKnudsen . Muti : 1 965 , G . Bakus . Muti : 1 968 , A . H a

vens . R ig i l i : 1 957, J . S. Gar t h . Run i t : 1 959, J . SGar th . On r eef , usual ly under r ocks and in coral r ubble.

Pet r o lis thes coccineus (Owen )Enewetak : 1 969 , su rge channel and blow ho le about 20i t back fr om outer r eef edge, C . A . Ch i ld . Engebi : 1 967 ,

algal r i dge, J . W . Knudsen .

Petro l is thes decocan thus Or tmannEnewetak : 1 967 , algal r idge, J . W . Knudsen . Enewetak :1 969, outer r eef r im and surge channels along nor thone-thi r d of i s land , C . E . Dawson and C . A . Ch i ld .Enewetak : 1 969 , sur ge channel and blow ho le about20 ft back from outer r eef edge, C . A . Ch i ld .

253

Petr o lis thes e legan s H aigEnewetak : 1 965, 1 967 , J . W . Knudsen . Enewetak :1 969 , C. A . Ch i ld . Between Enewetak and Bokandre

tok : 1 966 , J . W . Knudsen . Medr en : 1 965 , J . W . Knudsen . Mut i : 1 968 , J . W . Knudsen . Rujor u : 1 946 ,

M . W . Johnson . Reef flat and algal r idge,under r ocks

and fr om cor als , includingPoc i llopora.

Petr ol is thes f imbr iatus Bor r adai leAn iyaan i i : 1 957, J . S. Gar th . An iyaan i i : 1 967 , J . W .

Knudsen . Bogen : 1 966 , J . W . Knudsen . Bokandr etok :

1 967 , J . W . Knudsen . Bokandretok : 1 968 , A . H avens .

Enewetak : 1 957 , J . S. Gar th . Enewetak : 1 965 to 1 968 ,

J . W . Knudsen . Enewetak : 1 968 , A . H avens . Enewetako r G ugegwe [Kwajalein Ato l l]: 1 946 ,

F . C . Ziesenhenne.

Between Enewetak and Bokandretok : 1 966 , J . W .

Knudsen . Engebi : 1 956 , s tudent co l lector . Engebi

1 957 , J . S. Gar th . Igur in : 1 957 , 1 959 , J . S. Gar th . Igu

r in : 1 965 , J . W . Knudsen . Jei roju : 1946 , J . P . E . Mor r ison . Jier u r o : 1 946 , M . W . Johnson . Medr en : 1 957,

1 959 , J . S . Gar th . Medren : 1 965, J . W . Knudsen . Muti :1 957 , J . S. Gar th . Muti : 1 965, 1 968 , J . W . Knudsen .

Mut i : 1 965 , G . Bakus . Mut i : 1968 , A . H avens . Pi i r aai :

1 968 , A . H avens . R ig i l i : 1 956 , D . Reish . R igi l i : 1 959,

J . S. Gar th . R ig i l i : 1 966 , 1 968 , J . W . Knudsen . Run i t :

1 959 , J . S. Gar th . Enewetak Atol l , no fur ther locali tydata: 1944 , W . A . Bar tos . Enewetak Atol l , no fur therlocal i ty data : 1 946 , F . C. Ziesenhenne. On reef , underrocks and fr om r ubble; occas ional ly from corals .

Pet r olis thes lamar cki i (Leach)An iyaan i i : 1 957 , J . S. Gar th . Bokandr etok : 1 967 ,

J . W .

Knudsen . Bokandr etok : 1968 , A . H avens . Enewetak1 965, 1 967 , 1 968 , J . W . Knudsen . Enewetak : 1 968 ,

A . H avens . Engebi : 1 957 , J . S. Gar th . Igur in : 1 957 ,

1 959, J . S. Gar th . Medr en : 1 959 , J . S. Gar th . Medr en :1 965, J . W . Knudsen . Mu i : 1 968 , A . H avens . Mut i :1 957, J . S. Gar th . Muti : 1 965 , G . Bakus . Mut i : 1967 ,

1 968 , J . W . Knudsen . Mut i : 1 968 , A . H avens . Pi i raai :

1 968 , A . H avens . R ig i li : 1 957 , 1 959, J . S. Gar th .R ig i li : 1 966 , J . W . Knudsen . Run i t : 1 959 , J . S. Gar th .

Run i t : 1 965 , J . W . Knudsen . Enewetak Ato l l , no fur therlocal i ty data: 1 946 , F . C. Ziesenhenne . On r eef, usual lyunder r ocks and in cor al r ubble .

Petr ol is thes mosoki i M i yakeEnewetak : 1 969 , surge channel and blow hole about20 ft back fr om outer reef edge,

C . A . Ch i ld .

Pet r olis thes pen ici l latus (H el ler )Off Bokandr etok : 1 968 , A . H avens . Enewetak : 1 968 ,

J . W . Knudsen . Enewetak : 1 968 , A . H avens . Enewetak :1 969 , C . A . Ch i ld . Between Enewetak and Bokandre

tok : 1 966 , J . W . Knudsen . Medr en : 1 965 , 1 968 , J . W .

Knudsen . R ig i l i : 1965 , J . W . Knudsen . On r eef , to

m depth , under rocks and r ubble, and in Por ites ,Acropora, and Poc i llopor a.

Pet r o l is thes pubescens St impsonBokandr etok : 1 968 , channel southwes t of is land , underr ock , A . H avens .

Pet r o lis thes [undescr ibed sp . 1]R ig i l i : 1965 , no r th end of i s land fr om l ive Poc illopor aelegans , J . W . Knudsen .

254 GARTH . HAIG , AND KNUDSEN

Petr ol is thes [undescr i bed sp . 3] 1 967 , J . W . Knudsen . Reef flat and r i dge, some speciEnewetak : 1 967 , J . W . Knudsen . Enewetak : 1 968 , mens fr om co r als .

A . H avens . Enewetak : 1 969 , C . A . Ch i ld . Engebi :

TABLE 1

Data on Col lect i ons of Anom u r a‘

Col lect or (s ) Iden t i fi er Depos i tory

Bar tos , W . A . USNM

Br unenmeis ter , S. L .

USNM , RMNH

Coatswor th , J .Co l in , P . , and D . M . Devaney

Dawson , C. E . USNM , RMNH

Donaldson , L ., and E . H eld

Fielding , A .

Fusar o , C .

Gar th , J . S.

Gor eau , T . , and R . Nesh ida AH F , MPRL

MacCoy, C. V .

Mor r i son , J . P . E .

Reese, E . 5 .

Rees e, E . S., and

R . A . Boo loot ian

Bei sh , D .

Rober ts , J .Sather , B . , and R . StevensonSchu ltz , L . P .

Sm ith , A . , and J . Coatswor thSm i th , T .

Student co l lector sSwerdloff , S.

Ziesenhenne, F . C .

‘AH F , A l lan H ancock Foundation ; BPBM , Ber n ice P . B ishop Museum ; MPRL , M id-Pacifi c

Resear ch Labo rator y , Enewetak ; RMN H , R i jksmuseum van Natu u r l ijke H istor ie, Leiden ; USNM ,

Nat ional Museum of Natur al H i story , Sm ithson ian Insti tution .

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256 GARTH , HAIG , AND KNUDSEN

Daldorf io (or Par thenope) sp .

Knudsen .

H or r ovio e legans De ManGar th .

?H eterocryp ta sp .

Knudsen , H avens .

Par thenope (Au locolombr us) cu r visp in is (M ier s)Knudsen .

Par thenope (Au loco lombr us) hop lonotus (Adams andWh i te)Knudsen .

Par thenope (Au loco lombr us) sp .

Knudsen , H avens .

Par thenope (Pseudolambr u s) sp .

Knudsen .

Par thenope sp .

Knudsen .

Thyro lombr us er osus (M ier s )Knudsen .

Thyr o lombr us sp .

Knudsen .

Fam i l y ATELECYCL IDAEKr ous s ia in teg r o RathbunKnudsen , H avens .

Kr ous s ia sp . , cf . mar quesa SereneKnudsen .

Krous s ia n itida St impsonGar th .

Kr ous s ia r os tr ipes F . MullerH avens .

Krous s ia r ugu losa (Kr aus s )EMBL , Gar th , Knudsen ,

'

H avens , Ch i ld .

Fam i l y PORTUN IDAECor upa tenu ipes DanaZiesenhenne, Knudsen , H avens .

Cotop t r us inoequol is (Rathbun )Knudsen .

Cotop t r us n itidus A . M i lne Edwar dsGar th , Z iesenhenne.

Cotop t r us r athbunoe Serene

Ziesenhenne.

Cotop tr us ?tr uncat ifr on s De ManH avens .

Char ybd is (G on iosup r adens) erythrodoctylus (Lamar ck)H aven s , F ield ing .

Coelocar cinu s fol iotus EdmondsonH avens .

L ibis tes vi llos us RathbunKnudsen , H avens .

L is socar cinu s holo thu r icola Str eetsKnudsen .

L is socar cin us or bicular is DanaGar th , Shoup , Knudsen , H avens .

Por tunus (Ache lous) gr anu laras (H . M i lne Edwar ds )Z iesenhenne , Knudsen , H avens .

Por tunus (Ache lous) sp . , nr . or bicular is (R ichter s )Gar th , Z iesenhenne.

Po r t unus (H ellenu s) long isp inos us Stephenson and

Campbel l .

Reish, Gar th , Ziesenhenne,Knudsen , H avens .

Thalam i ta admete (H er bs t)EMBL , Reish, Gar th , Ziesenhenne,

Knudsen, H avens ,Ch i ld .

Thalam ita bouv ier i Nobi l iKnudsen .

Thalam i ta chap tol i i (Audouin)Knudsen .

Thalam ita coer u leipes Jacqu inotKnudsen , H avens .

Thalam i ta cor r ugata Stephenson and Rees

Knudsen .

Thalam ita dakin i Mon tgomer yKnudsen , H avens , Ch i ld .

Thalam ita g lor iens is Crosn ier

Mor r ison .

Thalam ita graci l ipes (A . Mi lne Edwar ds)Gar th , Ziesenhenne, Knudsen , H avens .

Thalam ita in teg r o Dana

Pomeroy and Kuenz ler .

Thalam ita ocu lea A lcockKnudsen .

Thalam ita p icta StimpsonEMEL , Gar th , Ziesenhenne, Knudsen , H avens , Ch i ld.

Thalam i ta p i lumnoides Bor r adai le

Gar t h , Knudsen .

Thalam ita p r ymno (H er bs t)Knudsen , H avens , Ch i ld , Field ing .

Thalam i ta quodr i lobata M ier sKnudsen .

Thalam ita sex loboto M ier sKnudsen .

Thalam ita s imo H . M i lne EdwardsKnudsen .

Thalam ita sp icer i EdmondsonGar th , Knudsen , H avens , H ighsmi th .

Thalam ita sp in imana Dana

EMEL , Gar th .Thalam ita s t impson i A . M i lne Edwar dsKnudsen .

Th alam ita wokens is EdmondsonKnudsen , H avens .

Thalam i ta yo ronens is SakaiKnudsen .

Thalam i ta sp . , nr . ououens is RathbunKnudsen .

Thalom ito ides quadr idens A . M i lne Edwar dsBanner , Gar th , Reese, Knudsen , USGS (foss i l) .

Fam i ly XANTH IDAEActaea sp . , nr . bocki OdhnerKnudsen .

cavipes Dana

Knudsen , H avens .

Actaea morgar it ifer o OdhnerKnudsen .

Actaea pu lche l la modes ta (De Man )Knudsen .

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA)

Actaea quodr iar eoloto Takeda and MiyakeBanner , Gar th , Knudsen .

Actaea sp .

Knudsen , H avens .

Actoeodes consobr inus (A . M i lne Edwards )Gar th , Reese, Knudsen , H avens .

Actaeodes hi r s ut is s imus (R i ippell)USGS (foss i l) .

Actumnus on telmei WardKnudsen .

Actumnus asper (Ruppell)Knudsen .

Actumnus setifer (De H aan)Knudsen .

Actumnus sp .

Knudsen .

Actumnu s (or P i lumnus ) sp .

Knudsen .

Atergot is ?di l itatus De H aan

de G r uy.

Atergot is flor idus (L innaeus)Knudsen .

A tergatop s is s ignata (Adams and Wh i te)Knudsen , Bur ke.

Banareio nobi l i i (Odhner )Knudsen .

Banar eio par vula (Krauss )Knudsen .

Corp i lius convexus (For sskal)Johnson , Ziesenhenne, Knudsen , H avens , Field ing .

Corp ilius maculatus (L innaeus)Knudsen , F ielding .

Chlorodiello corallicole Miyake and TakedaKnudsen .

Chlorodiello cytherea (Dana)Ziesenhenne, Knudsen , H avens .

Chlorodiello laeviss ima (Dana)Reish, Gar t h , Z iesenhenne, Reese, Knudsen , H avens .

Chlorodiello n igra (For sskal)Reish, Banner , Gar th , Shoup , Knudsen , H avens .

Cycloxan thop s cavatus RathbunGar th , Knudsen , H avens .

Cyma andreossyi (Audou in)Banner , Reese, Knudsen .

Cyma dep lonotus A . M i lne Edwar dsZiesenhenne, Reese, Shoup , Knudsen , Ch i ld .

Cymo melanodactylus De H aan

Gar th , Reese, Knudsen .

Cyma quodr i lobatus M ier sKnudsen .

Dacryop i lumnus erem ita Nobi l iKnudsen , H avens .

Dacryop i lumnus r athbunoe Bals s

Knudsen , H avens .

Dai r a per lata (H er bs t)H avens .

Domecia g labra AlcockBanner , Gar th , Ziesenhenne, Reese, Shoup , Knudsen ,

H avens , Ch i ld .

257

Domecia hisp ida Eydoux and Sou leyet

Reese, Shoup , Knudsen , H avens .

E r iphia scabr icu lo Dana

Mor r ison , EMBL , Gar th , Ziesenhenne, Knudsen ,

H avens , Ch i ld .

E r iphia sebana (Shaw and Nodder )Mor r ison , EMBL , Gar th , Ziesenhenne, Bakus , Knudsen , H avens , Ch i ld .

Et is us bifr on tal is (Edmondson)Knudsen .

Et is us deman i OdhnerJohnson , Knudsen , Gar th , H avens .

Etis us sp . , nr . deman i OdhnerKnudsen , H avens .

E t isu s den tatus (H er bs t)Gar th , Z iesenhenne, Knudsen , H avens , Ch i ld .

E t is us electr a (H er bs t)Reish, Knudsen .

E t isus fron talis DanaKnudsen .

E t is us laeviman us Randal lUSGS (foss i l) .

Et is us molokoiens is (Rathbun)Gar th , H avens .

Et is us sp lendidus RathbunF ield ing , USGS (foss i l) .

Et is us sp . 1

Knudsen , H avens .

Et isus sp . 2

Knudsen , H avens .

Et is us sp . 3

H avens .

Euxan thus exscu lp tus (H er bs t)Gar th .

Euxan thus (or H ypocolpus ) sp .

Knudsen .

Goi llordiellus r ueppell i i (Kraus s)Z iesenhenne, Knudsen .

Goi llordiellus s uper ci l iar is (Odhner )Rei sh, Banner , Gar th , Ziesenhenne, Reese, Shoup,Knudsen , H avens , Ch i ld .

G lobop i lum n us g lobosus (Dana)Knudsen , H avens .

H eter op i lum n us sp . , cf . long ipes (St impson)Knudsen , H avens .

Lachnopodus ponopens is (Rathbun)Knudsen , H avens .

Lachnopodus s ubocu tus (St impson)Knudsen , H avens , Field ing .

Lachnopodus tohi tens is De Man

EMEL , Gar th , Knudsen , H avens , Chi ld , Bur ke.

Lep tod ius dovaoens is War dH aven s , Ch i ld .

Lep todius exar otus (H . M i lne Edwar ds)Reish, Gar th , Buss ing ,

Knudsen , Chi ld .Lep tod ius gr aci l is (Dana)Ziesenhenne, Bakus , Knudsen , H avens .

Lep tod iu s n ud ipes (Dana)Knudsen , H avens .

258 GARTH , H AIG , AND KNUDSEN

Lep tod ius sangu ineus (H . M i lne Edwar ds )Z iesenhenne, Mor r ison , Johnson , EMBL , Gar th ,Bakus , Knudsen , H avens , Ch i ld .

Lep tod ius woioluonus RathbunKnudsen , H avens , Ch i ld .

L iocorp i lodes arm iger pocif icus BalssLadd , Banner , Gar th , Reese, Shoup , Knudsen ,

H avens , H ighsm i th .

L iocorp i lodes biungu is (Rathbun)Ladd , EMBL , Reish, Gar th , Knudsen , H avens , Ch i ld ,H ighsm i th .

L iocorp i lodes in teger r im us (Dana)Reish , Knudsen , H avens .

L iocorp i lodes pum i lus (Jacqu inot)Ladd , EMEL , Knudsen .

L iomer a be l la (Dana)Bar tos , Mor r ison , Banner , Gar th , Ziesenhenne,

Shoup , Bakus , Knudsen , H avens , H ighsm i th .L iomera coe lata (Odhner )Banner .

L iomera loev is (A . M i lne Edwar ds)Gar th .

L iomera mon t icu losa (A . M i lne Edwar ds )Reese, Knudsen .

L iomera pa l l ida (Bor radai le)Knudsen , H avens .

L iomera r ugoto (H . M i lne Edwar ds)Ziesenhenne, Gar th , Knudsen , H avens .

L iomer a s t imp son i (A . M i lne Edwar ds )Knudsen .

L iomer a tr is t is (Dana)Gar th .

L iomera sp .

Knudsen .

Lophozozym us dodone (H er bs t)Knudsen , Ch i ld .

Lophozozymus incis us (H . M i lne Edwar ds )Johnson .

Lophozozym u s pu lchellus A . M i lne Edwar dsShoup , Knudsen , H avens .

Lybia caes t ifera (A lcock)Knudsen .

Lybia tes se l lato (Latr ei l le)Z iesenhenne, Stokes , Knudsen , H avens .

L ydia onn u l ipes (H . M i lne Edwar ds )Knudsen , H avens , Field ing .

Macr omedaeus n udipes (A . M i lne Edwar ds )Gar th , Knudsen , H aven s , Ch i ld .

Maldivia palmyrens is RathbunKnudsen , H ighsm i th .

Mold ivio t r iu ngu icu loto (Bor r adai le)Knudsen , H avens , H ighsm i th .

Medaeus e legan s A . M i lne Edwar dsKnudsen , H avens .

Medaeus o r natus DanaKnudsen , H avens .

Neoxan thios imp ressus (Lamar ck )Gar th , Z iesenhenne, Knudsen , Bur ke.

Par actoeo retusa (Nobi l i )Knudsen , H avens .

Par actoeo r ufopunctota (H . Mi lne Edwar ds )Banner , Gar th , Knudsen , H avens .

Paractoeo r ufop unctoto f . p lumoso Gu inotKnudsen .

Paractoeo tumu loso (Odhner )Knudsen .

Par omedoeus s imp lex (A . M i lne Edwar ds )Knudsen , H avens , Ch i ld .

Por op i lum n us cor oll iophi lus Takeda and M iyakeShoup , Knudsen , H avens .

Porop ilumnus ?incer tus Takeda and M i yakeKnudsen .

Par axon th ias natatas (Dana)Ladd , Reish , EMBL , Banner , Gar th , Ziesenhenne,

Knudsen , H avens , H ighsmi th .Par oxan th ias pachydoctylus (A . M i lne Edwar ds )Knudsen , H avens .

Phymodius ?gr anu latus (Targ ion i-Tozzett i )

Knudsen .

Phymod ius layson i RathbunReese, Knudsen , H avens .

Phymod iu s mon t icu los u s (Dana)Gar th , Knudsen .

Phymodius n itidus (Dana)Gar th , Knudsen , H avens .

Phymodius ungu lotu s (H . M i lne Edwar ds )Banner , Reish, Gar th , Reese, Shoup , Knudsen ,

H avens .

P i lodius ar eolatus (H . M i lne Edwar ds )Ziesenhenne, Johnson , Ladd , EMBL , Gar th ,Z iesenhenne, Shoup , Bakus , Knudsen , H avens ,

H ighsm i th .P i lodius flavus RathbunReese, Knudsen , H avens .

P i lodius melanodactylus (A . Mi lne Edwar ds )Knudsen .

P i lod ius p i lumno ides (Wh i te)Mor r i son , Rei sh, Gar th , Knudsen ,

H avens .

P i lod ius pug i l DanaJohnson , Reish, Banner , Z iesenhenne , USGS (foss i l) .

P i lod iu s s cabr icu lu s Dana

Knudsen , H avens .

P i lod ius sp in ipes H ellerReish, Banner , Gar th , Knudsen , H aven s .

P i lum nus ander son i De Man

Knudsen , H avens .

P i lumnus caer u lescens A . M i lne Edwar dsKnudsen , H avens , Ch i ld .

P i lum nus ?e legans De ManKnudsen .

P i lum nus long icom is H i lgendor fBanner , Gar th , Knudsen , H avens , Ch i ld .

P ilum nus r anson i For es t and Gu inotKnudsen .

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260

Cyclogr ap s us long ipes St impsonKnudsen , H aven s .

Cyclogr ap s us sanctoecr ucis Gr i ffinKnudsen .

Geogr ap s us cr in ipes (Dana)H eld , Knudsen , H avens , F ield ing .

Geogr ap sus gr ayi (H . M i lne Edwar ds)H eld , Knudsen , H avens , Field ing .

G rapsus in termedius De ManKnudsen .

Grapsus long i tor sus DanaEMEL , H eld , Ch i ld , H avens .

G r aps us tenu icr us tatus (H er bs t)EMBL , Gar th , Ziesenhenne, Reese and Booloot ian ,

Knudsen , H avens .

Metasesarma r ous seoux i H . M i lne Edwar dsKnudsen .

Metopogr ap s us thukuhar (Owen)Knudsen , H avens .

Pachygr aps u s m inutas A . M i lne Edwar dsEMBL , Reish, Gar th , Ziesenhenne, Bakus , Knudsen ,

H avens , Ch i ld , H ighsm i th .

Pochygr ap sus p lon ifr ons De ManGar th , Bakus , Knudsen , H avens , Ch i ld .

Pochygr op s us p l icatus H . Mi lne EdwardsEMBL , Bakus , Knudsen , H avens , Ch i ld .

Per cnon abbr eviatum (Dana)Knudsen , H avens , Ch i ld .

Per cnon p i l imonus (A . M i lne Edwar ds )Knudsen , H avens , Ch i ld , Field ing .

Per cnon p lan is s imum (H er bs t)EMBL , Bakus , Knudsen , H avens , Ch i ld , H ighsm i th .

P logus io dep r essa tuber culata Lamar ckKnudsen , H avens .

P logu s io immacu lata Lamar ckF ield ing .

P logu s io speciosa DanaKnudsen , H avens , Ch i ld , F ielding .

Pseudogr apsus albus St impsonEMBL , Knudsen , H avens .

GARTH , HAIG , AND KNUDSEN

Sesarma (H olometOp us) sp .

Knudsen .

Fam i ly GECARClNlDAEGecarco ideo lalandi i H . M i lne Edwar dsKnudsen , Field ing .

Fam i ly PiNNOTH ERlDAEXan thas ia mur igero Wh i teGar th .

Fami ly OCYPODlDAEMacr Ophthalm us (Macr ophthalmus) telescop icus (Owen)Knudsen .

Macr Ophthalmus (Mopsocar c inus) bo sci Audou in and

Sav ignyGar th , Knudsen .

Ocypode cer atophthalmo (Pal las )EMBL , Gar th , Ziesenhenne, Reese and Booloot ian ,

Bakus , Ch i ld .

Ocypode cordimana Desmar es t

Knudsen , Field ing .

Par acleis tosomo (or Cleis tosomo) sp .

Knudsen .

Uca tetragonon (H er bs t)Knudsen .

Genus and spec ies incer tae sedis

Knudsen .

Fam i l y H APALOCARCINIDAECryp tochir us corolliodytes H el lerBakus , Knudsen , W i jsman .

H apalocarcinus mar s up iolis StimpsonReese, Shoup, Knudsen , H avens .

Neotr og locarcinus dowydoff i (Fize and Serene)Reese and Stevenson , BPBM .

Pseudocryp tochi r us cr escen tus EdmondsonPichon .

Species 1 , incer tae sedis

Knudsen .

Species 2, incer tae sedis

Knudsen .

TABLE 2

Data on Co l lect i on s of B rachyu r a‘

Collector (s )

Bakus , G . J. , and B . H . Buss ingBanner , A . H .

Bar tos , W . A .

Burke, R .

Ch i ld , C . A .

Coatswor th , J .

Iden t i fi er Depos i tory

‘AH F , Allan H ancock Foundation ; BPBM , Bern ice P . B i shop Museum ; MPRL , Mid

Paci fi c Research La bor atory , Enewetak ; USNM , National Museum of Natu ral H istory ,Sm i thson ian Ins ti tution .

TXan thidae on ly ; the Por tun idae were ident ified by Stephenson and Rees

(Th is table continued on next page. )

CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA)

TABLE 2 (con t’d)

Col lector (s ) Iden ti fier Depos i tor y

Gareau , T . , and R . Nesh idaH avens , A . D .

Ladd , H . S.

Mor r ison , J . P . E .

Palumbo , R .

Pichon , M .

Reese, E. S. , and R . A . Booloo t ian

Reese, E. S. , and R . A . StevensonReish , D .

Ryan , E .

Shoup , J .Stokes , D .

W i jsman , M .

Ziesenhenne, F . C.

‘

I'Xanthidae on ly ; the Por tun idae wer e ident i fied by Stephenson and Rees

26 1

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264 CUTRESS AND ROWE

TABLE 1 (con t ’d)Fam ily STICHOPODIDAEStichopus chlor onotu s Br andt : Lawr ence , 1 980 .

‘Stichopus har r ens Selenka, 1 867.

‘Stichopus vari egatus Semper , 1 868 .

”(St ichop us ?var iegatus juven i le.

Thelenota ananas (Jaeger ) : H umes , 1 973 ; Lamber son , 1 978 .

Thelenota anax H . L . Clar k : Rowe and Do ty , 1977 ; Lamber son , 1 978 .

Order APOPIDAFam i ly CH IRIDOTIDAEChi r idoto r igi da Semper : Lawr ence, 1980 ; Lawr ence and Gu i lle, 1 982 .

Fam i ly SYNAP’I'IDAE

‘Euop to godefir oyi (Semper ), 1 868 .

‘Pendekop lectono n igra (Semper ), 1 868 .

Synapta macu lata (Cham isso and Eysenhar dt) : Bakus , 1 968 , 1973 .

‘New publi shed Enewetak record .TVery small specimen .

maur itiana, and St ichopus hor r ens at Enewetak . Bakus( 1 968 , 1 973 ) r epor ted on defense mechan isms and the

eco logy of several species . Enewetak was one s i te for a

s tudy (Bakus and Green , 1974) on changes in toxi ci ty of

holothur ians wi th relat ion to var ious lati tu des , water temperatu r es , and habi tats . Pr esence of a tumor in one of 1 0

specimens of - H . (Mer tens iothur io) leucosp i loto was

r epor ted by Sm i th et ai . H umes ( 1973) recor ded aparas i t ic copepod on Thelenota ananas . In thei r r epor t onGuam ho lothur ians , Rowe and Do ty ( 1977) includedEnewetak in the general dis tr ibution of T . anax . Someaspects of the biology of H . (H .) atr a wer e s tud ied byWebb et ai . ( 1977) and Eber t Lawr ence ( 1 980)r ecor ded num ber s and biomass of eight species of

ho lothur ians on the Enewetak r eef flat . The organ ic compos i tion of the body wal l of s ix species of Enewetakholothur ians was analyzed by Lawrence and Gu i lle

B IOGEOGRAPH YAccor d ing to dis tr ibut ion summar ies for var ious Indo

Pac i fic species (H . L . Clar k , 1921 , 1 946 ; A . M . Clar k andRowe, 1 8 of the 20 holothur ian species known at

present fr om Enewetak Atol l are w idely d is tr i butedthroughout the Indo Wes t Paci fi c . The other two ,

Thelenota anax and Chi r idoto r ig ida, range fr om nor thernAus tral ia nor thwar d to the Ph i l ippines and eas twar d to theWes t and Cen tr al Paci fic . “

Of the 1 8 Indo Wes t Paci fic species , those mos t w idel ydi s tr ibuted are H o lothur ia (P lotyperona) diffici lis (Red Seato the H awai ian and Eas ter is lands) , H . (Thym iosycio) arenico lo (Red Sea to the H awai ian ,

Galapagos , and CocosIs lands ; Car ibbean ; Bermuda) and H . (T .) impat iens (RedSea to H awai i , Car ibbean , Med i ter r anean) . Twelve otherspecies r ange fr om the shor es of Eas t Afr ica or the Red

Juven i le specimens of thr ee spec ies (Act inopyga ?mour i t iona,

Labidodemos ?semper ianum , and St ichop us

?var iegatus ) have been col lected at Enewetak , and figur esof thei r sp icules have been included . Due to the elemen t ofdoubt r egar d ing thei r identi fi cat ion and the di s t inctivenessof thei r co lor and sp icules , all three forms have beenincluded in the key that fo l lows .

Sea to the Cen tral Paci fic : Act inopyga maur itiana,H . (H olodeima) atr a,

H . (T .) h i l la, H . (Mer tens iothur ia) per

vicox , H . (M .) leucosp i loto, St ichopus chlor onotus ,

S . har r ens, and Euop to godeffr oyi to the H awai ian Is lands ;

St ichop us var iegatus , Th elenota ananas , and Synop tu lo

macu lata to the L ine and (or ) Society is lands ; and

Pendap lectana n igr a to the Mar shall Is lands . The otherthree have not been r epor ted fr om A fr ica or the Red Sea .

Afr ocucum is afr icanus ranges fr om Maur i tius to Fi j i ;Bohadschia argus , fr om the Seychel les is lands to the

Mar shal l Is lands , and Labidodemos semper ionum , from theMald ives to the H awai ian and Society is lands .

Of the 78 ho lothur ian species recor ded fr om Paci fi ci s lands (A . M . Clar k and Rowe, on ly 20 ar e

represented at Enewetak . On the other hand , a vas t majori ty (9096) of the Enewetak species also ranges into the

Indian Ocean , compar ed w i th 68% of the total Paci fi cspecies r ecor ded by Clar k and Rowe. Wi th the poss i bleexcept ion of the sur pr i s ingly low number of spec ies found ,the compos i t ion of the ho lothur ian fauna at Enewetakcould be pr edi cted fr om the geog raph ical pos i t ion of thatatol l and may be ind icative of the reproduct ive s tr ategiesof the w idespr ead spec ies . Unfor tunately , l i ttle is known of

lar val developmen t of the species or length of lar val l i fe.

Inc idental l y , i t may be noted that Osh ima ( 1 91 6) repor tedbrood-car e in Af rocucum is af r icona in Japan , which is

HOLOTHURIANS 265

KEY TO SPECIES

1 . Body sausage-

shaped ; podia present , as pedicels (tube feet) and pap i llae; body wal l somet imes th ickand muscular , not s ticky to touch ; spicules tables , rods or plates , never anchor s , anchor plates , orwheelsBody worm- l ike; pod ia absent ; body wal l thin , usual ly s t icky to touch ; spicules anchor s , anchor p lates ,and wheelsTen tacles dendr i t ic (r ichly branched) ; ped icels in double r ows along each ambulacr um ; spicules in bodywal l lar ge knobbed , lent icular , per forated plates and smooth , per forated plates and rods (F ig . 3a);co lor un i formly deep v io let , almos t black in l i fe, gray-brown in alcohol ; body length up to 7 cm (F ig .

6a) Af r ocucum is af r icana (Semper )Tentacles peltate or peltato dig i tate (leaf-shaped); pod ia usually clear ly di fferentiated in to ven tral pedicels and dor sal papi l laeGonads in two tufts , one oh ei ther s ide of dor sal mesentery ; body s tout , flattened ven tral ly in to dist inct sole, squar ish in cr oss sect ion or ar ched dor sal ly , wi th large dor sal and lateral papi llae; amongspicules ei ther C-

shaped rods or dichotomous ly branched rodsGonads in s ingle tuft to left of dor sal mesenter y ; body ei ther cyl indr ica l or ar ched dor sally and flattened in to so le ven tral l y , wi th numer ous , smal l , dor sal papi l lae; spicules tables , flat buttons , and/orbr anched rods , never C-

shaped r odsSpicules in body wal l minute grains and s lender , d ichotomous ly branched r ods (F ig . I f)Spicules tables , C or S-

shaped rods , per forated rods and, in some forms , rosettesDor sal papi l lae un i ted into groups of two to eight , the larges t gr oups form ing s tellate figu res ; color ofspecimens at Enewetak yellowi sh r ed, reddish brown , or light brown ; in some specimens , tr ansver sereddi sh black l ines between dor sal papi llae; body lengt h up to 70 cm (F ig . 5c) Th elenota ananas (Jaeger )Dor sal sur face w i th numerous small and some large papi llae, several pap i llae often joined basal ly butnot form ing s tellate figu res ; color l ight gray or yellowi sh gray wi th spots of tan , brown , or red and, in

some specimens , a broken networ k of redd ish black between large papi llae; body length up tocm (F ig . 5a, b) Thelenota anax H . L . Clar kRosettes absen t or , at mos t , very r ar e; tables smal l wi th smal l disk ; no large tack-like tables (F ig . 1 d);dor sal papi l lae limi ted to ambulacr a ; co lor in li fe usually un i formly ver y dar k gr een except dor salpap i l lae t ipped w i th or ange or s car let ; at Enewetak , some specimens dar k o live to blue-black ; bodylength up to about 30 cm (F ig . 5d) St ichop us chloronotus B randtRosettes common ; tack- l ike tables pr esen t or absen t 7

In the papi llae, large r obus t tables wi th con ical , usually s ingle-po inted spi re and large disk wi thnumer ous per forations (F ig . l o) ; color dar k olive green mo ttled wi th deep br own ish green in li fe, dullocher in alcohol ; body length up to about 1 6 cm (F ig . 5e) Stichopus hor r ens SelenkaNo large tack- like tables in papi llae 8

Tables (in specimen examined) small , 35 to 42 pm high and wi de; 0 and S-shaped spi cules 35 to 55um long , rosettes 1 9 to 46 um long (F ig . 1a) ; color of specimen exam ined un i fom'

i ly dar k br own ,

almos t black ; co lor s repor ted in li terature ei ther var iegated whi te or gray wi th br own and yellow ,some specimens showing tinges of pink , lavender , or r ed, or un i formly dar k or l ight brown ; bodylengt h common ly 30 to 40 cm but appar ently r each ing more than 1 00 cm St ichop us ?var iegatus SemperTables (in specimen exam ined) 39 to 70 um h igh , 39 to 62 um acr oss disk ; C-shaped spicules up to70 um long ; r osettes to 54 um long (F ig . 1b); co lor mot tled br own and or ange

-yel low dor sal ly , un iformly d i r ty or ange-yel low ventral ly ; body length cm (F ig . 5i) St ichopus ?var iegatus juven i leBody wall wi th d ichotomous ly br anched X-

shaped spicules (rosettes ) and r ods , never tables 1 0

Tables always among spicules 1 2

No large calci fi ed anal teeth but fi ve gr oups of anal papi llae; sp icu les _

laterally smoo th r ods andr osettes (F ig . 3b); color br own-

g r ay w i th go lden br own or black-r inged dar ker blotches ; body length upto about 30 cm (F ig . 6c) Bohadschia argus JaegerAnus sur rounded by fi ve conspicuous , cal ci fi ed teeth 1 1

(Key to Species con tinued on page

266 CUTRESS AND ROWE

E FF i g 1 A , St ichopus ?var iegatus Semper . a, tables , 0 , S and X-s haped spi cules and r osettes , dor sal i n tegumen t; b, table di sksuppor ti ng r ods , ten tacles .

B , Sti chopus ?var iega tus , juven i le. a, tables . dor sal and ven tr al in tegumen t; b, C-s haped spi cules and r oset tes , dc, per for ated r ods , ven t r al pedi cel s ; d, r ods , ten tacles .

C, St ichopus hor r ens Selenka . a, lar ge table, s ide v iew and di sk of lar ge table, dor sal papi l la; b, smal ler tables ,and tops of spi r es , dor sal i n tegumen t; c, C-s haped spi cules , dor sal and ven tr al i n tegumen t; d, r oset tes . dor sal i n tegumenpor ti ng r od , ven tr al pedi cel ( r edr awn fr om F i s her , 1 907, PI . 70, F ig .

D , St ichopus chlor ono tus B r an dt. a, tables . s ide v iew and tops of spi res , dor sal and ven tr al in tegumen t; b, di sks of tables .c, d, r osette and C-s haped spi cu les , dor sal and ven t ral i n tegumen t; e, r od , ven tr a l pedi cel ( r edr awn fr om M i t suk ur i . 1912,

27 and Theé l , 1 886 , Pl . 7, F i g .

E , H olo thu r ia per vicax Selen ka. a , tables , s ide v iew and di sk ; b, table wi th r udimen tar y spi r e; c, r ods of var i ous shapes

r od ; al l fr om dor sal and ven tr al i n tegumen t ( r edrawn fr om F i sher , 1907, PI . 68 , F ig .

and ven tr al i n tegumen t; c , sp inous r ods and per for ate r ods , ven tr al ped icels ; d, r od. ten tacle.

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268 CUTRESS AND ROWE

1OOEm

F ig . 2A , H o lot hu r ia (H o lodeima) at r a Jaeger . a. tables , s ide vi ews , di s ks and a cr own ; b, X-s haped and i r r egular rod-l ike

spi cu les , ven tr al i n tegumen t; c, smal l gr anule-l i ke depos i ts , ten tacles .

B , H o lot hur ia (M er tens iothu ri a) leucosp i lo to (B r andt). a, tables ; b, buttons and r ods , dor sal and ven tr al i n tegumen t; c. rods ,

ten tacles .

C, H olothu r ia (Thym ios yc io) h i l la Les s on . a, b, tables and butt on s , dor sa l and ven tr al in tegumen t, and per forated r od , dor sal

pap i l la ; c, r ods , ten tac les .

D . H o lothur ia (Plo typer ona) d iffi c i l is Semper . a, b, tables and butt ons , dor sa l and ven tr al i n tegumen t; c, per for ated plate, dor salpapi l la; d, r ods , ten tac les .

E , H o lo thur ia (Thym iosyc io) or en ico lo Semper . a, b, c, tables , button s and per for ated rods , dor sal and ven tr al i n tegumen t; d. rods .

ten tacles .

F, H o lothu r i a (Thym iosyc io) impa t iens (Fot skal ). a , b, c , tables , buttons , per for ated r ods , dor sal and ven tr al in tegumen t; d. r ods ,

ten tacles .

HOLOTH URIANS 289

B - H

Db,Fa, b

of r icana (Semper ). a , th i ck, per f or ated and sp i ned di sks . dor sal and ven t r al In tegumen t; b, r ods , dorsaln t ; c, r ods , ten tacles .

a roset tes and r od, dor sal in tegumen t and papi l la; b, r od and row t tes , ven tr al in tegum en t; c, r ods ,oy an d Gaimard). a, r ods and r oset tes , dor sal in tegumen t; b , rods , ven tr al in tegumen t; c, r ods ,

a, r ods and r oset tes , dor sal in tegumen t; b, r ods and smal l gr ains , ven tr al ln tegu

di sks , s i de v i ew and f rom cr owns , dor sal an d ven t r al In tegumen t ; b, r ods . ven t r al

sp i cu les , sp ined and smooth . dorsal and

270 c CUTRESS AND ROWE

‘f i'l l AM

i g . 4A , Ch i r idoto r i g ida Sem per . a , b , whee ls , r ods and g r a i n s , body wal l and i n tegumen t; c, r ods , ten tacles .

B , Euop to godeff r oyi (Semper ). a , b, c, an chor , an chor plate and r osett es , body wal l and i n tegumen t; d, rod , ten tacles (as repor t ed

an d fi gu r ed by F i s her , 1907, p. 722, PI . 81 , F ig . 3b; not found i n Enewetak specimen nor men ti oned by Sem per ).C, Pendekop lectono n ig r a (Semper ). a, b, c, an chor , an chor plate and r osettes , body wal l and i n tegumen t.D , Synap ta macu lata (Cham i sso and Eysenhardt ). a, b, c, d, an chor , young an chor s , anchor plate and roset tes , body wal l andi n tegum en t.

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272 CUTRESS AND ROWE

22 . Anchor s ver y large, 85 to 1 00 um long and 60 to 70 um across flukes ; anchor plates also large, upto 75 um long ; rosettes small dichotomous ly br anched rods (F ig . 4d) ; color yellow-brown to grayishwhi te wi th or w i thout pat ter n of dar k gr ay spots ; body length up to about 2 m (F ig . 7i )

Synap ta macu lata (Cham isso and Eysenhardt )Anchor s and plates not mass ive, up to 30 pm and 22 um long , r espect ively ; r oset tes smal l , i r r egular ,per forate buttons (F ig . 4c); color trans lucen t reddish brown , wi th many smal l , d i ffuse whi te spots ;body length to about 20 cm (F ig . 79)

APPEN D IX

Col lect ion Recor ds

These r ecor ds ar e on ly of ho lothur ians co l lected for therefer ence col lect ions at the Enewetak Mar ine B io log i calLabor ator y (EMBL ) (later the M id-Paci fic Mar ine Laborator y Mos t of the spec imens were co l lected in

1 955 by Char les E. Cu t ress at snor keling depths and wereidenti fied by Ber tha Cu t r ess . Dur ing subsequen t year s , a

few specimens were added to the co l lections by Rober t W .

H iatt , J . J . Naughton , J . S. Pear se, G . A . H iatt , and

Janet Lamber son . These wer e identi fied by F . W . E .

Rowe, B . Cu t r ess , and J . Lamber son . Recor ds of

Enewetak ho lothur ians men tioned in publ i shed l i terature(Table 1 ) ar e not included here.

Af r ocucum is af r icanaAn iyaan i i (Anan ij): May 8 , 1 955 , inner reef , at tached tounder sur faces of loose cor al or r ock . Enewetak : underrocks around quar r y .

Act inopyga maur i tianaMedr en : May 10 , 1 955 , inner reef, between cor al in t idepoo ls and surge channels . Two commensal worm s the

same co lor as the ground color of the ho lothur ian werefound among the tube feet of one specimen and werepreser ved separately . Seen but not co l lected in 1 955 at

An iyaan i i (Anan ij) , Igur in , Run i t , and Japtan . Enewetak :ocean r eef off r unway .

(F ig . 6 on page

Pendekop lectona n igr a (Semper )Act in ipygo ?maur itiana juven i leAn iyaan i i (Anan ij) : May 1 1 , 1 955 , inner r eef , under r ubble.

Bohadschia argusNancy (E lle) : lagoon s ide, m .

H o lothur ia (Thym iosyc io) or en icoloBun i tz June 20 , 1955 , bur ied in sand under loose coralon causeway . Sand Is land : channel .

H olothur ia (H olodeima) atraRun i t : June 20 , 1955 , in open on sandy or coral bot tomof t ide pools . Seen but not co l lected in 1 955 at Igur i n ,

An iyaan i i (Anan ij) , Medr en , and Japtan . Enewetakocean reef nor th of i s land .

H olothur ia (P lotyperona) diffici l isAn iyaan i (Anan ij) : May 8 and June 21 , 1 955 , inner r eefat tached to under sur faces of coral or r ubble. Enewetak :under r ocks around quar r y .

H olothur ia (Thym iosyc io) h i l laAn iyaan i i (Anan ij) : June 21 , 1 955 , attached to undersur faces of cor al or r ubble . Enewetak : ocean s ide, nor tend, under r ocks jus t below beach ; Sand Is land ; Enjebi :ocean s ide.

H o lothur ia (Thym iosycio) impatien sRun i t : June 20 , 1955, under rock on causeway .

H olothur ia (Mer tens io thu r ia) leucosp i lo toRun i t : June 20 , 1955, under loose coral in poo ls on

causeway . Igur in : May 1 1 , 1 955, inner r eef , under r ubble. Seen but not co l lected in 1 955 at An iyaan i i

(Anan ij) , Medren , and Japtan . Enewetak : ocean s ide,

nor th end“

(Co llect ion r ecords cont inued on page 275 .

F ig . 6A , Af r ocucum is af r i cano (Semper ). Preser ved specimen fr om Enewetak , cm long .

B , Lab idodemos ?semper ianum , juven i le. Pr eser ved Enewetak specimens , and cm long .

C, Bohadsch ia ar gus Jaeger . L i ve specimen in s i tu ; Cocos Lagoon , Guam (photogr aphed by Char les B i r keland).D , Lab idodemos semper ionum Selenka. Pr eser ved Enewetak specimen , 20 cm long .

E , H olo t hur ia (Thym iosycio) h i l la Lesson . L i ve specim en fr om H awai i , 1 9 cm long .

F , H o lot hur ia (Plotyper ona) d ifi i ci l is Semper . Pr eser v ed Enewetak specimen , 6 cm long .

G, H olothur ia (H olodeim a) at r a Jaeger . L i ve specim en fr om H awai i , 30 cm long .

H , H o lot hur ia (Mer tens io thu r ia) per vicox Selenka. L ive spec im en from H awai i , 18 cm long .

I , H o lo t hur ia (M er tens iothu r ia) leucosp i loto (B r andt). L i ve specim en from H awai i , 20 cm long .

(F ig . 7 on page

F i g . 7A , B , H o lo thur ia (Th ym iosyc io) or en icolo Sem per . a, pr eser ved specimen fr om Enewetak , 8 cm long ; b, l ive specimen fr oH awai i , 1 9 cm long .

C, H o lo t hu r ia (Thym iosyc io) impat iens (For skal ). Pr es er ved Enewetak specimen , cm long .

D , Act i nopyga maur i t iana , juven i le. Pr eser ved specimen fr om H awai i , 20 cm long .

E . Act inopyga ?maur i t iona , juven i le. Pr eserved Enewetak specimen , cm long .

F , Ch i r ido to r ig ida Sem per . Pr eser v ed Enewetak specimen , 5 cm long .

G , Pendekop lectano n ig r a (Semper ) . Pr eser ved Enewetak speci men , 12 cm long .

H , Euop to godeff r oyi (Sem per ). Pr es er ved Enewetak specim en , 21 cm long .

I , Synop tu lo macu lata (Cham i ss o and Eysenhardt ). Smal l pr eser ved specimen fr om Enewetak , 21 cm long .

HOLOTH URIANS 273

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Chapter 25

Ech inodermata O ther than H o lo thuro ideaof Enewetak Ato ll

DEN N IS M . DEVAN EY (deceased)

Ber n ice P . B is hop MuseumH ono lu lu , H awai i 968 1 7

IN TRODUCT IONThe echinoderm fauna at Enewetak became known

through publ i cations by A . H . Clar k ( 1952, 1 954) thatr ecor ded nonho lothu r ian species fr om sever al of the

Mar shal l Is lands . These two publ i cat ions included n ine

As ter o idea, n ine Oph iur o idea, and 1 9 Ech ino idea fr omEnewetak . The Cr ino idea and H o lothuroidea were not

r eco rded.

fr om the ato l l .In the late 1 9505 and ear ly 1 9605 , Fred C . Ziesen

henne, then wi th the A l lan H ancock Foundat ion (AH F ) ,was asked by the M id-Paci fi c Resear ch Labor atory (MPRL )to prov ide an ident i fied r eference collect ion of nanholothur ian echinoderms . H is ident ifi cat ions , based on specimens that he co l lected and those co llected by s tuden ts andr esear cher s wor k ing at MPRL , prov ided not on l y a valuable r eference co l lect ion but also increased the number of

taxa known from Enewetak . An unpubl ished repor t , wh ichincludes habi tat informat ion and keys to man y of the

species , was prepar ed by F . C . Z iesenhenne and

R . A . Boo loot ian and was left at the laboratory for use byv is i t ing r esear cher s . The holothur ians were being s tud iedseparatel y by Ber tha Cu t r ess , and her wor k has beenupdated in a separate chapter for the pr esen t volume (seeCu t r ess and Rowe , Chapter

Since these ear ly effor ts , new s tud ies and add i t ionalrecor ds of ech inoderms have been comp i led by MPRL personnel and other r esear cher s wor k ing at Enewetak . Th ischapter br ings together these numerous r ecor ds (Table

A lthough the major i ty of Enewetak ’

s shal low-water (toabout 1 00 m ) ech inoderms is r easonably well known , faunafrom deeper water s , especial l y in pas ses and off the sea

war d r eef face, i s s t i l l incompletely known . Less than 196of the species so far r ecor ded have been taken in waterdeeper than 1 00 m .

In compar ison w i th the Mar shal l Is lands as a who le,

knowledge of the Enewetak ech inoderms is cons iderable.

At th is one ato l l , 8696 of all Mar shal l Is lands spec ies ar e

now known to occur at Enewetak .

CRINO IDEA

A lthough cr ino ids had been r epor ted fr om the Mar shallIs lands by G is lén ( 1 940) and A . H . Clar k ( 1952 , no

spec ies fr om Enewetak were included . Cr inoids col lected inthe late 1 9505 and ident i fied by F . C. Ziesenhenne

included Coman thus bennett i and Comas ter gr aci l is , speciesr ecor ded pr ev ious ly in the Mar shal l Is lands fr om Rongelapand B ik in i by A . H . Clar k ( 1952) and from A r no by A . H .

Clar k Banner and Banner ( 1968) repor ted the

alpheid shr imp , Synalpheus deman i , fr om C . bennett i collected at Enewetak . Zmar z ly (per sonal commun i cat ion)doubts the assoc iat ion of alpheids w i th th is cr inoid and,

based on her s tudies , feels that the hos t was a di fferentspec ies . One of the Enewetak specimens of C . bennettiexamined by the author and Ann Field ing had a s inglealpheid, S . s t imp son i , assoc iated ; two other alpheids wer ealso assoc iated w i th other cr ino id spec ies at Enewetak(Devaney and B r uce, Chapter 22 , th is vo lume) . One of

these 8 . car inatus , was also noted in assoc iat ion wi th cr i

no ids at Enewetak by Banner and Banner H umes( 1 972) r epo r ted a new spec ies of copepod fr om C. ben

nett i , and Bar to l in i et ai . ( 1 973) exam ined pigments fr omEnewetak spec imens of th is cr ino id .

In 1976 , Devaney and Field ing exam ined Enewetak cr ino ids for thei r symbionts ; cr us taceans , po l ychaetes , myzostomes , and other inver tebr ates were noted and col lected . Itwas thought that on l y Coman thus bennett i and Comas tergr aci l is wer e r epr esented in the mater ial examined . Subsequen tl y , however , due to a more intens ive s tudy on

Enewetak cr inoid ecolog y and thei r symbionts by Zmar zlyin 1980 , two add i t ional species (Coman thina schlegel i and

Coman thus par vici r r us ) were r ecogn i zed among the 1976

mater ial . The cr us taceans as soc iated w i th these cr ino idshave been tabulated in the pr esent volume (Devaney andB r uce, pp . 225 and more in format ion was g iven in

Zmar zly The smal l 1 0-armed antedon id, Do romet ro

nana, was col lected in 1 976 off one of the souther n w indwar d is lets (Anan ij) beneath coral in l imes tone rubble at 1 8

DEVANEY

TABLE 1

Check l i s t of Ech i noderm ata O ther than H olo thu r oidea f r om Enewetak A tol l

Phy lum ECH INODERMATASubphy lum CRINOZOAClass CR INOIDEAFam i ly COMASTERIDAEComan thina schlegel i (Carpenter , Zmar z ly, 1984

Coman thu s bennetti (M ii ller , Banner and Banner , 1 968 ; H umes , 1972; Bar to l in i et al . , 1973 ; Zmar z ly, 1 984 .

Coman thus par vici r r us (M ii ller , Zmar zly, 1984 .

Comas ter gr aci lis (H ar t laub , Zmar z ly, 1 984 .

Fam i ly EUDIOCRIN IDAEEudiocr inus ten u iss im us G is lén , 1 940 : Zmar zly, 1 984 .

Fam i ly ANTEDON IDAEDo rometr o nana (H ar t laub, Zmar z ly, 1 984 .

Subphy lum ASTEROZOAClass STELLEROIDEASubc lass ASTEROIDEAFam i ly ASTROPECT IN IDAEAs t ropecten po lyacan thus Mu l ler and T r oschel , 1 842 : A . H . Clark , 1 952 .

Fam i ly OREASTERIDAE‘Chor ios ter gr on u lotus L ii tken , 1 869 .

Cu lcita novaegu ineae M i l ller and T r oschel , 1 842 : A . H . Clark , 1952 ; H umes , 1971 ; B ruce, 1 979 .

Fam i ly OPH IDIASTERIDAE‘TF r om io hem iop lo Fisher , 1913 .

Pr om ic m i llepo rello (Lamarck ,t F rom io bolan sae Per r ier , 1875: A . H . Clar k , 1 952 .

F rom ia mon i l is Per r ier , 1 875.

Gomophia egyp t ioca Gray , 1 840 .

‘Leias ter speciosus von Mar tens , 1 866 .

L inckio laevigata (L innaeus , A . H . Clark , 1952 .

L inckio m u lt iforo (Lamarck , A . H . Clar k , 1952 ; 1 954 .

‘Neoferdina cum ingi G r ay , 1 840 .

‘Ophidios ter gran ifer L i i tken , 1 871 : A . H . Clark , 1 952 .

Ophidios ter robi llardi de Lor io l , 1885 : Mar sh , 1 977 .

Oph idios ter lo r iol i Fi sher , 1906 : A . H . Clark , 1952 .

Fam i ly ASTEROPSEIDAE‘As ter ops is car in ifera (Lamar ck ,Fam i ly ASTER IN IDAE‘§As ter ino anomala H . L . Clar k , 1921 .

Fam i ly ASTER IN IDAEAs ter ino bu r ton i Gr ay , 1 840 : A . M . Clark and Rowe, 1 971 .

As ter ino cepheo (M ii ller and T roschel , A . H . Clark , 1 952 .

Fam i ly ACANTH ASTERIDAEAcan thas ter p lanci (L innaeus , A l len , 1972 ; H ighsm ith , 1 980 .

Fam i ly MITH RODIDAE‘M ithrod io clovigero (Lamar ck ,Fam i ly ECH INASTER IDAEEch inos ter luz on icus (Gray ,Oth i lia luz on ica Gray : A . H . Clark , 1952; Lawr ence and Gu i lle, 1982 .

Subc lass OPH IUROIDEAFam i ly OPH IOMYXIDAE‘Neop lax crass ipes Koehler , 1 922 .

‘Ophiomyxo aus tralis L ii tken , 1869 .

Fam i ly GORGONOCEPH ALIDAE‘As t r oboo nuda (L yman ,

‘New r eco rd at Enewetak .TEnewetak specimen s determ ined by F . C. Z iesenhenne at AFH (1 dry) and MPRL ; those repor ted as E . hem iop lo fr om B ikin iby A . H . Clar k ( 1952) probable juven i les of other species (L . M . Mar sh , per sonal commun ication . )t Cons ider ed probably r efer r able to F . m i lleporello after r eexam i nation by L . M .

-Mar sh (per sonal -commun icati on ).§Cons ider ed as poss ible syn . of As ter ino bu r ton i by A . M . Clark and Rowe, 1 971 , p. 68 .

(Th is table continued on next page. )

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DEVANEY

TABLE 1 (con t’d)

Fam i ly ECH INOMETRIDAE (cont ’d)Echinometr o oblonga (de B lainv i l le,

Echinometra mathaei var . oblonga (de B lainvi l le) : A . H . Clark , 1 952 .

Ech inos tr ephus ocicu lotus A . Agass i z , 1 863 : A . H . Clark , 1 952; Pomeroy and Kuenzler , 1967 ; Russo , 1 980 ; Lawr ence,

and Gu i lle, 1 982; Eber t , 1982 .

‘H eterocen t r otus mamm i llatus (L innaeus ,H eterocen trotus tr igonar iu s (Lamarck , A . H . Clark, 1 952; Odum and Odum , 1 955; Lawrence and Dawes , 1 969;H umes , 1970 ; Eber t , 1982 .

Fam i ly DIADEMATIDAE‘D iadema savignyi M ichel in , 1 845 .

Ech inothr ix calamari s (Pal las , A . H . Clark , 1952; H umes , 1977.

Ech inothr ix diadema (L innaeus , A . H . Clark, 1952; Lawr ence and Gu i lle, 1982 ; Eber t , 1 982.

Fam i ly PARASALENIIDAE .

Par osolen io gr ot ioso A . Agass i z , 1 863 : A. H . Clar k, 1 952; H ighsm i th , 198 1 .

Fam i ly TEMNOPLEURIDAEMesp i l io g lobu lus (L innaeus , A . H . Clark , 1 952 .

Fam i ly TOXOPNEUSTIDAECyr techin us ver r ucu lotus (Lu tken , H ighsmith , 198 1 .

‘Toxop neus tes p i leo lus (Lamarck ,Tr ipneus tes gr at i llo (L innaeus , A . H . Clark , 1 952.

Fam i ly ECH INONEIDAEEchinoneus obnormalis de Lor io l , 1 883 : A . H . Clark , 1952.

Ech inoneus cyclos tomus Leske, 1 778 : A . H . Clark , 1 952 .

Fam i ly SCH IZASTERIDAE‘D ip loporos ter savignyi (Fou r tau ,

Fam i ly FlBULARlIDAEEchinocyamus aus tr alis (Desmou l ins ,Fibu lar ia aus tr alis Desmoul ins : A . H . Clark , 1952 .

‘Echinocyomus cr ispus Mazett i , 1893 .

Echinocyam us megopetolus H . L . Clar k , 1914 : A . H . Clark , 1952.

Fibu lar ia ovulum Lamarck , 1 788 : .A . H . Clark , 1 952 .

Fibu lar ia vo lvo L . Agass i z and Desar , 1 847 : A . H . Clark , 1 952 .

Fam i ly SPATANG IDAEMoretio p lanu lata (Lamar ck ,Maretic ovota (Leske, A . H . Clark , 1 952 .

Fam i ly BRISSIDAEB r i ss us latecor inatus (Leske, Pomeroy and Kuenzler , 1 967.

Meta l io dierona H . L . Clar k, 191 7 : A . H . Clark , 1 952 .

‘Meta lia spatagus (L innaeus ,Rhynobr iss us hem ias tero ides A . Agass i z , 1 879 : Pomeroy and Kuenz ler , 1967 .

Fam i ly CLYPEASTERIDAE‘Clypeas ter ret icu latus (L innaeus ,Fam i ly LAGANIDAELogan um depr essum L . Agass i z , 1841 : A . H . Clark , 1 952 .

‘New r ecord at Enewetak .

to 21 m and was ident i fied by the author . Addi tional specimens of th is spec ies have been found under cor al r ubble inthe lagoon . Dor omet ro nana had been recor ded fr om Ebonand Jalu i t by G i s lén In 1980 at Enewetak , a

five-

armed Eudiocr inus , ten tat ively ident i fied as E . tenu iss im us , was found in recesses on the s teep seawar d dropoffnear R i gi l i is let .

“

G is lén (1940) recor ded thi sl '

species or igi

nal l y fr om Jalu i t Ato l l in the Mar shal ls .

Six cr ino id species ar e now known fr om Enewetak shallow water s ; all but one (E . ten u is s im us) occur in otherlndo-Wes t Paci fic ar eas outs ide the Mar shal l Is lands (A . M .

Clar k and Rowe, Add i t ional cr ino ids r ecor ded fr omother Mar shal l Is lands include: Cenomet ro bel la var .

magn if ica from Jalu i t (G is lén , as wel l as

Stephanometr a indica (as S . p rotectus ) from Ebon and

Jalu i t (G i s lén , 1940) and (as S. indica p r otectus ) fr omRongelap and Ronger ik (A . H . Clar k ,Six of the n ine cr ino ids known fr om the Mar shall

Is lands have been found at Enewetak . Seven spec ies are

known from Guam and 21 from Palau (Meyer and

Macu rda , Enewetak shar es fou r spec ies w i th Guamand five w i th Palau .

i ER THAN HOLOTHUROIDEA 28 1

Oph iocom idae

The ophiocom ids ar e d iver se at Enewetak r epresentedby 1 6 species in five gener a w i th a new r ecor d ,Ophiar thr um elegans . One specimen of Ophiop s i la remainsto be iden t i fied to spec ies .

Oph ioner eididae

Two species of the ophioner eids are r ecogn i zed :Ophionereis por r ecto and Ophionereis sp . , wh ich r esemblethose r ecor ded by A . H . Clar k (1952) fr om B ik in i Ato l l asOphionereis degener i .

Oph iodermatndae

F ive ophiodermat ids occu r at Enewetak . Ophior ochnaaff in is , Ophiorochnello gorgon ia, and Ophiocon is perm ix ta

are new recor ds for the Mar shal l Is lands . Ophiopeza sp i

noso was r ecor ded (as Oph iopezella sp inosa by A . H .

Clar k , 1952) fr om B ik in i and Ronger ik Atol ls . The on lyother ophiodermat id r ecor ded from the Mar shal l Is lands isOphiopezella (Ophiopeza) dubiosa fr om Jalu i t (Koehler ,

An addi t ional specimen of the uncommonOph ior ochnello par visp ina, repor ted fr om Enewetak byDevaney was located in the A l lan H ancock Foundation fr om the in ter t idal lagoon , July 1 7 , 1960 .

Oph i u r i dae

Of the four ophiur ids known from Enewetak on ly one,

Ophiop locus imbr icotus , a w idespread warm water IndoWes t Paci fic species has been prev ious ly repor ted fr omth is ato l l . All four oph iur ids have been repor ted fr om elsewher e in the Mar shal l Is lands (A . H . Clar k ,

The Col lection Recor ds l is t the habi tats of the newlyr epor ted species of Oph iur o idea. Table 2 l is ts commentson ophiur o id color , anatomy , d isc and arm lengt h s i ze, and

repos i tor y .

ASTERO IDEA

As t r opect i n idae

The fam i ly As tropect in idae is represented by a s inglew idespr ead lndo-Wes t Paci fic species , As t ropecten po lyacan thus , co l lected in the lagoon (A . H . Clar k , Two

smal l specimens were co l lected at B ik in i Ato ll in July 1946and ident i fied by F . C. Z iesenhenne (AH F ) .

O r eas ter i dae

The fami l y Or eas ter idae is r epresented by two species .

Cu lcita novaegu ineae has been co llected on the ocean reefat Enewetak (A . H . Clar k , 1 952) and from the lagoon and

channel . Symbiot ic crus taceans were recor ded from thisspecies by H umes ( 1 971 ) and B r uce A secondor eas ter id, Chor ias ter gr onu lotus h i ther to unknown fromthe Mar shal l Is lands , was discovered dur i ng the H awai i

282 DEVANEY

TABLE 2

Commen ts on Color , D i sc and A rm Lengt h , Si ze, and Repos i tor y

Neop lax cras s ipes , co lor deep brown with wh i te mark ings , one specimen dry wi th disc diameter (d d) 13 mm , AH F .

Ophiomyxa aus t ral is , co lor in alc . dul l s traw , arm spines tipped wi th r ed-brown , di sc and mouth l ight gray , 1 alc . MPRL .

As troboa nuda, co lor above d isc purp l ish black also main arm axes except l ighter be lts of hook lets , a darker band between eachr ow , br oader above and nar r owly cur ved distal ly ar ound s ides of arm s ; nar r ow depr essed longi tudinal dark line down centerof arm s , mor e distal arm br anches qu ite l ight in co lor ; lower sur face of arm s l ighter than above wi thout dark bands orlongitudinal l ine, a few distal br anches wi th some dark color not un iform , 1 dry, d d. 85 mm , MPRL .

Amph iu ro luetken i , r ed and wh ite disc wi th reddish arms .

Amph iod ia sp . , wh i te disc , wh ite arm s , 1 dry, di sc to 3 mm , arm 60 mm , BPBM .

Mocrophiothr ix longipedo, fr om Araanbir u , 1 alc . d d. mm , arm 300 mm , MPRL ; from Engebi , 1 alc , dd . mm ,

arm 132 mm , MPRL ; fr om Engebi , 1 spec. , disc 1 3 mm , arm 155 mm , MPRL .

Macr ophiothr ix rhabdoto , 1 dry , d d. 8 mm , arms broken , MPRL ; 1 alc. dd . 8 mm , arm s broken , AH F .

Oph iothelo donae, disc blue or gr een , arms annulated wi th narr ow band of l ight or dark acr oss , 1 0+ dry, disc to 4 mm usuallywi th s ix arm s or in fi ss iparous s tage with on ly thr ee arms , BPBM .

Ophiothr ix (Acon thOphiothr ix ) pr oteus , 1 dry, d d. 9 mm , arm s 1 00 to 1 10 mm , BPBM .

Ophiothr ix (Acon thoph iothr ix ) purpurea, color of wide,dark str ipe or along m iddle or nar r ow dark str ipe bordered by l ight l ines on

dor sal s ide of arm s ; arm spines banded dar k and l ight ; radial sh ields bar e, but between , r adial ly and inter r adial ly ,either granular s tumps or stumps and a few long spines , marginal ly and on ventral s ide, disc wi th stumps and spines widely spaced,1 alc . , d d. 7 mm ,

BPBM .

Ophiothr ix (Ophiothr ix ) t r i l ineata,fr om Enewetak Is let , 1 spec . , gonads r ipe; from Japtan Is let , 1 spec . gonads r ipe; fr om

Medr en , 2 spec . , gonads r ipe .

Ophian thr um elegans , l iv ing co lor dark black , arm s r ed and wh itish band wi th some smal l dark speck l ing between arm spines withi r r egular dark and l ight marks , fr om Enewetak , 1 alc . , d d. 1 2 mm , BPBM ; Sand Is let , 2 alc . , d d. 10, dd . 1 3 mm ,

BPBM ; Lagoon , color dark black with red arm s , 1 dry, BPBM .

Oph ionereis sp . , from Enewetak Is let , 2 dry, d d. 5, d d. 6 , BPBM .

Ophiorochno aff in is , 1 alc . d d. 1 2 mm , MPRL .

Ophior ochnello gorgon ia, from Enewetak Is let , 1 dry, d d. 12 mm,BPBM ; fr om Rabaion Is let , 3 dry, d d. 14 to 1 6 mm , AH F .

Oph iocon is perm ix to, 1 alc . , d d. 2 mm , BPBM .

Oph iolep is super ba , fr om Enewetak Is let , 1 alc . , d d. 24 mm , AH F .

Ophiolep is sp . after 0 . cin ta, from Enewetak Is let, 1 dry d d . 13 mm , 1 dry , d d. mm , BPBM ; fr om Lagoon , 1 spec . d d. 7

mm , AH F .

Oph iur a kinberg i , fr om Engebi Is let , 1 juv . , alc . , d d. 3 mm , MPRL .

Under sea Resear ch Laborator y dives in 1 98 1 at a depth of1 22 to 1 30 m .

Oph idias ter i dae

Th i s fami ly has . the lar ges t number of speciesr epr esen ted at Enewetak , w i th at leas t 1 0 now r ecor ded .

Four have been r epo r ted pr ev ious ly from the Mar shal lIs lands ; all are w idespr ead lndo-Wes t Pac i fi c species .

As ter opseidae

The As ter opseidae i s r ecor ded for the fi r s t t ime in theMar shal l Is lands in the form of A s ter op s is cor in ifer o , a

spec ies wh ich r anges in the Indo-Wes t Pac i fic from H awaithrough Ocean ia and into the Ind ian Ocean .

A s ter i n idae

The As ter in idae at Enewetak ar e repr esented by tw<species and poss ibly a th i r d (As ter ino coronata) determ ine<by F . C . Z iesenhenne . A ll three have been pr ev ious lg

r ecor ded fr om the Mar shall Is lands (A . H . Clar k ,Acan thas ter idae

Acan thas ter p lanei , the crown -

of-tho r ns sea s tar , i

known at Enewetak fr om s tud ies of i ts pr edator y act iv i tyon scler act in ian co rals (H ighsm i th , 1 980 ,

1 982) and a f is l

s ym biont (A llan ,

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234 DEVAN

Rober t s , or i g . det . F . C . Ziesenhenne, ver i fi ed by D . M .

Devaney (AH F ) .Ophiomyxo aus t ralisEnewetak Is let : Augus t 1 957, 7 to 8 m , R ichar dN i sh ioka, or ig . det . F . C. Ziesenhenne, ver i fied byD . M . Devaney (MPRL ) .

Gor gonocephal idae

As t r oboo nudaKidrenen Is let : May 26 , 1976 , ocean s ide, depth of 1 3m in a cave on the seawar d s ide jus t nor th of Kidr enenIs let , det . D . M . Devaney (MPRL ) .

Amph iur idae

Amphiphql is squamoto

Lagoon : July 1 960 , shal low , J . H . Rober ts (Stn . 252-B) ,id. F . C . Ziesenhenne; 1 dry (AH F ) . Enewetak Is let : July25, 1 957,

“ found abundantl y” in the s ludge area of the

sewage plan t discharge l ine; the on ly ar ea wher e s i lt andsediment wer e found on the reef; call . and id. F . C.

Ziesenhenne; 80 alc . (30 MPRL ; 55 AH F ) . Run i t Is letJuly 8 , 1959 , in ter t idal , ft tide, co l l . J . S. Gar th andF . C. Ziesenhenne, id. F . C. Ziesenhenne; 1

Japtan Is let : July 1 9, 1 957, seawar d r eef; col l . and id.

F . C. Ziesenhenne; 20 to 30 alc . (MPRL ) . Engebi Is let :July 21 , 1 959 , in ter t idal , ft tide, co l l . J . S. Gar thand F . C . Ziesenhenne, id. F . C . Ziesenhenne; 4 alc .

(AH F ) . Igur in Is let : July 1 1 , 1 959, inter t idal , ft t ide,

col l . by J . S. Gar th and F . C. Ziesenhenne, id. F . C.

Z iesenhenne, 1 alc . (AH F ) . Enewetak Is let : Aug . 9,

1965 , under coral , rock , quarr y , D . R . Stokes , det .

D . M . Devaney , l spec . (BPBM) .Amphiod io sp .

Lagoon : Sept . 24 , 1 980 , ai r l i ft sample of lagoon bottom , 35 it off Enewetak Is let , call . and id. D . M . Deva

ney, 1 dry (BPBM ) .

Oph iot r ich idae

Mocr ophiothr ix longipedo

Ar aanbi r u Is let : July 22 , 1 959 , id. F . C. Ziesenhenne

(MPRL ) . Engebi Is let : July 21 , 1959; id.

F . C . Ziesenhenne; 2 spec . alc . (MPRL ) . Igur in Is let :September 1 956 , outer r eef , 2 to 3 ft depth , col l .D . Watson , id. F . C. Ziesenhenne, 1 spec . (MPRL ) .Jap tan Is let : July 1 8 , 1959 , id. F . C. Ziesenhenne; 4

spec . (MPRL ) . (Note, at leas t 1 0 addi t ional lots at AH F[not exam ined by D . M . Devaney , det . byF . C.

Mocr ophiothr ix r habdo to

Enewetak Is let : Feb . 24 , 1957, call . A . H . Banner , id.

F . C . Z iesenhenne, 1 spec . dry (MPRL ) . Enewetak Is let :Feb . 27 , 1 957 , lagoon s ide about 1A of lengt h of souther n t ip of i s let , sandy bo ttom ca . 2 m deep , heads of

Poci llopor a and br anch ing Acr opo r a, A . H . Banner (Stn .

B . E. id. F . C . Ziesenhenne, ver i fied (insofar as poss ible) by D . M . Devaney , 1 spec . alc . (AH F ) .

Ophiothelo danae

Enewetak Is let : May 21 , 1 975, depth 8 ft , lagoon ,cling

ing to soft nephtheid cor al co lony that was attached tochunks of dead coral bur ied in sandy bottom ,

co l l .

ER THAN HOLOTHUROIDEA 285

G is lén , T . , 1 940 , A Co l lection of Cr ino ids from the South SeaIs lands , K . s vensko Vetensk Akad . H and l . Ser ies 3 , 18

3- 16 .

H ighsm ith , R . C. , 1 980 , Bur r owing by the B ivalve Mo l luscL i thophaga cur ta in the L iv ing Reef Coral Mon t ipo r a ber ryiand a H ypothes is of Recipr oca l Lar val Recru itment , Mar . B io l .(Ber lin) , 56 : 1 55-1 62 .

1 98 1 , Coral B ioer os ions at Enewetak : Agents and Dynam ics ,In t . Rev . Gesam ten . H ydrobiol . , 66 : 335

-375 .

H umes , A . G . , 1 970 , Clavisodal is heter ocen tr ot i gen . et sp . n . , a

Cyclopo id Copepod Paras itic on an Ech ino id at En iwetokAto l l , Jour . Par as i t . , 56 : 575-583 .

1 971 , Cyclopo id Copepods (Stell icom i t idae) Paras itic on SeaStar s fr om Madagasca r and En iwetok Ato l l , J . Paras i tol . , 57 :1 330- 1 343 .

1 972 , Pseudan thes s ius coman thi n . sp . (Copepoda, Cyclopo ida) Assoc iated wi th a Cr ino id at En iwetok Ato l l , Poc . Sc i . ,26 : 373-380 .

1 977 , Pseudan thess id Copepods (Cyclopoida) Associatedwi th Cr ino ids and Ech ino ids (Ech inodermata) in the T r opica lWestern Paci fic Ocean , Sm ithson . Con tr ib . Zoo l . No . 243 , pp .1 4 3 .

K ing , C . E . 1 964 , Relative Abundance of Species and

MacAr thu r 5 Model , Eco logy , 45 : 71 6-727.

Lawr ence, J . M . , 1970 , The Effects of Star vation on the L ipidand Car bohydrate Levels of the T ropical Sea U rch inEchinometra mathaei (de B lainvi l le), Pac . Sci . , 487-489 .

and C. J . Dav ies , 1969, A lgal Growth over the Ep ider rn is of

Sea U r ch in Spines , J . Phyco l . , 5 269 .

and A . Gu i lle, 1 982, Or ganic Compos ition of T r opica l , Po larand Temper ate-Water Ech inoderm s , Comp . B iochem . Phys io l . ,728 283-287 .

Mar sh , L . M . , 1 977 , Coral Reef As tero ids of Palau, Car o l ineIs lands , M icr ones ica, 1 3 251 -28 1 .

Meyer D . L . , and D . B . Macurda, Jr . , 1 980 , Eco logy and Di s tr ibution of the Shal low-Water Cr ino ids of Palau and Guam ,

M icr ones ica, 16 59-99 .

Pomeroy , L . R . , and E . J . Kuenz ler , 1 967, Phosphoru s Tur noverby Coral Reef An imals , Sympos ium on Radioeco logy, D . J .Nelson , and F . C . Evans Ann Ar bor , Michigan ,

pp . 474-482 .

Russo , A . R . , 1 980 , B ioeros ion by Two Rock Bor ing Echinoids(Echinometra mathaei and Echinos t r ephus ocicu lotus ) on

Enewetak Ato l l , Mar shall Is lands , J. Mar . Res . , 38 : 99

Sloan , N . A . , A . M . Clark , and J. D . Taylor , 1 979, The Ech inoderm s of Aldabr a and Thei r H abitats , Bu l l . B r . M us . (Nat .H is t .) Zoo l . , 37 : 8 1 - 1 28 .

Zmar z ly, D . L . , 1 984, Di s tr ibution and Eco logy of Shal low-WaterCr ino ids at Enewetak Ato l l , Mar shal l Is lands , wi th an Annofated Check l ist of Thei r Symbionts , Poc. Sc i . , 38 : 1 05-1 22 .

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288 ELDRED

known d i s tr ibut ion of D ip losoma s im i lis (as separated fr omD ip losoma vi rens ) includes Indones ia , the Gr eat Bar r ierReef , F i j i , Japan , and ques t ionably the Ph i l ipp ine Is lands .

D idem n um gr ande, a w idespr ead spec ies , is found in

Japan , Ph i lipp ine Is lands , Malays ia, Aus tralia , Tasman ia,

south Ar abia, and Palmyr a in the L ine Is lands . D idem num

qu incunciale has been known pr ev ious l y on ly from Zan z ibar and Tr ididemnum cyclops fr om Aus tral ia and Madagas

car .

Tokioka (1 967) s tudied the ascidians in the U . S.

National Museum and recor ded four species fr om otherato l ls in the Mar shal l Is lands . These are:

D ip losoma vi r en s H ar tmeyer , 1909 (Lep toc l in um

vi ren s) — EbonAscidio aper ta Slu i ter , 1 904 - RongelapPo lycar pa cryp tocorpa (Slui ter , 1 885) — B ik in iPo lycarpa iwoyomae Tok ioka, 1 950

— Ar no

No specimens wer e found fr om Enewetak .D idemn ids show an increase in diver s i ty towar d the

Indo-Malayan region . The didemn id fauna of Enewetakcon tains a number of w idespr ead species , showing i ts

s trong faun i s t ic affin i ty to the Indc es t Paci fic .Seven species of lar vaceans (Ger ber , 1 98 1 ) and two

species of thal iaceans (salps ) (G i lmar tin , 1958) have been

Chapter 27

Annotated Checklis t of the F is hes ofEnewetak Ato ll and Other M ars ha ll Is lands

JOH N E . RANDALL and H ELEN A. RANDALL

Ber n ice P . B ishop MuseumH onolulu, H awai i 968 1 7

IN TRODUCT IONIntens ive fish co l lect ing was car r ied ou t in the Mar shall

Is lands in 1946 and 1947 in connect ion w i th and fo l lowingthe atom bomb tes ts of Operat ion Cros s roads . These col

lect ions wer e depos i ted in the Nat ional Museum of Natur alH is tor y of the Sm i thson ian Ins t i tut ion in Wash ington ,

D . C . Add i t ional Mar shall Is lands fi shes were taken in

1 948 and 1 949 dur ing exped i t ions of the Appl iedF isher ies Labo r ator y of the Un iver s i ty of Wash ington . All

these fi shes , plus co l lections fr om the Mar iana Is lands ,were s tud ied by Leonar d P . Schultz , then Curator of

F i shes of the Nat ional Museum of Natur al H is tor y , and hisco l laborator s , W i lber t M . Chapman , Ear l S. H er ald ,Er nes t A . Lachner , Ar thur D . Welander , and Lor en P .

Woods . The r esult was the valuable thr ee-volume wor k onthe fishes of the Mar shal l and Mar iana Is lands ( 1 953 to

A total of 543 species of fi shes was r epor ted fromthe Mar shal l Is lands in these thr ee vo lumes . U n for tunately ,a s tudy of the fam i ly w i th the larges t number of species ,the Gobi idae, was not completed .

Ver y few fi shes wer e repo r ted from the Mar shal lIs lands pr ior to 1953 . G i inther (1 873 to 1 91 0) l is ted 3 1species fr om the atol ls of Ebon , Jalu i t , and Majur o [thoughhis Angu i l la maur itiana and Monocan thus oyr audi are probable local i ty er r or s , and his Echeneis scutota (Remora aus

tr al is) is suspect]. Kendal l and Goldsbor ough ( 1 91 1 )recor ded 58 species . In a s tudy of po isonous fishes of the

Mar shal ls , H i yama (1 943) i l lus tr ated 7 1 species in color .

The text (tr ans lated from the Japanese by Van Campen ,

1 950) deals mor e w i th the toxici ty of these fishes thanthei r class i fi cation ,

but the h igh qual i ty of mos t of the f igures enables one to make pos i t ive ident i ficat ions to

species .

Donald W . Str asbur g (1 953 ) subm i tted a r eport to the

Off ice of Naval Resear ch ent i tled F is hes of the Souther nMar s ha l l Is lan ds . H e tr eated 345 species in thi s repor t ,lar gel y fr om col lect ions made at Ar no . H e wor ked closelyw i th Schultz et al . and had access to thei r manuscr ipts onMar shall Is lands fishes . H e l i s ted 26 species of fi shes thatwer e not included in the manus cr ipts of Schultz et al . at

that t ime ( later a few of these fishes wer e r epor ted inVols . 2 and 3 of F ishes of the Mar s ha l l and Mar ianas

Is lands ) .lchthyo log ical activ i ty in the Mar shal l Is lands cont inued

in the year s fo l low ing the r esear ch of Schultz et al . , largelyas a r esult of the es tabl ishment of the M id-Pac i fic Resear chLabor ator y on Enewetak (in i t iall y named En iwetok Mar ineB iolog ical Laborator y) . A lso of impor tance has been the

pr esence of a number of scuba d iver s at Kwajalein w i th aninteres t in fishes . Some of these are exper t underwaterphotographer s , and thei r effor ts to captur e the r ar e or

unusual spec ies on fi lm have prov ided new r ecor ds of

fi shes fr om the ar ch ipelago .

Randal l (1 986) r ev iewed all the new r ecor ds and new

species of fi shes of the Mar shal l Is lands that have beendocumen ted s ince Schultz et al . , including those of

Str asbur g ’

s Off ice of Naval Resear ch r epor t (s ince few cop

ies were d i s tr ibuted) . A to tal of 86 such fishes was foundin the l i ter atur e, and another 1 06 new r ecor ds were addedby Randal l . A lso there ar e about 45 species of fishes fromthe Mar shal ls wh ich ar e s t i l l undescr ibed .

In add i t ion to new r ecor ds and new species , man ychanges in the names of fi shes as l is ted by Schultz et al .have been made in r ecent publ ications .

The pr esent paper pr ov ides a checkl i s t (Table 1 ) of ther eef , shor e, and epipelagic fishes known fr om the Mar shal lIs lands . We r ecor d a total of 8 1 7 spec ies in 338 gener aand 92 fam i l ies . Deep-water fishes have been om i tted from

290 RANDALL AND RANDALL

TABLE 1

Check l i s t of t he F i shes of the Mar shal l Is lan ds

Phy lum CH ORDATAClass CH ONDRICH TH YSSubclas s ELASMOBRANCH IIOrder H EXANCH IFORMESFam i ly H EXANCH IDAEH exanchus g r iseus (Bonnater r e, Randal l , 1 986 .

Order LAMNIFORMESFam i ly G INGLYMOSTOMATIDAENebr ius concolor

‘

R i i ppell , 1 837

G ing lym os toma fer rug ineum Schu ltz , 1 953 .

Nebr ius fer rug ineus Randal l , l 980a .

Fam i ly ISURIDAECar char odon car char ias (L innaeus , Schultz , 1 953 .

Fam i ly CARCH ARH IN IDAECar char h inus olbimarginatu s (Ruppell , Schultz , 1 953 .

Car char h inus amblyr hynchos (B leeker ,Car char h inus men is or r ah Schultz , 1953 . Gar r ick ( 1 982) concluded that the true C. men isor rah (Valenciennes in M i i llerand H en le) is a jun ior synonym of C . fa lciform is (B ibr on in M ii ller and H en le) .

Car char h inus falciform is (B ibron , Gar r ick , 1982 .

Car char h inus galapagens is (Snodgrass and H el ler , Randal l , 1 980a.

Car char h inus l imbatus (Valenciennes , Randal l , 1 980a .

Car char h inus long im on us (Poey ,Car char h inus brachyurus Schu ltz , 1 953 . Gar r ick (1 982) stated that the specimens Schultz identi fied as C . brachyurusare C . long imon us . Gar r ick po inted ou t that C . moou (Les son) is an o lder name for longimonus , but he gave reasons for

retain ing the long-

es tabl ished longimonus .

Car char h inus melanop ter us (Quoy and Gaimard, Schultz , 1953 .

Galeocer do cuvier (Peron and Les ueur , Schu ltz , 1 953 .

Negapr ion ocu tiden s (Ruppell ,H em igaleop s fos ter i Schu ltz and Welander , 1953 . Gar r i ck and Schu ltz in G i lber t ( 1963) referr ed H em igaleop s fos ter i tothe synonymy of Negapr ion ocu tidens .

Tr iaenodon obes us (Ruppell , Schultz (1 953 ) class ified th is shark in the T r iakidae; recent author s place i t InCarchar h in idae; however , A . Bar anes and G . Di ngerkus (MS) plan to reclass i fy i t in the H em igaleidae.

Order RAJIFORMESFam i ly DASYATIDIDAEDasyat is sp . Sighted by the sen ior author at Enewetak .Taen iu ra melanosp i los B leeker , 1 853 : Randal l , 1 980a.

Taen iu ra brocki Schultz , 1 953 .

Urogymnus sp . A specimen co l lected in the Enewetak Lagoon is under study by the sen ior author .

Fam i ly MYL IOBAT IDAEAetobatus mar inar i (Euphrasen , Randal l , 1 986 (after Strasburg).Fami ly MOBUL IDAEMan ta olf r edi (Kr efft , Schu ltz , 1953 .

Class OSTEICH TH YESSubclass .

ACT INOPTERYGII‘Or der ELOPIFORMESFam i ly ALBUL IDAEAlbu la g los sodon to (For sskal , Randal l , 1 986 . Shak lee and Tamaru ( 1 98 1 ) have shown that there are two

species of A lbu la in the H awai ian Is lands (and elsewhere in the Indo-Pacific r egion), neither of which is A . vu lpes (L innaeus) ,now known to be r es tr icted to the At lantic .

Or der ANGUILL IFORMESFam i ly MOR INGU IDAEMo r ingua fer r ugi nea B l iss , 1 883 .

Mor ingua macrocephala Schultz , 1 953 .

Mo r ingua macr ochi r Schu ltz , 1 953 . Gos l ine and Strasburg ( 1956) showed the remarkable d i fferences of Immatur eH awai ian Mor ingua from adults and between the sexes (including ver tebral counts ) , as have Cast le and 5de (1976)for the western At lantic Mor ingua edwa rds i (Jordan and Ba il inan) . Gos l ine and Strasburg identified thei r H awai ianmater ial as M . macrochi r B leeker . Cast le (1 968a) bel ieves the H awai ian species (and thus pr esumably the specimensfrom the Mar shal l Is lands identified by Schultz as macrochir ) is M . fer r ugineo. Also , Cas tle regards M . macrocephala(B leeker ) as the immatur e form of macrochi r . (Th i s table cont inued on next page.)

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292 RANDALL AND RANDALL

TABLE 1 (con t’d)Fam i ly MURAEN IDAE (cont ’d)U r op teryg iu s conco lor R i i ppell , 1 837 : Schultz , 1953 . Schultz ’s identi fi cat ion should be confi rmed .

U rop teryg ius fuscogu ttatus Schultz , 1 953 .

U rOp terygiu s komar McCosker and Randal l , 1 977: Randal l , 1986 .

U r op teryg iu s macrocephalus (B leeker ,U rOp terygius reidi Schultz , 1 953 . Gos l ine (1 958) placed U . r eidei Schultz (1 943 ) in the synonymy of U . kn ight i (Jordanand Starks ) .U rop terygius kn ight i Schultz , 1 953 . McCosker et al . (1 984) have r eferr ed U . kn ig ht i and U . necturus (Jordan and Gi lber t)to the synonymy of U . macrocephalus , now known to be a tr ans -Pacific species .

U rop teryg iu s marmoratus (Lacepede, Schultz , 1953 .

U r Op terygius s up rafor atu s (Regan , Schu ltz , 1953 .

U rop terygiu s den tatus Schu ltz , 1 953 , from Johnston Is land , was synonym i zed by Gos l ineU rOp teryg ius xan thop ter u s B leeker , 1859 : Schu ltz , 1 953 .

Fam i ly CONGR IDAEA r iosoma scheelei (St romman ,

A r iosoma obud Schultz , 1 953 . P . H . J . Cas tle (per sonal commun ication ) wrote us , i t seem s h ighly l ikely that A . obudH er r e is a jun io r synonym of A . s cheelei , a species wi th low ver tebral counts . Three Bi shop Museum specimens fr omEnewetak have 1 1 6 to 1 20 ver tebr ae (low for the genus —

see Castle, l 968b) .

Conger cinereus cinereus R ii ppell , 1828 .

Conger noo rdz ieki i Schu ltz , 1953 . Kanazawa (1958) placed C . noo rdz iek i i B leeker in the synonymy of cinereus .

Gorgas ia sp . An apparent undescr ibed species ; 1 9 specimens fr om Enewetak , wi th 1 8 1 to 1 89 ver tebrae.

H eteroconger has s i (Klausew i tz and Eibl-Eibes feldt , Bahlke and Randal l , 1981 ; Randall , 1986 .

Taen ioconger hass i Matsuura, 1 984 .

Poec i loconger fasc iatus G’

Lin ther , 1 87 1 : Randal l , 1 986 .

Fam i ly OPH ICH TH IDAEAp ter ichtus klaz ingai (Weber , Randal l , 1 986 . McCosker (1 977) has shown that th is gener i c name r eplacesVermo . A lso some species of Ap ter ichtu s have been m is taken ly descr ibed in Sphogebr anchus and Caecu la .

B r achysomophis sou rop s is Schu ltz , 1 953 .

Callechelys marmoratus (B leeker , Schultz , 1953 .

Callechelys melonotoen io B leeker , 1 864 : Schultz , 1953 .

Lep tenchelys p innocep s Schu ltz , 1 953 . McCosker ( 1970) stated that th is taxon is a jun ior synonym of C . melonotoen io .

Ci r r icoecu la john son i Schultz , 1953 .

Leiur anus sem icinctus (Lay and Bennett , Schu ltz , 1 953 .

M u raen ichthys gymnotus B leeker , 1 864 : Schu ltz , 1 953 .

M u raen ichthys lat icaudata (Og i lby , Schultz , 1 953 .

M u raen ichthys macr opterus B leeker , 1857 : Schultz , 1 953 .

Mu r aen ichthys s chu ltz ei B leeker , 1 857 : Schu ltz , 1 953 .

Mu r aen ichthys s ibogae Weber and de Beaufor t , 1 916 : Schultz , 1 953 .

Myr ichthys bleeker i Gos l ine, 195 1 .

Myr ichthys sem ic inctus Schultz , 1 953 . Gos l ine ( 1951 ) prov ided a new name for Oph is u rus fasciatus var . sem icinctus

B leeker , preoccupied by O . sem icinctu s Lay and Bennett .

Myr ichthys co lubr inas (Boddaer t , Schultz , 1953 .

Myr ichthys macu losus (Cuv ier , Schu ltz , 1 953 .

Myr oph is u r 0 p ter us (Temm inck and Schlegel , Schultz , 1 953 .

Ophichthus sp . Under s tudy by John E . McCosker .

Phyllophichthu s xenodon tus Gos l ine, 1 951 : Randal l , 1 986 (after Strasburg) .Schismor hynchus labial is (Seale,

Lep tenchelys labia l is Schultz , 1 953 . McCosker ( 1 970) descr ibed a new genus , Schism orhynchus , for labia lis .

Schu ltz idio johns tonens is Schultz and Woods , 1 949 : Schu ltz , 1 953 .

Schu ltz idio ret rOp inn is (Fowler , Schultz , 1 953 .

Order CLUPEIFORMESFam i ly CLUPEIDAED uss um ier ia has selt i B leeker , 1 850 : Schultz and Welander , 1 953 .

H er klots ichthys quadr imacu latus (R ijppell , Randal l , 1 986 [after Str asbur g , who identi fied th is clupeid fr om Ar no

as H arengu lo kunzei

Sp ratello ides ot rofasc iotus Schu ltz , 1 953 .

Sp r atel lo ides del icatu lus (Bennett , Schultz and Welander , 1953 .

O rder SALMON IFORMESFam i ly SYNODONT IDAESou r ida g raci l is (Quoy and Gaimard, Schultz , 1 953 .

SW OdU S bino tatu s SChU ltZl 1 953(Th is table cont inued on next page

ANNOTATE D CHECKLIST OF F ISHES 293

TABLE 1 (con t’d)

Fam i ly SYNODONT IDAE (cont ’d)Synodus eng lemon i Schultz , 1953 . Cressey ( 198 1 ) r ev ised Synodus ; he r ecogn i zed th is species and S . binototus as val id .

Synodus var iegatus (Lacepede, Schultz , 1 953 .

Synodus sp .

Order GONORYNCH IFORMESFam i ly CH AN IDAEChonos Chonos (For sskal , Randal l , 1 986 .

Or der GOBIESOCIFORMESFam i ly GOB IESOCIDAEL iobranchia s tr io Br iggs , 1 955 : Schu ltz, 1 966 .

Or der LOPH IIFORMESFam i ly ANTENNAR IIDAEAn ten nar ius anolis (Gos l ine, Randal l , 1 986 .

An tennar ius coccineus (Lesson , Schultz, 1 966 .

An tennar ius p ictus (Shaw and Nodder , Randal l , 1 986 . Theodore W . Pietsch (per sonal commun ication)informed u s that A . p ictus i s a sen ior synonym of A . chi ronectes Lacepede (the name used by most r ecent author s forth is species ) .An tennar ius r andoll i A l len , 1 970 : Randal l , 1 986 .

An tennar ius rosaceus (Sm i th and Radcl i ffe,

Tr ichophryne rosaceus Schultz , 1 966 .

An tennatus tuberosus (Cuvier , Randal l , 1986 .

An tennar ius bigibbus Fowler , 1 928 .

Order GAD IFORMESFam i ly BREGMACEROT IDAEB regmacer os nectoban us Whi tley , 1941 .

B regmaceros mcclellandi Schultz , 1 953 . Mar shal l Is lands mater ial (including a B ishop Museum specimen fr om the

Enewetak Lagoon) was r eidentified as B . nectobanus , fo l lowing D ’

Ancona and Cavinato

Order OPH IDIIFORMESFam i ly OPH ID IIDAEB r otu la townsendi Fowler , 1 900 : Schultz , 1960 .

Fam i ly BYTH ITIDAEB rosmOphyciops pou tz kei Schultz , 1960 . Cohen and N ielson ( 1 978) are fo l lowed in cons ider ing Byt hi t idae a fam i ly forth is and r elated fi shes .

D inemat ichthys i luocoeteo ides B leeker , 1 855 : Schultz , 1960 . A badly needed r ev is ion of the genus D inemat ichthys and

al l ies has just been commenced by A l legra N . Sedor of the Un iver s i ty of Souther n Ca l i for n ia. More than one speciesof the genus occur s in the Mar shal l Is lands ; i t is not clear that the name i luocoeteoides can be confidently appl iedto any of them .

Fam i ly CARAPIDAECorap us homei (R ichardson , Schu ltz , 1960 .

Carapus mou r lan i (Peti t, Schu ltz , 1960 . Schul tz identi fied his mater ial as C. mou r lan i wi th cons iderableuncer tainty ," though po inted ou t that his specimens ar e clear ly distinct fr om his C. homei (wh ich Ar no ld, 1 956 , l istedas a sen ior synonym of mour lan i) .

Jordan icus graci lis (B leeker , Schultz , 1 960 .

Order ATH ERINIFORMESFam i ly BELON IDAEAblennes h ians (Valenciennes , Schultz , 1 953 .

Platybelone argalus p latyu ra (Bennett ,Be lone p latyu ra Schultz , 1 953 .

St rongylu ro incisa Schultz , 1 953 .

Rhophiobelone robus ta Schultz, 1953 . Mees ( 1962) and Par in (1967) placed th is taxon in the synonymy of S. incisa.

Ty losur us crocod i lus crocod i lus (Peron and Lesueur , 1 821 )Fam i ly H EMIRAMPH IDAEEu lep tor hamphus vi r idis (van H asselt , Schultz , 1953 .

H ypo r homphus acutus acutus (G i i n ther , Co l lette, 1 974 .

H ypo r homphus acutus Schultz , 1 953 .

H ypor homphus affin is (G i i n ther , Par in et al . , 1980 .

H ypo r homphus duss um ier i Schu ltz , 1 953 .

H ypo r homphus dus s um ier i (Valenciennes , Par in et al . , 1 980 .

H ypo r homphus lat icep s Schultz , 1 953 .

Oxypor homphus m icropterus m icropter us Par in et al . , 1 980 .

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294 RANDALL AND RANDALL

TABLE 1 (con t'd)‘Fam i ly EXOCOET IDAE

Cyp selu r us an toncich i Woods and Schu ltz , 1953 .

Cyp selu r us sp i lonotop ter us (B leeker , Woods and Schultz , 1953 .

Cyp selu r us un ico lor (Valenciennes , Woods and Schu ltz , 1953 .

Exocoetus vo l itan s L innaeus , 1 758 : Woods and Schultz , 1953 .

Parexocoetus men to men to (Valenciennes ,Parexocoetu s men to Woods and Schultz , 1953 .

P r ogn ichthys sp . Woods and Schu ltz , 1 953 .

Fam i ly ATH ER IN IDAEA ther inomor us lacunos us (B loch and Schneider ,P ranes us p ingu is Schultz , 1 953 . Wh itehead and Ivan tsoff (1 983) placed A ther ino p ingu is Lacepede in the synonym y ofA ther inomor us lacunos us .

A ther ion elymus Jordan and Starks , 1 90 1 .

A ther ion e lymus asper Schu ltz , 1 953 . Walter Ivant soff (per sonal commun ication) does not recogn i ze Schu ltz ’s subspeciesof A . e lymus .

H ypoather ino bar nes i Schultz , 1 953 .

H ypoather ino ovolaua (H er r e,

A llanetto ovolaua Schu ltz , 1 953 . Gener ic change fr om Wh itehead and Ivan tsoff , 1 983 .

Stenother ina panatela (Jordan and R ichardson ,

Stenother ina temm incki i Schultz , 1 953 . Walter Ivan tsoff (per sonal commun ication ) has corr ected th is m i s identi fi cationby Schu ltz .

Fam i ly ISON IDAEIso hawai iens is Gos l ine, 1 952 : Randal l , 1 986 .

Order BERYCIFORMESFam i ly ANOMALOPIDAEPhotoblephar on palpebr atus (Boddaer t , Randal l , 1 986 .

Fam i ly H OLOCENTRIDAEMyr ip r is t is odus ta B leeker , 1 853 : Randal l , 1 986 .

Myr ip r is t is amoena (Castelnau ,Myr ip r is t is argyr om u s Woods , 1953 . Greenfield ( 1974) placed M . argyr om us Jordan and Evermann in the synonym y ofM . amoen us (Cas telnau) .

Myr ip r is t is ber ndt i Jordan and Evermann , 1 903 : Woods , 1953 .

Myr ip r is t is kun tee Cuv ier , 1 83 1 .

Myr ip r is t is m u lt i radiatu s Woods , 1 953 . Green field ( 1 974) s ynonym i zed M . m u lt i radiatus G ijn ther wi th M . kun tee.

Myr ip r is t is mu rdjan (For s skal , Woods , 1953 .

Myr ip r is t is bowd i tchoe Woods , 1 953 . Randal l and Gueze( 1 98 1 ) placed th is species , M . par videns Cuv ier , andM . ax i l lar is Valenciennes in the synonymy of M . m u rdjon .

Myr ip r is t is p ral in ia Cuv ier , 1 829 : Woods , 1 953 .

Myr ip r is t is vio lacea B leeker , 1 851 .

Myr ip r is t is m icr ophthalmus Woods , 1 953 . G r eenfield ( 1974) synonym i zed M . m icr ophthalmus B leeker wi th vio lacea.

Myr ip r is tis vittata Cuv ier , 1 83 1 : Randal l , 1 986 .

Neon iphon orgen teus (Valenciennes ,“H o locen t rus laevis Woods , 1953 . Sh im i zu and Yamakawa (1979) placed H o locen t rus laeve Gunther in the synonym y oforgen teus . Randal l and H eems tra ( 1 985) have shown that Neon iphon Castelnau replaces the gener ic nameF lommeo Jordan and Evermann .

N eon iphon oper cu lar is (Valenciennes ,H o locen t r u s oper cu lar is Woods , 1 953 .

Neon iphon sommaro (For sskal ,H o locen t r u s sommaro Woods , 1 953 .

Plect rypop s l ima (Valenciennes ,H olo t r ochys l im o Woods , 1953 . Woods and Sonoda ( 1 973 ) showed that H olot rochys Gii nther is a gener ic synonym of

Plect rypop s G i ll .Sargocen t r on coudimocu lotum (Ruppell , Randal l , 1 986 . Matsuura and Shim izu ( 1 982) noted thatSargocen t ron Fowler is an o lder name than Adio ryx Starks .

Sargocen t r on diadema (Lacepede,

H o locen t rus diadema Woods , 1 953 .

‘Par in (1 960 ; trans lation 1 963 ) l i sted n ine species of Cyp selu r u s , in addition to the three included here,three of P rogn ichthys , and

Parexocoetus brachyp ter u s brachyp ter us fr om Ocean ia. Poss ibly some of these species wi ll eventual ly be found in the Mar shall s area . Ran

dal l ( 1955a) repo r ted Cyp selu r u s s u tton i and P rogn ichthys albimacu latus fr om the nearby Gi lber t Is lands .

(Th is table continued on next page. )

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TABLE 1 (con t’d)

Fam i ly SCORPAEN IDAE (cont ’d)Ptero is an tennata (B loch , Schultz , 1 966 .

Pter ois radiata Cuv ier , 1 829: Schu ltz , 1 966 .

Pter ois vo l iton s (L innaeus , Kendal l and Go ldsbor ough , 19 1 1 ; A l len and Eschmeyer , 1 973 .

Sco rpaenodes br ocki (Schultz ,H ypomocr us brocki Schu ltz , 1 966 .

Scorpaenodes guamen s is (Quay and Gaimard, Schultz , 1 966 .

Scorpaenodes h i r sutus Sm ith , 1957: Randal l , 1 986 .

Scorpaenodes kelloggi (Jenk ins , Schu ltz , 1 966 .

Scorpaenodes por vip inn is (Garr ett , Schultz , 1 966 .

Scor paenop s is diabo lus (Cuv ier ,Scorpaenop s is gibbosa Schultz , 1 966 . Eschmeyer and Randall (1 975) s tated that the tr ue 8 . gibbosa appear s to beres tr icted to the Indian Ocean .

Scorpaenop s is fowler i (Pietschmann , Eschmeyer and Randal l , 1975 . Th is smal l species is on ly tentatively placed inthe genus Scorpaenop s is .

Scorpaenop s is sp . A new species wh ich wi l l be descr i bed by W i ll iam N . Eschmeyer and Kaza Rama Rao .

Sebas top is tes cyonos t igma (B leeker ,Sebas top is tes olbobr u nnea Schu ltz , 1 966 . Eschmeyer and Rama Rao (MS) have shown that Scorpaena olbobr u nnea

G i i n ther is a jun ior synonym of cyonos t igmo.

Sebas top is tes maur i t iana (Cuv ier ,Sco rpaena cor all icola Schu ltz , 1 966 . Eschmeyer and Randal l (1 975) placed 8 . corall icola (Jenk ins ) in the synonymy ofS . ball ieu i (Sauvage) . Eschmeyer and Rama Rao (MS) r egard Sebas top is tes ball ieu i as a H awai ian endem ic . The closelyr elated S. maur it iana is a w ide-

r ang ing Indo Paci fic species that does not occur in H awai i .Sebas top is tes s t rongio (Cuv ier ,Sebas top is tes bynoen s is Schultz , 1 966 . Sebas top is tes bynoens is R ichardson is a jun ior synonym of Parascorpaena p icta (Cuvi(Eschmeyer and Rama Rao , MS) . The B ik in i specimen identified as S . bynoens is by Schu ltz is s t rongio .

Synanceia ver rucosa B loch and Schneider , 1 80 1 : Schu ltz , 1 966 .

Taen ionotus tr iacon thus Lacepede, 1 802 : Schultz , 1 966 .

Fam i ly CARACANTH IDAECar ocon thu s macu latus (Gray , Schultz , 1 966 .

Car ocon thus un ip in no (Gr ay ,Car ocon thus un ip innus Schu ltz , 1 966 .

Fam i ly APLOACTINIDAECocot ropus sp . An undescr ibed species which wi l l be named by Stuar t G . Pos s .

Fam i ly PLATYCEPH AL IDAEThysanophrys oren icolo Schultz , 1966 .

Thysanophrys ch i ltonae Schultz , 1 966 .

Thysanophrys malayanus (B leeker ,Thysanophrys pap i llo labium Schultz , 1 966 . Les l ie W . Knapp (per sonal commun ication) regards th is species as a jun iorsynonym of malayanus .

Wak iyus we lander i Schu ltz , 1 966 .

Order PEGASIFORMESFam i ly PEGASIDAEEu rypegas us dracon is (L innaeus , Randal l , 1 986 .

O rder PERCIFORMESFam i ly SERRAN IDAEAn th ias bor t lettor um Randal l and Lubbock , 1 981 .

An th ias bico lor Randal l , 1 979 .

An th ias dispar (H er re, Randal l and Lubbock , 198 1 .

An th ias pascalus (Jordan and Tanaka, Randal l and Lubbock , 198 1 .

An th ias p leu rotaen io B leeker , 1 857 : Randal l , 1 986 .

An th ias randoll i Lubbock and A l len , 1978 .

An th ias sm i thvan iz i Randal l and Lubbock , 1 98 1 .

An th ias ven t r al is ven t r a lis Randal l , 1 979 .

Anyperodon leucog r omm icu s (Valenciennes , Schultz , 1953 .

Be lonoper ca chabanaudi Fowler and Bean , 1 930 : Schultz , 1953 .

Cephalophol is argus (B loch and Schneider , Schultz , 1 953 . Randal l and Ben-Tuv ia (1 983) have shown thatCephalophol is guttatus (Bloch) i s an ear l ier name for this species , but they have peti tioned the Inter nat ionalComm i s s ion for Zoo log ical Nomenclature to suppr ess C . guttatus (B loch) in order to retain the long-

establ ished C . argus .

Cephalopho l is leopar du s (Lacepede, Schultz , 1 953 .

Cephalopho lis m in iotus (For s skal , Schultz , 1 953 .

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ANNOTATED CHECKLIST OF FISHES 297

TABLE 1 (con t’d)

Fam i ly SERRAN IDAECephalophol is sexmacu loto (R i i ppell , Randal l and Ben-Tuvia, 1 983 .

CephoIOphol is sonnerati (Valenciennes , Randal l , 1 986 .

Cephalophol is sp i loparaeo (Valenciennes ,Cephalophol is anolis Randal l , 1986 .

Cephalophol is u rodeto (B loch and Schneider ,Cephalophol is urode las Schultz , 1953 .

Ep inephelus caer u leop unctatus (B loch , Kendal l and Go lds bo rough , 1 91 1 ; Randal l , 1 986 .

Ep inephelus cyonOpodus (R ichardson ,

Ser ranus flavocaer u leus H iyama, 1943 . Many author s have m is taken ly r egarded E . flovocaer u leus (La cepede) as asynonym of E . hoedt i i (cyonopodus) .

Ep inephelus kohler i Schu ltz , 1953 . Katayama (1 960) and Randal l (1 980a) placed th is species in the synonymy ofE . hoedti i . Ep inephelus cyonopodus is a sti l l o lder name for the species (Randal l and Wh itehead ,

Ep inephelus fasciotus (For sskal ,Ep inephelus emoryi Schultz , 1953 . Katayama (1960) cor r ectly r efer red emo ryi to the synonymy of fasciotus .

Ep inephelus fuscogu ttatus (For sskal , Schultz (1 953 ) confused two species under the heading E . fuscogu ttatus .

H i s i l lustr ation (P late 26 A) is E . m icrodon . Randal l (1964a) dis tingu i shed these two related species .

Ep inephelus hexagonotus (B loch and Schneider , Schultz , 1 953 .

Ep inephelus lanceo latus (Bloch, Schultz , 1 966 .

Ep inephelus macrosp i los (B leeker , Ep inephelus macr osp i los Schu ltz, 1 953 .

Ser r anus howlandi G ii nther , 1 883 and Ep inephelus sp i lotus Schultz , 1 953 ar e synonyms . Randal l (1 955a)er r oneous ly r egarded E . sp i lotus as a synonym of E . corallicola (Valenciennes ) .

Ep inephelus macu latus (B loch ,Ep inephe lus medu r ens is Schultz , 1953 . Randal l ( l 98oa) r egarded E . medu rens is (Gunther ) as a synonym of

E macu latus .

Ep inephe lus melonos t igmo Schultz , 1 953 . Randal l (1955a) prov i s ional ly placed th is species in the synonym y ofE . hexagonotus . It is now known to be val id .

Ep inephelus mer ra B loch , 1 793 : Schultz , 1 953 .

Ep inephelus m icrodon (B leeker ,Ep inephelus fuscogu ttatus Schultz , 1953 (in part ) .Ep inephe lus hor r idus Schultz , 1 966 . Ser ranus hor r idus Valenciennes is a jun ior synonym of E . fuscoguttotus (For sskal) .

Ep inephelus ongus (B loch ,Ep inephelus s ummona Schultz , 1 953 . Randal l and Ben-Tuv ia (1 983 ) have shown that E . s ummona (For sskal) is endem icto the Red Sea. The r elated E . angu s occur s elsewhere in the lndo Pacif ic .

Ep inephelus soc ial is (G ii nther , Schultz , 1953 . Randal l (1980a) m is taken ly used a photo of E . ongu s fr om Ponape toi l lus trate E . soc ial is .

Ep inephelus sp i lotocep s Schultz, 1 953 . Randal l ( 1 955a) was in er r or refer r ing th is species to the synonymy ofE . hexagonotus .

Ep inephelus touvino (For sské l ,Ep inephelus elongatus Schultz , 1 953 . Katayama (1 960) and Randal l (1 964a) r egarded th i s species as a synonym of

E . touvino .

G r oci la albomargi nata (Fowler and Bean , Randal l , 1 986 .

G ramm is tes sex l ineatus (Thunber g , Schultz , 1953 .

G r amm is tops ocellatus Schultz , 1 953 .

L iop r opomo pal lidum (Fowler ,Yp s igr amma pa l lida Schultz , 1 953 .

L iop rOpomo sasum i (Jordan and Seals ,

Yp s igr amma l ineata Schu ltz , 1 953 . Randal l ( l 955a) refer r ed th is species to the synonymy of L iop r opomo sus am i .Yps igramma br ocki Schultz , 1 953 . Randal l (1 955a) stated that Y. br ock i i s a probable synonym of L iop ropomo sus am i .Th is fi sh is also a poss ible hybr id of L iOp rOpoma s usum i and L . pa l l idum ; i t is sti l l under study .

L iop r opomo sp . A smal l species with two br oad r ed s tr ipes on s ide of body wh ich extend into caudal fin ; i t wi l l bedescr ibed by Randal l and Tay lor who ar e r ev is ing the genus .

L uzon ichthys wo itei (Fowler ,L uzon ichthys robus tus Fou rmano i r , 1 977 . Randal l (1 98 1 b) regarded L . r obus tus as a probable synonym of L . woi tei .

P lect ron thios fou rmanoi r i Randal l , 1 980 .

P lect ron thios nanus Randal l , 1 980 .

P lect r opomus laevis (Lacepede,

Plect ropomus leopardus Schultz , 1 953 (non -Lacepede) .Plect r opomus melanoleucus Randal l , l 980a. H oese et al . , 1 98 1 , suspected that P . laevis (Lacepede) wou ld have pr ior i tyover P . melanoleucus (Lacepede) ; th is has been confi rmed (Randal l and H oese, in press ) . What Schultz ca l led P . leopardus is

the mos t common co lor phase of thl s spec1es .

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TABLE 1 (con t’d)

Fam i ly SERRAN IDAE (cont’d)Plect ropom us ol igacon thus (B leeker , H iyama, 1 943 ; Randal l , 1 986 .

Plect r opom us areo latus (Ruppell ,P lect ropom us t r uncatus Schultz . Randall and H oese (in pr ess) have shown that P . t r uncat us Fowler is a jun ior synonym of

P . ar eo lat us .

Pogonoper ca punctata (Valenciennes , Randall , 1 986 .

Pseudogr amma bi linear is (Schultz ,Apo r op s bi l inear is Schultz , 1953 . Randal l (1 955a) synonym i zed the genus Apo rop s with Pseudog r ammo .

Pseudogr amma po lyacan tha (B leeker , Schu ltz , 1 953 .

Pseudogr amma sp . A new species to be descr ibed by J . Randal l who plans a r ev is ion of the genus .

Var io la lout i (For sské l , Schultz , 1 953 .

Fam i ly PSEUDOCH ROMIDAEPseudochr om is ou r eo mar shallens is Schultz , 1 953 .

Pseudochrom is tapeinosomo B leeker , 1 853 : Schultz , 1 953 .

Pseudop les iop s revellei Schu ltz , 1 953 .

Pseudop les iop s r osoe Schultz , 1 953 .

Pseudop les iop s sorgen t i Schultz , 1 953 .

Pseudop les iop s sp . One specimen from Kwajalein .

Fam i ly ACANTH OCLINIDAEAcon thop les iop s h iatt i Schultz , 1953 .

Fam i ly PLESIOPIDAECollop les iop s alt ivel is (Steindachner , Randal l , 1 986 .

P les iop s coer u leol ineatus R ii ppell , 1 835 .

Ples iop s me los Schu ltz , 1 953 . Schultz ( 1 966 , addendum ) (after Inger 1955) r eiden ti fi ed hi s Mar shall Is lands mater ialas P . coer u leol ineatus .

Ples iop s corall icola B leeker , 1 853 .

P les iop s n igr icans Schultz , 1953 . Inger ( 1 955) has shown , fr om mater ial avai lable to him for his r evi s ion of P les iops ,

that the tr ue P . n igr icans (Ruppell) is endem ic to the Red Sea.

Fam i ly CIRRH ITIDAEAmblyc i r r hi tus bimacu lo (Jenkins ,Ci r r hi to idea bimacu la Schultz , 1960 .

Ci r rh it ichthys oxycephalus (B leeker , Gunther , 1 873 ; Randall , 1 963a.

Ci r rh i tus p innu latus (B loch and Schneider , Schultz , 1 960 .

Isoc i r r hi tus sexfosciotus (Schultz ,Ci r r hi to idea sexfasciato Schultz , 1 960 . Randall ( 1 963a) cr eated the monotypic genus Isoc i r r hi tu s for th is species .

Oxyc i r r h ites typus B leeker , 1 857 : Randal l , 1 986 .

Paraci r r hites or cotu s (Cuv ier ,Gymnoc i r r hites or cotu s Schultz , 1 960 .

Paraci r r hites for s ter i (B loch and Schneider , Schultz , 1 960 .

Paraci r r hi tes hem is t ictus (G ii n ther , Schu ltz , 1 960 .

Fam i ly KUHL IIDAEKuhl ia margi nata (Cuv ier , Schul tz , 1 953 .

Kuhl ia m ug i l (Bloch and Schneider ,Kuhlia taen iu ra Schul tz , 1 953 . Randal l ( 1 973 ) showed that K . mugi l is an ear l ier name for K . taen iura (Cuv ier ) .

Fam i ly PRIACANTH IDAEH eter op r iacon thus cr uen tatus (Lacepede,

Pr iacan thus cr uen tatus Schultz , 1953 .

Pr i acan thus hom r u r (For ss kal , Schultz , 1 953 .

Fam i ly APOGON IDAEApogon coccineus Ruppell , 1838 .

Apogon er ythr inus Lachner , 1 953 . Sm i th (1 96 1 ) could find no reason for maintain ing A erythr i nus Snyder as adist inct species .

Apogon cyonosoma B leeker , 1 853 .

Apogon novae gu ineae Lachner , 1 953 . The author and Er n es t A . Lachner have determ ined that the specimenstentat ively ident i fied by Lachner as A . novaegu ineae appear to be A . cyonosoma. Marshal l Is lands specimens di fferfr om typ ica l cyonosomo in having yellow instead of o range s tr ipes .

Apogon doryssa (Jo rdan and Seale, Randall , 1 986 .

Apogon el l io tt Day , 1 875 : Randa ll , 1 986 (after Strasburg) .Apogon everman n i Jordan and Snyder , 1 904 : Randal l and Bii hlke, 1 98 1 .

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TABLE 1 (con t’d)Fam i ly

_

CARANG IDAE (cont ’d)Corango ides fer dou (For s skal ,Cor ango ides g i lber t i Woods , 1 953 . W i l l iam F . Sm i th-Van i z (per sonal commun ication) has determ ined that the trueferdou is the species some r ecent author s such as Woods have labe l led gi lber t i , and the species usual ly identi fied asC . ferdou jordan i N icho ls is C. or thogr amm us (Jo rdan and G i lber t) .

Corangoides or thog ramm us (Jordan and Gi l ber t ,Corango ides ferdou jordan i Woods , 1953 .

Corangoides p lagiotoen ia B leeker , 1 857 : Randal l , 1 986 .

Caranx ignobi l is (For sskal , Randal l , 1 980a.

Car anx lugubr is Poey , 1 861 : Woods , 1 953 .

Car anx melampygus Cuv ier , 1 833 : Woods , 1 953 .

Car anx sexfasciotus Quoy and Gaimard, 1 825 : Woods , 1 953 .

Decapterus m u r oads i (Temm inck and Schlegel , Woods , 1 953 .

E lagat is bip innu loto (Quoy and Ga imard, Woods , 1953 .

G nathanodon Speciosus (For s skal , Schultz , 1 966 , addendum ; Randal l , 1986 (after Str asburg) .Scombe r o ides lyson (Fo r sskal ,Scombe r o ides sanct i -petr i Woods , 1 953 . Sm i th-Van i z and Staiger (1973) have shown that S . lyson is an ear l ier namefor th is species than S. s onct ipet r i (Cuv ier ) .

Se lor cr umenophtholmu s (B loch ,Tr ochu rop s cr umenophtholm us Woods , 1 953 .

Trach inotus boi llon i (Lacepede, Woods , 1 953 .

Trach inotus blochi i (Lacepede,Randal l , 1 986 .

Fam i ly CORYPH AEN IDAECor yphaena hipp u r us L innaeus , 1 758 : Schu ltz , 1953 .

Fam i ly LUTJAN IDAEAphareus furco (Lacepede,

Aphareus fu r cotus Schultz , 1 953 .

Ap r ion v i r escens Valenciennes , 1 830 : Schultz , 1 953 .

L u tjon us bohar (For sskal , Schultz , 1 953 .

Lutjanus fu lvus (B loch and Schneider , Randal l , 1 986 (after Str asburg) .Lutjanus marginatus Kendal l and Go ldsbo r ough , 1 91 1 .

Lutjanus flavipes H iyama, 1 943 . Randal l ( 1 973) noted that L . fu lvus pr edates L . vo igiens is (Quay and Gaimard) andL . marginatus (Cuv ier ) , the names usual ly given to th i s common lu tjan id .

Lutjanus g ibbus (For s skal , Schultz , 1953 .

An th ias her aldi Schultz , 1953 . H eem s tra ( 1972) showed that th i s taxon is a juven i le L . gi bbus .

Lutjanus kosm i ra (For s skal , Schu ltz , 1 953 .

Lutjanus monos t igma (Cuv ier ,Lutjan us m onos t igm us Schultz , 1953 .

Lutjanus sem icinctus Quoy and Gaimard, 1 824 : H iyama, 1943 ; Randal l , 1986 .

Lutjan us vitta (Quay and Gaimard, Randal l , 1 986 (after Str asburg) .Lutjanus sem icinctus Quoy and Gaimard, 1 824 : H iyama, 1 943 ; Randal l , 1 986 .

Lutjanus vi tta (Quoy and Gaimard, Randal l , 1 986 (after Strasburg) .Maco lor n iger (For s skal , Schultz , 1953 .

Fami ly CAESIONIDAECoes io caer u lau r ea Lacepede, 1 80 1 : Randal l , 1 986 .

Coes io ter es Seale, 1 906 .

Coes io xan thonotus Schultz , 1 953 .

P ter ocoes io mar r i Schultz , 1 953 .

Pterocaes io kohler i Schu ltz , 1 953 . Kent E . Ca rpenter , who is r ev is ing the fam i ly , be l ieves thi s is the same species asP. mar r i .

Pter ocoes io t i le (Cuv ier , Schultz , 1953 .

Pterocaes io sp .

Fam i ly H AEMUL IDAEP lecto rh inchus obscu r us (Gunther , Randal l , 1986 .

Plector hinchus p icus (Cuv ier , Randal l , 1 986 .

Fam i ly LETH RIN IDAEG nathoden tex au reolineatus (Lacepede, Schultz , 1 953 .

Gym noc ron ius m l OdO"(B leeker , Randal l , 1 986 (after Str asbur g) used the name m icrodon for the Mar shallIs lands Gym noc r on ius w ith reservations . The genus Gymnocr on ius needs r ev is ion .

Lethr inus ambo inen s is B leeker , 1 854 : Randa l l , 1 980a (after Sato ,

L the r i n us var iegatus SChU ltZ'1 953 .

(Th i s table cont inued on next page. )

ANNOTATED CHECKLIST OF F ISH ES 30 1

TABLE 1 (con t’d)

Fam i ly LETH RIN IDAE (cont ’d)?Lethr in us r et icu latus Schultz , 1 953 . Schultz prov ided no i l lustrations of any of the five species of Lethr inu s he

repor ted fr om the Mar shal l Is lands , and Sato (1 978) made reference to on ly two of them , kollop ter us and m in iatus

(e longatus) . The sma l l specimen s ident i fied as r et icu latus by Schu ltz are on ly tentatively r egarded as ambo inens isher e.

Lethr inus e longatus Valenciennes , 1 830 .

Lethr in us m in iatu s Schultz , 1 953 . Randal l and Wheeler (MS) have determ ined that the true L . m in iatu s (B loch andSchneider ) is a species most author s have ident ified as L . chrysos tomus R ichardson .

Lethr inus hor ak (For s ské l , Schultz , 1 953 .

Lethr in us kollop ter us B leeker , 1856 : Schultz, 1 953 .

Lethr inus r amak (For sskal , Randal l , 1 986 .

Lethr in us xan thochi lus Klunz inger , 1870 : Randal l , 1 980a .

Lethr inus m icrodon Schultz 1 953 .

Monotox is gr andocu l is (For s skal ,Fam i ly NEMIPTERIDAEPen topodus con in us (Cuv ier , Randal l , 1 986 .

Scolop s is lineatus Quay and Gaimard, 1 824 .

Scolops is cancel latus Str asburg , 1 953 (m imeo r epor t to ONR) ; H iatt and Strasburg , 1 960 . Bar r y C . Rus sel l (per sonalcommun ication) has informed us that S . lineatus is an o lder name for th is species than S . cancellatus (Cuv ier ) .

Fam i ly GERREIDAEGer res orgyr eu s (B loch and Schneider , Schultz , 1 953 .

Fam i ly MULL IDAEMu llo ides flavo lineotu s (Lacepede,

Mul lo idichthys samoens is Lachner , 1960 .

M u llo ides von icolen s is (Valenciennes ,Mul lo idichthys ou r iflammo Lachner , 1 960 .

Mu l lo idichthys von icolens is Lachner , 1 960 .

Mu llo ides pfluger i Steindachner , 1 90 1 : Randal l , 1 986 .

Par upeneus bor ber ino ides (B leeker , Randal l , 1 986 .

Upeneus ot r ocingu lotus Kner , 1 870 . Lachner ( 1 960) er r ed in cons ider ing U . ot roc ingu lot us as a synonym of P t r i/osc iatus .

Par upeneus bor ber inu s (Lacepede, Lachner , 1 960 .

Par upeneus bifosciatu s (Lacepede, Randal l , 1 986 .

Par upeneus cras s i labr u s Lachner , 1 960 .

Par upeneus cyclos tomus (Lacepede, Lachner , 1 960 .

Par upeneus hep tocon thus (Lacepede, Randal l , 1 986 .

Par upeneus m u lt ifosciatus (Quoy and Gaimard,

Par upeneus t r ifasciotus Lachner , 1 960 . Randal l (MS) r egards P t r ifosc iotu s (Lacepede) as a synonym of

P . bifasciatus (Lacepede) . The species in the centr al and wes ter n Pacific identified by most author s as t r ifasciotusshould be ca l led P . mu lt ifasc iotus . It is not known fr om the wester n Indian Ocean .

Par upeneus p leuros t igma (Bennett , Lachner , 1 960 .

Upeneus toen iop ter us Cuv ier , 1 829 : Randal l , 1 986 .

Fam i ly PEMPH ERIDIDAEParap r iocan thus berycifo rmes Franz , 1 910 : Schultz , 1953 .

Pempher is oualen s is Cuv ier , 1 83 1 : Schultz , 1 953 .

Fam i ly KYPH OSIDAEKyphosus ciner ascens (For sské l , Schultz , 1953 .

Kyphosus vo ig iens is (Quoy and Gaimard, Randal l , 1 986 .

Fam i ly EPH IPPIDIDAEPlatox or bicular is (For sskal , Randall , 1 986 (after Strasburg) .Fam i ly CH AETODONT IDAEChaetodon aur iga For s skal , 1 775 : Woods , 1953 .

Chaetodon bennett i Cuv ier , 1 83 1 : Burgess , 1978 .

Chaetodon cit r inellus Cuv ier , 1 83 1 : Woods , 1 953 .

Chaetodon eph ipp ium Cuv ier , 1 83 1 : Woods , 1 953 .

Chaetodon klein i i B loch , 1 790 : Woods , 1 953 .

Chaetodon l ineo latus Cuv ier , 1 83 1 : Burgess , 1 978 .

Chaetodon lunu la (Lacepede, Woods , 1953 .

Chaetodon melannotus B loch and Schneider , 1 80 1 : Woods , 1 953 .

Chaetodon mer tens i i Cuv ier , 1 83 1 : Woods , 1953 .

Chaetodon meyer i B loch and Schneider , 1 80 1 : Woods , 1953 .

Chaetodon or nat is s im us Cuv ier 1 83 1 : Woods , 1 953 .

(Th is table cont inued on next page . )

302 RANDALL AND RANDALL

TABLE 1 (con t’d)

Fam i ly CH AETODONT IDAE (cont ’d)Chaetodon punctatoj'asciatu s Cuv ier , 1 83 1 : Woods , 1953 .

Chaetodon quadr imacu latus Gr ay , 1 833 : Woods , 1 953 .

Chaetodon raffles i Bennett , 1 830 : Burges s , 1978 (fr om underwater photo by Scott Johnson , but local i ty Kwajalein ,not

Enewetak) .Chaetodon r et icu latus Cuv ier , 1 83 1 : Woods , 1953 .

Chaetodon semeion B leeker , 1 855 : Woods , 1 953 .

Chaetodon t inker i Schultz , 1 951 : Randal l , 1 986 .

Chaetodon tr ifoscial is Quay and Gaimard, 1825 .

Megap r otodon s t r igangu lus Woods , 1 953 .

Chaetodon tr ifascial is Mungo Par k , 1 797 : Woods , 1 953 .

Chaetodon u l ietens is Cuv ier , 1 83 1 .

Chaetodon [alcu la Woods , 1 953 . Burges s (1978) wr ote that C . falcu la B loch is an Indian Ocean species that is often not

d is tingui shed fr om the Paci fic C . u l ietens is .

Chaetodon un imocu lotus B loch , 1 787 : Woods , 1 953 .

Chaetodon vogabundus L innaeus , 1 758 : Randal l , 1 986 (after Str asburg) .For cip iger flav is s im us Jordan and McG regor , 1 898 .

F or cip iger long i r os t r is Woods , 1 953 . Randal l and Caldwel l (1970) showed that most author s have used the name

longi r os t r is for the mos t common of the two species of the genus (the one wi th the shor ter snout); the cor r ect name forth is bu tt erflyfi sh is F . flavis s im us .

F or c ip iger long i ros t r is (B roussonet , Randal l and Caldwel l , 1970 .

H em itau r ichthys po lylep is (B leeker , Burgess , 1 978 .

H en iochus acum inatus (L innaeus , Fowler , 1928 .

H en iochus ch rysos tomus Cuv ier , 1 83 1 .

H en iochus perm u tatus Woods , 1 953 . Burgess (1978) reidenti fied Woods ’

mater ial of this species fr om the Mar shal lIs lands as H . chr ysos tomus .

H en iochus monocer os Cuv ier , 1 83 1 : Woods , 1 953 .

H en iochus var ias (Cuv ier , Randal l , 1 986 .

Fam i ly POMACANTH IDAECen t ropyge bico lo r (B loch , Randal l , 1986 .

Cen t ropyge bisp inos us (G ii n ther , Woods and Schultz , 1953 .

Cen t ropyge flavis s im us (Cuv ier , Woods and Schultz , 1953 .

Cen t r opyge heraldi Woods and Schu ltz , 1 953 .

Cen t r opyge lor icu lu s (G i i n ther , Randal l , 1986 .

Cen tropyge mu lt ico lor Randal l and Wass , 1974 .

Cen t ropyge mu lt ifasciatus (Sm ith and Radcl i ffe, Randal l , 1 986 .

Cen t ropyge vrol iki i (B leeker , Randal l , 1 986 (after Str asburg) .G en ican thus be l lus Randal l , 1 975 : Randal l , 1 986 .

G en ican thus wotanobei (Yasuda and Tom inaga, Randal l , 1 986 .

Pomacan thus imperator (B loch, Woods and Schu ltz , 1 953 .

Pygop li tes diacan thus (Boddaer t , Woods and Schultz , 1953 .

Fam i ly POMACENTRIDAEAbudefduf sep temfasciotu s (Cuv ier , Woods and Schultz , 1953 .

Abudefdu/ sex/osciotus (Lacepede,

Abudefduf coe les t in us Al len , 1975 . A l len , Bauchot , and Desou tter ( 1978) placed A . coeles t inus (Cuv ier ) in thesynonymy of A . sexfasciotu s .

Abudefduf sordidus (For s skal , Woods and Schultz , 1 960 .

Amblyglyphidodon aureus (Cuv ier ,Abude/du] aureus Woods and Schultz , 1960 .

Amblyg lyphidodon curacao (B loch ,Abudefduf cur acao Woods and Schultz , 1960 .

Amblyg lyphidodon leucogas ter (B leeker , Randal l , 1 986 .

Amph ipr ion chrys op ter us Cuv ier , 1830 .

Amph ipr ion bicinctus Woods and Shultz , 1960 . A l len (1 972a) found specimens of the true A . bicinctus Ruppell on lyfr om the Red Sea.

Amph ipr ion melanop u s B leeker , 1 852 : Woods and Schultz , 1960 .

Amph ipr ion pe r ider aion B leeker , 1 955 : Woods and Schultz , 1960 .

Amph ipr ion t r icinctu s Schultz and Welander , 1 953 .

Chr om is acar es Randal l and Swerdloff , 1 973 .

Chr om is ag i l is Sm i th , 1 960 : Randal l and Swerdloff, 1 973 .

Chrom is leucu r us Woods and Schultz , 1960 . (Th is table cont inued on next page. )

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TABLE 1 (con t'd)

Fam i ly MUGIL IDAEChaenomugi l leuciscus (G ijn ther ,Neomyxus chap tal i i Schultz , 1953 . James M . Thom son (per sonal commun ication) stated that Chaenom ugi l leuciscus isthe cor r ect name for the specimens Schu ltz identifi ed as N . chap tal i i (Eydoux and Sou leyet ) . Th is and the fo llowi ngmug i l id gener ic placements were made wi th his counsel .

Chelon labiosus (Valenciennes ,P l icom ugi l labios us Schultz , 1 953 .

Cr en im ugi l cren i labis (For s skal , Schultz , 1 953 .

L iz a voigiens is (Quoy and Gaimard,

Che lon vo ig iens is Schultz , 1953 .

Volamugi l engel i (B leeker ,Chelon engeli Schultz , 1953 .

Fam i ly SPH YRAEN IDAESphyraena bar r acuda (Walbaum , Schu ltz , 1 953 .

Sphyr aena for s ter i Cuv ier , 1 829 : Schultz , 1 953 .

Sphyr aena hel ler i Jenkins , 1 90 1 : Schu ltz , 1 953 .

Sphyr aena gen ie Klunz inger , 1 870 : Schu ltz , 1953 .

Fam i ly POLYNEMIDAEPo lydacty lus sexfi l is (Valenciennes , Schultz , 1953 .

Fam i ly LABR IDAEAnamp ses caer u leop unctotu s Ruppell , 1828 : Schu ltz , 1960 .

Anamp ses melanu r u s B leeker , 1 857 : Randal l , 1986 .

Anamp ses meleagr ides Valenciennes , 1 839 : Randal l , 1986 .

Anamp ses twis t i i B leeker , 1 856 : Schu ltz , 1960 .

Bodianus an thio ides (Bennett , Randal l , 1 986 .

Bod ianus ax i l lar is (Bennett , Randal l , 1 986 .

Bod ianus diono (Lacepede, Randal l , 1 986 .

Bod ianu s loxozonus (Snyder , Randal l , 1986 .

Chei l inus arenatus Valenciennes , 1 840 : Randal l , 1981 c .

Chei l inus bimacu latus Valenciennes , 1 840 : Randal l , 1 986 .

Chei l inus celebicus B leeker , 1 853 : Schu ltz , 1960 .

Chei l inu s chlor ou r us (B loch , Schu ltz , 1960 .

Chei l in us digr omm us (Lacepede, Schu ltz , 1 960 .

Chei l inus fasciatus (B loch , Schultz , 1 960 .

Chei l in us or ien ta lis G ii n ther , 1 862 : Randal l , 1986 .

Chei l in us oxycephalus B leeker , 1 853 : Schu ltz , 1 960 .

Chei l in us t r i lobatus Lacepede, 1 801 : Schultz , 1 960 .

Chei l in u s undulatus R ii ppell, 1 835 : Randal l , H ead, and Sander s , 1 978 .

Chei l in us un ifasciatu s Streets , 1 877 .

Chei l inus r hodochrous Woods , 1 960 .

Chei l io inerm is (For sskal , Randal l , 1 986 .

Cir r hi lobr us exqu is itus Sm ith , 1 957 : Randall , 1986 .

Cir rhi lobr us sp .

Ci r r hi lobr us temm incki Schultz , 1960 . Th is is an undescr i bed species al l ied to C . temm incki i B leeker(Randal l , MS) .

Ci r r hi lobr us sp . A new species , found on_

dense beds of benth ic algae in lagoons ofEnewetak and Kwajalein in to m (Randal l , MS) .

C i r r hi lobr us sp . A new species related to C . cyanop leu ra (B leeker ) (Randal l , MS) .Ci r r hi lobr us sp . A new species found in outer r eef areas in 30 m or mor e (Randall , MS) .Co r is aygu la Lacepede, 180 1 : Schultz , 1960 .

Cor is golmard (Quay and Gaimard, Schultz , 1 960 .

Cor is var iegota (R i ippell , Schu ltz , 1960 .

Cymo lu tes p roetex tatus (Quay and Ga imard, Schultz , 1 960 .

Cym olu‘tes torquatus (Valenciennes , Randall , 1986 .

Ep ibu lus ins idiator (Pal las Schultz , 1 960 .

Gomphosus var ius Lacepede, 1 80 1 : Schu ltz , 1 960 .

G omphosus t r ico lor Schultz , 1 960 . Str asburg and H iatt ( 1 957) discovered that G . t r ico lor Quay and Gaimard is themale of G . var ius .

H olichoer es biocellatus Schultz , 1960 .

H olichoe res chrys us Randal l , 1 980 .

H ol ichoer es hor tu lonus (Lacepede, Schultz , 1 960 .

(Th i s table continued on next page. )

ANNOTATED CH ECKLIST OF F ISHES 305

TABLE 1 (con t’d)

Fam i ly LABR IDAE (cont ’d)H ol ichoer es morgar itoceus (Valenciennes , Schultz , 1 960 .

H olichoer es marginatus Ruppell , 1 835 : Schultz , 1 960 .

H ol ichoeres melanu r us (B leeker ,H ol ichoeres hoeven i Schu ltz , 1 960 .

H ol ichoeres kallochroma Schultz , 1960 . Randal l (198Gb) concluded that Schultz m is identified the female of

H . melan u r us as H . hoeven i i B leeker [actual ly hoeven i i is the female of H . vr oliki i (B leeker )]and the male as

H . kallochroma (B leeker ) .Hol ichoeres melasmapom us Randal l , 1 980 .

H ol ichoeres r ichmondi Fowler and Bean , 1 928 : Randal l , 1 986 .

H ol ichoeres tr imacu latus (Quoy and Gaimard, Schultz , 1960 .

H olichoeres sp . Probably undescr ibed; close to H . hor tz feldi i (B leeker ) .H em igymnus fasc iatus (Bloch,

Schu ltz , 1 960 .

H em igymn us melap ter us (B loch , Schultz , 1 960 .

H ologymnos u s annu latus (Lacepede,Randal l , 1 982 .

Labr ichthys un i l ineotus (Gu ichenot ,Labr ichthys cyanotoen io Schultz , 1 960 . Randal l and Spr inger (1 973 ) showed that L . un il ineotus has pr ior ity overL . cyanotoen io B leeker .

Labr o ides bico lor Fowler and Bean , 1 928 : Schu ltz , 1 960 .

Labr o ides dim idiotu s (Valenciennes , Schultz , 1 960 .

Labro ides pecto r al is Randal l and Spr inger , 1 975Labr ops is al len i Randal l , 1 98 1 .

Labrop s is m icrones ica Randal l , 1 98 1 .

Labr op s is xan thonota Randal l , 1 98 1 .

Mocropharyngodon meleagr is (Valenciennes , Schultz , 1 960 .

Mocr opharyngodon pardal is Schultz , 1 960 . Randal l ( 1 978b) determ ined that M . pardal is (Kner ) is the female of

M . meleagr is .

1

Mocr opharyngodon negr osens is H er r e, 1932 : Randal l , l 978b.

Novocu lichthys taen iou r u s (Lacepede,

Xyr ichthys taen iou r us Schu ltz , 1960 .

Parachei l inus sp . An undescr ibed species r elated to P . filamen tos us A l len (Randall , MS) .Pseudochei l inu s evon idus Jordan and Evermann , 1 903 : Randal l , 1986 (after Str asburg) .Pseudochei l inus hexotoen io (B leeker , Schultz , 1 960 .

Pseudochei l inus octotoen io Jenkins , 1 900 : Schultz , 1 960 .

Pseudochei l inus tetratoen io Schu ltz , 1 960 .

Pseudochei l inus sp . A new species wi th a large ocellated black spot on s ide of caudal peduncle.

Pseudocor is yamosh i ro i (Schm idt , Sm ith-Van i z and Randal l (MS) di scuss th is species in thei r r ev is ion of the genus .

Pseudodox mo luccanus (Valenciennes , Randal l , 1 986 .

Pseudoju lo ides ceras inu s (Snyder , Randal l , 1 986 .

Pterogogus cr yptas Randal l , 1 98 1 .

Pter agogu s guttatus Schultz, 1960 . Randal l ( l 98 l c) r eidentified the specimen fr om Rongelap wh ich Schultz ca l ledP . gu ttatus (Fowler and Bean) .

Stethoju l is bandanens is (B leeker ,Stethoju l is ax i l lar is Schultz , 1 960 .

Stethoju l is linear is Schultz , 1 960 . Randal l and Kay (1974) showed that S . ax i l lar is (Quoy and Gaimard) is the in i tialphase of the H awai ian endem ic species 8 . balts ata (Quay and Gaimard) , and S. linear is Schultz is the term inal male of

S . bandanens is .

Stethoju l is s t r ig iven ter (Bennett , Schultz , 1 960 .

Thalas soma omblycephalum (B leeker ,Thalassoma amblycepha lus Schultz , 1 960 .

Thalassoma hardwicke (Bennett ,Tha lassoma hardwickei Schultz , 1 960 .

Thalass oma Iunare (L innaeus , Schultz , 1 960 .

Thalas soma lutescens (Lay and Bennett Schultz , 1960 .

Thalas soma purpureum (For sskal , Schultz , 1960 .

Tha las soma umbros tygma Schultz , 1 960 (in par t). Randal l and Edwards (1-984) have shown that T . umbros tygma (Ruppell)is a jun ior synonym of T . purpur eum based on the in itial phase (ho lotype of umbr os tygma exam ined in SenckenbergMuseum ) . The in i tial phase of purpureum is near ly identica l in co lor to that of T . t r i lobotum .

Thalas soma qu inquevittotum (Lay and Bennett ,Thalassoma qu inquevittotus Schultz , 1 960 .

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TABLE 1 (con t’d)Fam i ly LABR IDAE (cont ’d)Thalassoma tri lobotum (Lacepede,

Tha las soma fu scum Schu ltz , 1960 . Thalas soma fuscum (Lacepede) , descr ibed in Labr us , is a jun ior homonym of L fuscasGmel in , as po inted ou t by Randall , l 955a.

Tha las soma umbros tygma Schultz , 1 960 (in par t) .Wetmo r ello albofasciota Schultz and Mar shall , 1 954 : Randal l , 1983 .

Wetmor ello n igr op innato (Seale,

Wetm orello oce l lata Schultz and Mar shal l , 1 954 . Randall ( 1 983) determ ined that W . ocel lata i s a jun ior synonym of

W . n igr op in nata.

Xyr ichtys oneiten s is (Gun ther ,H em ipteronatus oneiten s is Schultz , 1 960 .

Xyr ichtys celebicus (B leeker ,H em ip teronotus celebicus Schultz , 1 960 .

Xyr ichtys pavo Valenciennes , 1 839: Randal l , 1986 .

Fam i ly SCAR IDAEColotom us car ol in us (Valenciennes ,Colo tomu s sp in idens Schu ltz , 1 958 ; Schultz , 1 960 . B ruce and Randal l ( 1 985) showed that Schultz and some otherauthor s have m is identified C . car ol inus as C . sp in idens (Quoy and Gaimard) .

Colotomu s sp in idens (Quoy and Gaimard, Randal l , 1 986 .

Cetoscor u s bico lor (R i ippell ,Chlor u r us bico lor Schultz , 1958 ; Schultz , 1960 .

Chlor u r us pu lche l lus Schultz , 1958 ; Schu ltz , 1960 . Randall (1963b) found that Cetoscor u s pu lche llus (Ruppell) is theterm inal male of C . bico lor .

H ipposcor us long icep s (Valenciennes ,Scarus har id Schultz , 1 958 ; Schu ltz , 1 960 . Randal l and B ruce (1 983) showed that H ipposcor us har id (For s skal) is anIndian Ocean species di s tinct fr om the Pacific H . longicep s .

Scarus alt ip inn is (Steindachner ,Scarus brevifil is Schultz , 1958 ; Schultz , 1 960 .

Scarus chlo rodon Schu ltz , 1 958 ; Schu ltz , 1 960 . Randal l and Choat (1980) pointed ou t that S. chlorodon Jenyns is the

term inal male (and a jun ior synonym ) of S. p ros iognothos Valenciennes . Scarus brevifil is (Gunther ) Is a jun ior synonym of

S. altip inn is based on the in itial phase.

Scor u s otropectorol is Schu ltz , 1958 ; Randal l , 1986 .

Scarus bleeker i (de Beaufor t , Randal l and Choat , 1980 .

Scarus dim idiotus B leeker , 1 859 : Randal l , 1 986 .

Scarus fes t ivas Valenciennes , 1 840 .

Scarus lunu la Schultz , 1 958 ; Schultz 1960 . Randall and Bruce ( 1 983) determ ined that S. fes t ivas i s an o lder name forS. lunu la (Snyder ) .

Scarus flavipectorol is Schul tz , 1 958 : Randall and Choat , 1 980 .

Scarus fo r s ten i (B leeker , Randal l , 1986 . Schu ltz (1 958 ; 1969) confu sed two scar ids under the name 8 . lep idus(non-Jenys ) . Randal l and Choat ( 1 980) fo l lowed but used the name 8 . tr ico lor B leeker . Randall ( 1986) has s hownthat the tr ue 8 . t r ico lor appar en t ly does not occur in the Mar shal l Is lands , but the r elated 8 . for s ten i does .

Scor us fr enotus Lacepede, 1 802 : Randal l , 1986 .

Scarus f ron ta l is Valenciennes , 1839 .

Scar us jones i Schultz , 1 958 ; Schultz , 1960 . Randal l and B ruce ( 1983) ment ioned that S. fr on tol is has pr ior i ty overS. jones i (Streets ) in thei r account of the related S. en neacan thus Lacepede.

Scor u s ghobbon For sské l , 1775 : Randall , 1 986 .

Scarus gibbus R'

Lippell , 1 828 .

Scarus m icr or hinos Schultz , 1958 ; Schultz , 1 960 . Sm ith (1 959) showed that Schultz ( 1958) was in er r or in applyi ng thename Chlor u r us gibbus (Ruppell) to the largest of the par r otfi shes , Bo lbometopon mur icatum (Valenciennes ) . Bo lbometoponmur icatum i s not known fr om the Mar shal l Is lands , but it occur s in the Gi lber ts and the L ine Is lands so It m ighteven tually be found in the souther n Mar shal ls . Scar u s m icr orhinos B leeker is r egar ded as a jun ior synonym of S.

gibbus by Randal l and B ruce fo llowi ng SchultzScarus g lobicep s Valenciennes , 1 840 : Schultz , 1958 ; Schu ltz , 1960 .

Scarus oer uginos us Schu ltz , 1958 , 1960 (in par t) . Randal l and B ruce ( 1 983) documented th is m is ident ifi cation . The

tr ue 8 . oer uginos us Valenciennes is a jun ior synonym of the Red Sea endem ic S. fer r ugineus For sskal (Randal l andOrmond,

Scarus n iger For sskal , 1 775 : Randal l , 1 986 .

Scarus ovicep s Valenciennes , 1839 : Randal l , 1 986 .

Scarus ps i ttacus For sskal , 1775 .

Scor u s for s ter i Schultz , 1958 ; Schultz , 1960 . Randal l and Onnond ( 1 978) demons tr ated that S . ps ittacus (the typespecies of Scor us ) i s a sen ior synonym of S. for s ten

‘

Valenciennes ; they descr i bed a neotype of S. ps ittacus .

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TABLE 1 (con t'd)

Fam i ly BLENN IIDAE (cont ’d)P logiotr em u s loudondus loudondus (Wh it ley , Sm ith-Van i z , 1 976 .

P logi otr em u s r hinor hynchos (B leeker , Randal l , 1 986 .

P logio tr em u s tapeinosomo (B leeker , Schu ltz , 1960 .

P r aealt icus sp . An un identi fi ed species fr om an underwater photogr aph taken in Kwajalein .

Rhabdoblenn iu s r hobdot rachelus (Fowler and Ba ll , Schultz and Chapman , 1 960 .

Rhabdoblenn ius snowi (Fowler , Schultz and Chapman , 1960 .

Salor ios fas ciotus (B loch , Randal l , 1 986 (after Strasburg) .Stanu lus seychellens is Sm i th , 1 959 .

Fallacir r ipectes m inutas Schul tz and Chapman , 1 960 and F . wells i Schultz and Chapman , 1960 . Sm ith-Van i z andSpr inger (1 971 ) relegated F . m inutas and F . wells i to the synonym y of Stanu lus seychellen s is .

Fam i ly TRIPTERYG IIDAEEnneap terygi us cop idata (Rosenblatt in Schultz ,Enneap terygi u s ch ica (Rosenblatt in Schultz ,Enneap terygius hem imelos (Knet and Steindachner , Schultz , 1 960 .

En neap terygi u s m in utus (Gunther , Schul tz , 1 960 .

Enneap terygius nanus Schu ltz , 1 960 .

H elcogr ommo hudson i (Jordan and Seale, H ansen , 1 982 .

Norfolkio brachylep is (Schultz ,Tr ip teryg ion brachylep is Schu ltz , 1 960 .

Fam i ly SCH INDLERIIDAESchindlen

‘

a p raemotu r u s (Schl indler , Schultz , 1960 .

Fam i ly CALL IONYMIDAECal l ionymus s imp l icicor n is Valenciennes , 1837 : Fr icke, 1 982 .

D ip logr omm us gor amens is (B leeker , Schu ltz , 1 960 .

Synchi r op u s loddi Schu ltz , 1 960 .

Synchi rop u s mor r i son i Schultz , 1960 .

Synch i r opus ocellatus (Pal las , Randal l , 1 986 .

Fam i ly GOB I IDAEAllom icrodesm us dorotheoe Schu ltz , 1 966 .

-Dawson ( 1 974) showed that th is species is not a

m icr odesm id; V ictor G . Spr inger (per sonal commun ication) class ifi es i t as a goby of the subfam i ly Xen is thm inae.

Amblyeleotr is fasciata (H er r e,

Amblyeleot r is gu tt ota (Fowler ,Amblyeleot r is s tein itz i (Klau sew i t z ,Amblyeleot r is wheeler i Polun in and Lubbock , 1 977 .

Amblygobius decus satus (B leeker ,Amblygobius pholaena (Valenciennes ,Amblygobius rainfordi (Wh it ley ,As ter rop teryx ens ifer us (B leeker ,As ter r op teryx sem ipunctotus R ii ppell , 1830 .

Bathygobius cocosens is (B leeker ,Bathygobius fuscus fuscas (R i i ppell ,Cabi l las tongarevoe (Fowler ,Callogobius boucho toe Gor en , 1979 .

Callogobius cen t r o lep is Weber , 1909 .

Cal logobiu s hos selt i (B leeker ,Callogobiu s macu l ip inn is (Fowler ,Callogobiu s ok inawoe (Snyder ,Callogobius s cloter i (Steindachner ,Callogobius sp .

Cryp tocen t r us s t r igi ll icep s (Jordan and Seale,

Ctenogobiops au roc ingu lu s (H er re,

Ctenogobiop s sp .

Eviato ofelei Jordan and Sea le, 1 906 .

Evio ta dis t igma Jordan and Scale, 1 906 .

Eviato fasc io la Kar nella and Lachner , 1 98 1 .

Ev io ta infu lato (Sm i th ,Eviato me lasma Lachner and Kar nella, 1 980 .

Eviota nebu losa Sm i th , 1 958 .

Evio ta sebr eei Jordan and Seale, 1 906 .

Evio ta smara dus Jordan and S l 1 906 .9 ea e

(Th is table cont inued on next page

ANNOTATED CHECKLIST OF F ISHES

TABLE 1 (con t’d)

Fam i ly GOB I IDAE (cont ’d)Eviato zon u ra Jordan and Seale, 1906 .

Eviota spp . (2 species ) .F us igobius neophyt us (Gunther ,F us igobius spp . (2 species ) .G natholep is onjerens is (B leeker ,G natholep is scopu los t igmo H er re, 1 953 .

Gobiodon cit r inus (R ii ppell ,G obiodon erythrosp i lus B leeker , 1 875 .

Gobiodon okinawoe Sawada, Ar ai , and Abe, 1 972 .

G obiodon r ivu latus (R i ippell , G l inther , 1 877.

H etereleot r is sp .

Is t igobius decoratus (H er r e,Is t igobius or natus (Ruppell ,Is t igobius r igi l ius (H er r e,

Kelloggello cen tralis H oese, 1 975 .

Lot i l io gr oci l iosa Klausew i tz , 1960 .

Macrodon togobius wi lbu r i H er r e, 1936 .

Op lopomops diacan thus (Schultz ,Op lopom us Op lopom us (Valenciennes ,Opuo nephodes E. K . Jordan , 1925 .

Pandoko pruinosa (Jordan and Seale,

Parogobiodon echinocepholus (Ri ippell , Gunther , 1 877 .

Par ogobiodon lacun icolus Kendal l and Go ldsbor ough , 191 1 .

Par ogobiodon melonosom us (B leeker ,Par ogobiodon modes tus (Regan ,

Parogobiodon xan thosomus (B leeker ,Pleur os icyo bi lobatus (Koumans ,Pleu r os icyo m uscar um (Jordan and Seale,

P r iolep is for cimen (Jordan and Evermann ,

Si lhouetteo sp .

St iphodon elegans (Steindachner ,Tr imma coes iu ro Jordan and Seale, 1 906 .

Tr imma eviotop s Schu ltz , 1 943 .

Tr imma noudei Sm ith , 1 956 .

Tr imma ok inawoe (Aoyagi ,Tr imma spp . (6 species ) .Valencienneo p uellor is Tom iyama, 1 956 .

Valencienneo sexgu ttatus (Valenciennes ,Valencienneo s t r igotus (Br oussonet ,Valencienneo sp .

Vonder hor s t ia am banoro (Fou rmano i r ,Vonder hor s t ia spp . (2 species ) .Xen is thm us chapm on i (Schultz ,Kraemer icus chapmon i Schultz , 1 966 . V ictor G . Spr inger (per sonal commun ication) advi sed us that K . chapmon i is

a kr aemer i id but a goby of the genus Xen is thm us .

Xen is thm us sp .

Fam i ly ELEOTRID IDAEOphieleot r is opo r os (B leeker ,Fam i ly KRAEMERIIDAEKr aemer io bryan i Schultz , 1 941 : Schultz , 1966 .

Kraemeri o samoens is Steindachner , 1 906 : Schultz , 1966 .

Fam i ly MICRODESMIDAEG u nnell ichthys monos t igma Sm ith , 1 958 : Str asbur g , 1 967 .

G unnell ichthys p leu r otaen io B leeker , 1 858 : Schultz , 1966 .

Paragobioides gr andocu lis Kendal l and Go ldsbor ough , 1 91 1 .

N emateleot r is helfr ichi Randal l and A l len , 1 973 .

Nemateleot r is magn if ica Fowler , 1 938 : Randal l and A l len , 1 973 .

Ptereleotr is cvides (Jordan and H ubbs , Randal l and H oese, 1985 .

P tereleo t r is hanae (Jordan and Snyder , Randal l and H oese, 1 985 .

Ptereleot r is heteroptera (B leeker , Randal l and H oese, 1985 .

(Th i s table cont inued on next page . )

3 10 RANDALL AND RANDALL

TABLE 1 (con t’d)

Fam i ly MICRODESMIDAE (cont'd)Ptereleot r is m icro lep is (B leeker , Randal l and H oese, 1 985 .

P ter eleot r is zebr a (Fowler , Randal l and H oese, 1985 .

Fam i ly ACANTH UR IDAEAcan thurus achi l les Shaw , 1 803 : Schu ltz and Woods , 1953 .

Acan thurus blochi i Valenciennes , 1 835 .

Acan thurus mota Schultz and Woods , 1953 . Randal l (MS) has shown that many author s have been m istaken in applyingthe name A . mota to th is fish . It is the ea r l iest name for the species that most authors have identified as A . bleeker i

Gun ther , 1 86 1 .

Acan thurus gu ttatus B loch and Schneider , 1 80 1 : Schultz and Woods , 1 953 .

Acan thurus l ineatus (L innaeus , Schu ltz and Woods , 1 953 .

Acan thurus mocu l icep s (Ahl , Randal l , 1 986 .

Acan thurus moto (Cuv ier ,Acan thurus bleeker i Schultz and Woods , 1 953 . See discus s ion under A . blochi i .

Acan thurus n igri cans (L innaeus ,Acan thurus olio la Woods and Schultz , 1 953 . Randall (1956) placed A . oliola Lesson in the synonymy of A . g loucopareius

Cuv ier ; both , however , are synonym s of A . n igr icans (L innaeus ) (Randal l , MS) .Acan thurus n igr icauda Du ncker and Mohr , 1 929 .

Acan thurus n igr icans Schultz and Woods , 1 953 . Acan thur u s n igr icans (L innaeus ) Is the species mos t author shave identi fied as A . g loucopar eius .

Acan thurus gahhm Randal l , 1 956 . Acan thur u s gahhm (For sskal) is an endem ic Red Sea species .

Acan thurus n igr ofuscus (For sskal ,Acan thurus elongatus Schultz and Woods , 1 953 (in par t) . Randall (1 956) indicated that n igr ofuscus i s the o ldes t namefor th is species ; A . e longatus (Lacepede) is not the same fi sh .

Acan thurus n igr or is Valenciennes , 1 835 .

Acan thurus elongatus Schultz and Woods , 1 953 (in par t) .Acan thur us ol ivaceus B loch and Schneider , 1 80 1 : Schu ltz and Woods , 1 953 .

Acan thurus pyr ofer us K i tt l i tz , 1 834 .

Acan thur us leucos ter non Schu ltz and Woods , 1953 . Randall ( 1 956) showed that th is is a m is identifi cation ;A . leucos ter non Bennett i s an Indian Ocean species .

Acan thurus thomp son i (Fowler ,Acan thur u s phi l ipp inus Schultz and Woods , 1 953 . Randal l (1 956) referr ed A . phi l ipp inus H er r e to the synonymy ofA . thomp son i .

Acan thurus t r ios tegus tr ios tegus (L innaeus , Woods and Schultz , 1 953 .

Acan thurus xan thop ter u s Valenciennes , 1 835 : Randal l , 1 980a .

Ctenochaetus binototus Randal l , 1 955b : Randal l , 1986 .

Ctenochaetus hawai iens is Randal l , 1955b : Randall , 1 986 .

Ctenochaetus margi natus (Valenciennes ,Ctenochaetus sp . H iyama , 1 943 .

Ctenochaetus cyanogu ttotus Randal l , 1 955b .

Ctenochaetus s t r iatu s (Quoy and Gaimard, Schultz and Woods , 1 953 .

Ctenochaetus s t r igos us (Bennett , Randal l , 1986 .

Naso onnu latus (Quay and Gaimard, Schultz and Woods , 1 953 .

Naso br evi r os t r is (Valenciennes , Schu ltz and Woods , 1 953 .

Naso hexocan thu s (B leeker , Schul tz and Woods , 1953 .

Naso l itu r otus (B loch and Schneider , Schultz and Woods , 1 953 .

Naso tube rosus Lacepede, 1 80 1 : Randall , 1 986 .

Naso un icom is (For sskal , Schultz and Woods , 1 953 .

Naso vlam ingi i (Valenciennes , Schultz and Woods , 1 953 .

Parocan thu r u s hepatus (L innaeus , Randal l , 1 98 1d.

Zebrasomo flavescens (Bennett , Schultz and Woods , 1 953 (In par t) .Zebr asomo scopas (Cuv ier ,Zebr as omo flevescens Schultz and Woods , 1953 (in par t) . Randal l ( 1955c) showed that Z . flavescens and Z . scopas aredi stinct species .

Zebrasomo ve l ifer um (B loch , Schultz and Woods , 1 953 .

Fam i ly ZANCL IDAEZanclus com u tu s (L innaeus , Woods , 1 953 .

Zanclu s canescen s Woods , 1 953 . Randal l ( 1 955a) demons trated that Z . canescens (L innaeus) is the late post lar val stageof Z . com u tus . The latter has pr ior i ty due to deci s ion of the fi r st rev isor , Cuv ier , 183 1 .

(Th is table cont inued on next page

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3 12 RANDALL AND RANDALL

TABLE 1 (con t ’d)

Fam i ly MONACANTH IDAEA lu ter us scr iptus (Osbeck ,Alu ter o scr ipto Woods , 1 966 .

Amanses scopas (Cuv ier , Woods , 1 966 .

B r acholu teres taylor i Woods , 1 966 .

Con ther hines dumer i li i (H o l lard ,Amanses cor o loe Woods , 1966 . Randal l ( 1964b) determ ined that Can ther hines cor oloe (Jordan and McG r egor ) is a

jun ior synonym of C . dumer i l i i .

Can ther h ines f r on t ic inctus (P layfai r and Gunther , Randal l , 1 986 .

Con ther hines pardal is (R ii ppell ,Amanses sandwichiens is Woods , 1 966 . Randall (1 964b) showed that C . sandwichiens is (Quay and Gaimard) is endem ic tothe H awai ian Is lands . The cor r ect name for the wide-

r anging species fr om elsewher e in the Indo-Paci fic is C. par da lis .

Oxymonacon thus long i ros t r is (B loch and Schneider , Woods , 1 966 .

Porolu teres p r ionu r us (B leeker , Woods , 1966 .

Paramonacon thu s cryp todon (B leeker , Woods , 1 966 .

Par amonacon thu s oblongas (Temm inck and Schlegel , Woods , 1966 .

Per vagor altem ans (Og i lby ,Per vagor melanocephalus mar shallens is Woods , 1 966 . Synonymy fr om H utch ins , 1986 .

Per vagor asp r icoudatus H o l lard, 1 854 .

Per vagor melanocephalus johns tonens is Woods , 1966 .

Per vago r melanocephalus (B leeker , This record based on one specimen r eported by H utch ins , 1 986 .

Fam i ly OSTRACIIDAELoctor io for nos in i (B iancon i , Woods , 1 966 .

Os t r acion cubicus L innaeus , 1 758 : Woods , 1 966 .

Os tracion meleagr is Shaw, 1 796 : Woods , 1 966 .

Fam i ly TETRAODONT IDAEAmblyr hynchotes honcken i i (B loch , Randal l , 1 986 (after Strasburg) .A r o thr on h isp idus (L innaeus , Woods and Schu ltz , 1966 .

A r othron man i len s is (Procé , Randal l (1 985) has shown that the strongly str iped A . man i lens is is a distinct speciesfr om the r elated unmarked A . immocu latus (B loch and Schneider ) .A r othron mappa (Lesson , Randal l , 1986 (after Str asbur g) .A r othron meleagr is (Lacepede, Woods and Schultz , 1 966 .

A r othr on n igr op unctatus (B loch and Schneider , Woods and Schu ltz , 1 966 .

A r othr on s te l latus (B loch and Schneider ,A r othron albo ret icu lotus Woods and Schultz , 1 966 .

Con thigas ter ambo inens is (B leeker , Woods , 1966 .

Con thigas ter bennett i (B leeker , Randal l , 1986 .

Con thigas ter coronata (Vai l lant and Sauvage, Randal l , 1 986 .

Con th igas ter ep i lamp r o (Jenk in s , Randal l , 1 986 .

Con thigas ter jan th inop tera (B leeker ,Con th igas ter jactator Woods . A l len and Randal l (1 977) regarded C . jactotor (Jenk ins ) as a H awai ian endem ic . Therelated w idespr ead species from elsewhere in the Indo-Paci fic reg ion is C . jan thinop tero .

Con th igas ter so landr i (R ichardson , Woods , 1 966 .

Con th igas ter va len t in i (B leeker , Randal l , 1986 .

Fam i ly D IODONT IDAED iodon hys t r ix L innaeus , 1 758 : Woods , 1966 .

D iodon l itu r os us Shaw , 1 804 : Randal l , 1 986 .

th i s l i s t . On ly a few such fis hes have been repor ted fr omthe ar ea . Schultz in Schultz et al . fo r example,

l i s ted on ly two myctophids , M yctop hum br achygnathos

(B leeker ) and D iaphus schm idt i Tan ing , both by name on ly .

Of the Gonos tomat idae, Gr ey ( 1 960) r eco r ded on lyD ip IOphos sp . (taen ia complex) , Gonos tomo at lan t icum No r

man , and G . ebel ing i Grey fr om the Mar shal ls r eg ion . R . K .

Johnson r ecent ly iden t i fied a g igantu r id (BPBM 26320 ,

mm SL ) taken w i th a m idwater tr awl at 1 1 °20 ’N ,

1 62°O7

’E at a depth of 0 to 200 m as Rosaur a inc

(B r auer ) .Ther e has not been adequate sampl ing of deep-wa

fishes fr om the Mar shal l Is lands or , for that matter , Inthe i s lands of Ocean ia in gener al (except the H awaiIs lands ) . Even spec ies of moder ate depths ( 1 00 to 300 1

such as snapper s of the gener a E tel is and P r is t ipomok

have not been repor ted fr om the Mar shal ls , though 50 ]su rel y occur ther e. If complete co l lect ions were made

ECKLIST OF FISHES 3 13

l is ted as s ynonyms below the cor r ect name and are

inden ted th ree spaces . When needed , an anno tat ion is prov ided to explain the name change and g ive the author i ty .

On l y those s ynon ym s are l i s ted that were used as val idnames in Schultz et al . The author g iven for each of theses ynon ym s i s the one respons ible for the Mar shal l Is landsr ecor d , no t the author of the species (un less named as new

in Schultz et ai . or in other paper s on the Mar shal l Is landsfi s h fauna) .We have el im inated the l is t ing of Schultz et al . each

t ime Schultz or one of his col laborator s is c ited in the

checkl i s t . Thus Schult z , 1 953 , impl ies Schultz in Schultz etal . , 1 953 . Sim i lar ly , we have shor tened Cuv ier (or Valenciennes ) in Cuv ier and Valenc iennes to jus t Cuv ier (or Valenc iennes ) .

Subgener ic categor ies have not been included in the

checkl is t .The or der of presen tat ion is phylogenet ic , appr ox imat

ing that of Gr eenwood et ai .We have not made a documen tat ion of the d i ffer en t

ato l ls at wh ich the var ious species of fi shes have been col

lected. Such l is t ing i s pr ov ided by Schultz et al . and byRandal l L i ttle d i ffer ence in the fish fauna can be

expected from one ato l l to the next w i th in the Mar shal lIs lands , par t icular l y those of about the same lat i tude.

Ther e ar e d i fferences , however , from the nor ther n to the

souther n Mar shalls . Mor e spec ies of fis hes ar e pr esentlyknown from the nor ther n ato l ls because these have beenmor e heav i l y co llected . Th is is espec ial l y tr ue of B ik in i andEnewetak . B ik in i i s the ato l l bes t r epr esented in the

mater ial at the Nat ional Museum of Natural H is tor y , butthe extens ive B i shop Museum co l lect ions made by thesen ior author and assoc iates over the las t 1 6 year s at

Enewetak have made it the local i ty w i th the mos t species .

We bel ieve,however , that the souther n ato l ls would have

a s l ightly r icher fish fauna i f the co l lect ing effor t wereequal . A num ber of consp icuous spec ies that have beenfound at Kwajalein or other souther n atol ls in the Mar shallshave not been obser ved at Enewetak . Examples are

Myr ip r is t is adus tas , Sargocen t r on r ubr um , An th iasbor t lettor um ,

Ep inephe lus caer u leop unctotus , P lect r opom us

ol igacon thus , Corango ides p lag iotoen ia, Pterocaes io sp . ,

Sco lop s is l ineatus , H en iochus var iu s , H ol ichoer es r ichmondi,

Scar us bleeker i , Amblygobius decus satus , Acan thur usmocu l icep s , Siganus vu lp in u s , and Bol is to ides consp ic i llum .

Th i s , of cou r se,i s not to say that they ar e tr uly absen t

from Enewetak .

H ow does the fis h fauna of the Mar shal l Is lands com

pare w i th the r es t of the Indo-Wes t-Pac i fic r eg ion? Of thees t imated total of spec ies of fishes in the wor ld ,about 8000 ar e shal low , warm -water mar ine spec ies(Cohen , The majo r i ty of these warm -water fishesoccur s in the vas t Indo -Wes t -Paci fic r egion . U s ing two di f

fer en t methods to ar r ive at an es t imate, Spr inger (1 982)concluded that there are about 4000 species of fishes inthe trop ical Indo -Wes t -Pac i fic r egion . Cohen how

ever , es t imated 3000 to 4000 for the Ind ian Ocean alone.

If Cohen is cor r ect , Spr inger adm i tted his es t imate for the

3 14 RANDALL AND RANDALL

Indo-Wes t-Pac i fic as a who le is much too low . By analyz ingthe 1 1 1 shor efish fami l ies which occur nonmarg inally on

the Paci fic Plate, Spr inger es t imated that ther e are 461

genera and an es t imated 1 3 1 2 species . Thus the Mar shal lIs lands , w i th 8 1 7 species , have 6296 of the total Pac i f icPlate fish fauna.

Proper compar isons between i s land gr oups of the

Pac i fic are pos s ible on l y when ther e has been a compar able co l lect ing effor t as wel l as an accur ate comp i lat ion of

species . Th is would seem to be the case on ly for the

H awai ian Is lands , Society Is lands , Samoa Is lands , souther nMar iana Is lands , and Eas ter Is land .

Randal l ( 198 1a) s tated that approx imately 6 10 speciesof fishes occur in the H awai ian Ar ch ipelago . By exclud ingthose wh ich are pelag ic , deep sea , or fr eshwater species ,he ar r ived at 420 wh ich could be r egar ded as r eef andshor e fi shes . Th is figure should be mod i fied to 460 to

include epipelag ic species , r ecen t new r ecor ds , and new

species (to make i t equ ivalent to es timates herein fromother is land gr oups ) . The extreme i so lat ion of the

H awai ian Is lands , bo th geogr aph i cal ly and hydrog r aphi

cally , accounts for i ts somewhat impover i shed fish fauna.

Randal l (1973) pr ov ided a prel iminar y checkl is t of 6 1 6fishes fr om the Soc iety Is lands . By el im inat ing those wh ichoccur in deep water or fresh water but adding new

r ecor ds , the lis t is r ev ised to 620 .

Wass ( 1984) has pr epared a checkl is t of fishes of the

Samoa Is lands . H is total for the l i s t is 958 , wh ich includes84 iden t i fied on l y to genus . If those wh ich occur in deepwater and in fr esh water are el im inated , along w i th dubious l i teratu re recor ds , the total is lower ed to 915 . The

Samoa Is lands lie marginal l y on the Pac i fic P late. Ther e isa continen tal componen t to the f ish fauna of Samoa due toi ts proxim i ty to the cont inental plate. These is lands are

mos t ly h igh is lands w i th a greater var iety of habi tats than

one finds on atol ls . Fur thermor e,they

from other i s land groups than ar e the

fore, a gr eater number of spec ies of

expected from Samoa .

Shepar d and Myer s (198 1 ) publ ishcheckl is t of the fishes of Guam and other

TABLE 2

Summar y of N on sys temat i c F i s h Resear ch at Enewetak

T ype of study F i sh taxa‘Refer en ce

Ecology and Behav iorAcoust ics and H o locen trus spp . , Myr ip r is t is spp .

env i ronment Ca rchar h in idaeReef fishesReef fi shesReef fishesChaetodontidae, Pomacentr idaeAcan thurus gu ttatus , A . tri os tegu s Scar us

Acanthur idae and Scar idaeFauna of nuclear Reef fi shestest cr ater‘T he scientific names g iven in thetable are those that appeared in the publ icat ion cited. Some of these names have been

changed (see check l is t her ein) .(Th is table continued on next page

i s lands wh ich lie on the mar g in of the Paci fic Plate.

Randal l and Cea Egana (1984) made a checklis t of ti lfi shes of Eas ter Is land . On ly 1 30 shor e and ep ipelagspecies have been found at th is smal l , r emote Sou

Paci fic is land . Recent collect ing by Randal l and colleagu

has raised the total to 1 55 spec ies .

Spr inger (1 982) has pr ov ided an anal ys is of endefishes of the Paci fic Plate . H e placed them in thr

categor ies . T ype 1 endem ics ar e those that are wi dely dtr ibuted on the Plate, Type 2 are those that are l imi teda few i s lands or is land gr oups , and T ype 3 are those thare found on l y at a s ing le is land or is land group . TlMar shal l Is lands have 28 of the 48 Type 1 Plate endem i

tabulated by Spr inger . Of the T ype 2 endem ics , Spr ingl i s ted on ly Pomochrom is ex i l is and Labrops is m icr ones 1fr om the Mar shal ls . T ype 3 endem i cs (the mos t commtype) ar e found main ly at h igh i s lands . Spr inger ment ionson ly three from the Mar shal ls , Acon thop les iops hiati

Amph ip r ion t r ic inctus , and Ci r r icoecu la johnson i . To the:

one m ight add H ypoather ino bar nes i , Pseudochrom is ouremar shallens is , and Aser aggodes sm ithi .

In the Paci fic , the H awai ian Ar chipelago has t l

h ighes t per cen tage of endemi sm among i t s shore fishew i th Eas ter Is land is next w i th Lor d H ov

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3 1 6 RANDALL AND RANDALL

Is land , M iddleton and El i zabeth Reefs , and Nor fo lk Is landcomb ined have about 1 296 endemi sm (Randal l ,Randal l ( l 978a) made an appr ox imat ion of 1 096 endem i smfor the fi shes of the Mar quesas . For the Mar shal l Is landsthe Type 3 endem i sm , based on species whi ch have beennamed , i s less than 196. Th is figure wi l l r ise s l ightly whensome new species cur rentl y known on l y from the Mar shal lsar e descr ibed . H owever , when ther e has been a compar able co l lecting effor t in the Centr al Paci fic , and in par t icular

_the G i lber t Is lands and eas ter n Carol ine Is lands (wh ich

l ie near the Mar shal ls and include ato l ls ) , i t i s expected

that mos t of the spec ies now r egarded as endemicMar shal l Is lands w i l l be found elsewher e.

In add i t ion to publ icat ions on the class i fi cat ion of

from the ato l l , there has beens tud ies deal ing w i th the Enewesummar i zes these and g ives the

invo lved .

The thr ee plates (Figs . 1 , 2 , and 3 ) of underwattog r aphs of fishes her ein were taken by the sen ior au

in the Mar shall Is lands .

Plates 1

F i g . 1 a, The gr ay r eef s har k. Ca r char h inus amblyr hynchos

(B leeker ), one of the thr ee mos t abundan t shar ks at

Enewetak ; r es pons i ble for four at tacks on man at the ato l l ;b. Gym nothor ax javan icus (B leeker ), the lar ges t m or ay of theIndo-Paci fi c r eg ion ; atta i n s a lengt h of at leas t 2 m ;

c, Poec i loconger fos c iotus Gun ther ; known fr om on ly fi vespec imens . T h i s one emerged fr om the sand fr om the effect ofich thyoc ide at Enewetak ; d , The squi r r el fi s h Sar gocen t r onm icr os toma (Gun ther ) . Photog r aph taken at n igh t; e, The

F ig . 2 a , A schoo l of the fu s i l ier Cas io caer u laur eus

Lacepede fr om the Kwajalei n lagoon . T h i s photo was the

bas i s for the fi r s t Mar s ha l l Is lands r ecord; b, The noctu r n a lIeth r i n id fi s h M ono tax is g r andocu l is (For s s kal ); has wel ldeveloped mo lar i fon n for cru s h ing m o l lusks , ech i no ids , etc ;

c, The black jack , Co r onx lugubr is Poey . Ci r cum tr opi ca l i ndi sui bu t i on ; d, The goatf i sh Par upeneus bor ber ino ides

(B leeker ) . Recen t co l lecti on s pr ov ided the fi r s t r ecor d for theMar s ha l l Is lands ; e, The co lor fu l dam se l fi s h Pomacen t r us

F i g . 3 a, A g r oup of t he butter fl yfi s h H en iochus chrys os tom us Cuv ier . U sua l ly seen i n pai r s or sm a l l agg r egation s ;b, The yel low angel fi s h Cen t r opyge hera ld i Woods and

Sc hu ltz ; named for the late Ea r l S. H er ald, who col lected m os tof the type spec im en s i n the Mar s ha l l Is lands i n 1946 ; c, Thegoby Amblyeleo t r is gu t to to (Fow ler ) and It s s ym biot i c par tnerA lpheus ochr os t iatus M i ya .

‘

The s hr i mp i s j us t r emov i ng a

load o f Sand and gr avel fr om the bur r ow ; d , The color fu lgoby N em ateleo t r i s helf r ich i Randa l l and A l len , named for

Ph i l ip H el fr i ch , the las t di r ector of the M id-Paci fi c Resea r ch

Spr i nger ; f, The gr ouper Ep inephelus m icr odon (B leekbei ng cleaned by the wr asse Labr o ides dim idiotus (Valeen nes ); g , The br i gh t v io let and yel low A n th ias bar t lett oRandal l and Lubbock , nam ed i n honor of N athan and Patr iBar t lett w ho d i scover ed i t at Kwajalein ; h , The r ed snapLu tjonus g i bbus (F or ss kal ). Photog r aph tak en at n igh t.

pavo (B loch ). B lue i n l i fe w i th yel low on the caudal fi n ; f, Tlanemonef i sh Amph ipr i on chrysop ter us Cuv ier . L i ke other sthe genus , i t l i ves s ym bioti ca l ly w i th sea anemones ; 9 , T i

wr asse Labr ops i s m icr ones ica Randa l l . Enewetak i s the ty;

local i ty . Adul ts feed on cor a l po lyp s ; h , The term inal male

wr asse H ol ichoer es t r imacu latus (Quay and Gaimar d). A c0 1

mon lagoon speci es i n sand and r u bble areas ar ound cor

heads .

Page 3.

Labo r ato r y at Enewetak ; e, The sur geon fi s h Acan thur us n ig

cans L i n naeus , di s ti ncti ve in the wh i te spot under the eye a.

yel low band at the base of the dor sal and anal fi n s ; 1, TIfi lefi s h Oxymonocon thus long i r os t r is (B loc h and Sc hneideblue-g r een w i th or ange spots . Feeds on cor a l po lyp s ; 9 , T i

female of the tr i gger fi s h Xan th ichthys aur omor g inatus (Be

net t ); gener al l y found at depths gr ea ter than 30 m ; h , T]clown tr i gger fi s h Bo l is to ides cons p ic i l lum (B loch ai

Sc hneider ). O ft en identi fied as B . n iger (Bon nater r e), but t Inam e i s a homon ym , hence i n val id.

Page 3]

ANNOTATED CHECKLIST OF F ISHES 3 17

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320 RANDALL AND RANDALL

ACKNO“ &EDGMEN TS

We thank the fol low ing per sons for ass is t ing us in

obtain ing the cor r ect scient i fic names for the fishes in thi scheckl is t : Gerald R . A l len , Ken t E . Carpen ter , P . H . J .Cas tle, B r uce B . Co l lette, Char les E . Dawson , W i l l iam N .

Es chmeyer , Thomas H . Fr aser , Ronald F r icke, Ronald A .

Fr i t zsche, Doug lass F . H oese, Walter Ivan t soff , SusanJewett , Rober t Kar l Johnson , Les l ie W . Knapp , Er nes t A .

Lachner , Kei ichi Matsuur a, John E . McCosker , Edwar d O .

Mur dy, Theodor e W . P ietsch , W i ll iam F . Sm i th-Van i z , V ic

tor G . Spr inger , James M . Thomson ,R i char d C . Wass , Jef

fr ey T . W i l l iams , and Dav id J . Woodland . The manuscr iptwas r ev iewed by Lor i J . Bell , Patr ick L . Col in , V icto r G .

Spr inger , and R ichar d C . Wass .

REF ERENCES '

A l len , G . R . , 1 972a, The Anemonefishes , T . F . H . Publ ications ,Neptune Ci ty , N . J .1 972b , Obser vations on a Commensa l Relationsh ip BetweenSiphom io fusco lineato (Apogon idae) and the Crown-

of-Thor nsStar fish , Acan thas ter p lanol, Cope ia, 3: 595-597 .

1 975 , Dom selfishes of the South Seas , T . F . H . Publ ications , Neptune City , N . J .M . L . Bauchot , and M . Desou t ter , 1978 , The Status of

Abudefduf sexfosciotus (Lacepede), a Pomacentr id Fish fr omthe Indo-West Pacific, Copeia, 2: 328

-330 .

and W . N . Es chmeyer , 1 973 , Turkeyfi shes at En iwetok , Poc .

D iscover y, 3 - 1 1 .

and J . E . Randal l , 1 977 , Rev iew of the Sharpnose Pufferfishes (Subfam i ly Canthigas ter inae) of the lndo-Pacific, Rec .

Aus t ral . M us . , 30: 475-51 7 .

Ar no ld , D . C.,1 956 , A Systematic Rev is ion of the Fishes of the

Teleost Fam i ly Car apidae (Per comorph i , B lenn io idea), wi thDescr ipt ions of Two New Species , Bu l l . B r i t . Mus . Nat . H is t . ,

Zoo l . , 4 : 247-307 .

Bakus , G . J. , 1 967 , The Feeding H abits of Fishes and Pr imar yProduct ion at En iwetok , Mar shal l Is lands , M icr ones ica, 3 :

1 35- 1 49 .

Bar low , G . W .,l 974a , Ext r aspeci f ic Impos ition of Social Gr oup

ing Among Sur geonfi shes (Pi sces : Acanthur idae), J . Zoo l . ,Lond . , 1 74: 333-340 .

1 974b, Contr asts in Social Behav ior Between Centr al Amer ican Cichl id Fi shes and Coral-Reef Surgeonfishes , Am . Zoo l . ,1 4: 9-34 .

Bohlke, J . E . , and J . E . Randall , 198 1 , Four New Garden Eels(Congr idae, H eter ocongr inae) from the Pacific and IndianOceans , Bul l . Mar . Sc i 31 : 366-382 .

and D . G . Sm i th , 1 968 , A New Xenocongr id Eel fr om the

Bahamas , with Notes on Other Species in the Fam i ly , Proc .

Acad . N at . Sc i . Ph i lade lph ia, 1 20 : 25-43 .

B ruce, R . W . , and J . E . Randal l , 1 985 , A Rev is ion of the IndoWest Paci fic Par rotfish Genera Colotom us and Lep tos cor us

(Scar idae: Spar isomat inae), [ado -Pacif ic Fishes , No . 5,pp .

1 -32 .

Bu l lock , T . H . , and J . T . Corwin , 1 979, Acoustic Evoked Activ i tyin the B rain in Sharks , J . Comp . Phys io l . , 1 29: 223-234 .

Burgess , W . E . , 1978 , Bu tterflyf ishes of the Wor ld, T . F . H .

Publ ications , Neptune City , N . J .Cast le, P . H . J. , 1 968a, A Contr ibution to a Rev is ion of the

Mor ingu id Eels , Depar tment of Ichthyo logy , Rhodes Un ivers tty , Spec . Publ . , No . 3 ,

pp . 1 -29 .

1 968b, The Congr id Eels of the Wes ter n Indian Ocean and

the Red Sea, Ichth . Bul l . Rhodes Un iv , No . 33 ,pp . 685-726 .

and J . E . Bdhlke, 1976 , Sexual D imorphi sm inSi ze and Ver

tebr al Number in the Wes ter n At lantic Eel Mor ingua edwar ds i

(Angui l l iformes , Mor inguidae), Bu l l . Mar . Sc i . , 26: 61 5-619.

Cohen , D . M . , 1 970 , H ow Many Recent Fi shes ar e There?, Proc .Cal if . Acad . Sc i . , 4ih ser . , 38 : 341 -346 .

1 973 , Zoogeog r aphy of the Fishes of the Indian Ocean ,

Eco l . Stud , 3 : 45 1-463 .

and J . G . N ielsen , 1978 , Guide to the Identi fi cation of Gen

er a of the Fi sh Order Ophidi i formes wi th a Tentat ive Class i fication of the Or der , NOAA Tech . Rep . NMFS Ci r . , 41 7:

vi i 72 .

Co l lette, B . B . , 1 974 , Geogr aph ic Var iation in the Central Paci ficH alfbeaks , H yporhomphus acutus (G i i n ther ), Poc . Sc i . , 28 :

1 1 1 - 1 22 .

and L . N . Chao , 1 975 , Systematics and Morpho logy of theBon i tos (Sar da) and Thei r Relatives (Scombr idae, Sardin i ),Fish . Bu l l .

,72 : 51 6-625 .

and R . H . Gibbs , Jr . , 1963 , Pr e lim inar y Field Guide to the

Mackere l and Tuna-L ike F ishes of the Indian Ocean (Scom br idae) , U . S. Nat . Mus . , Sm ithson . lnst . , Washington , D . C. ,

pp . v 48 .

Corwin , J . T . , 1 977a, Morpho logy of the Macula Neg lecta in

Sharks of the Genus Car char h inus , J . Morphol , 1 52:

341 -362 .

1977b, Ongo ing H ai r Cel l Production , Matur ation , and

Degeneration in the Shark Ear , Soc . Neu rosci . , 1 977 , 3 : 4 .

1 978 , The Relation of Inner Ear Structur e to the FeedingBehav ior in Sharks and Rays , Scann ing E lect ron M icrosc . , 2:

1 1 05- 1 1 1 2 .

Cr es sey , R . , 1 98 1 , Rev is ion of Indo-West Paci fic L i zardfi shes of

the Genus Synodus (P isces : Synodontidae), Sm ithson . Con tr ib .Zoo l . , 342: i i i 53 .

Cuv ier , G . , and A . Valenciennes , 1 831 , H is to i re Naturel le desPo issons , F . G . Levr au lt , Par is , 7: xx ix 53 1 .

D’

Ancona, U . , and G . Cav inato , 1 965 ,The Fi shes of the Fam i ly

B r egmacer ot idae, Dana-Repor t , 64: 1 -91 .

Dawson , C. E . , 1 974 , A Rev iew of the M icr odesm idae (P isces :Gobio idea) I . Cerdale and Clar kichthys wi th Descr iptions of

Thr ee New Species , Copeia, 2: 409-448 .

1 976 , Rev iew of the lndo-Paci fic Pipefi sh GenusChoero ichthys (Pisces : Syngnath idae) , wi th Descr iptions of

Two New Species , Pr oc . B io l . Soc . Wash . , 89: 39-66 .

l 977a, Rev iew of the Pipefish Genus Co rytho ichthys wi thDescr iption of Thr ee New Species , Copeia, 2: 295-338 .

1 977b, Synops i s of Syngnathine P ipefishes Usual ly Refer r edto the Genus Ichthyocomp us Kaup , with Descr ipt ion of New

Genera and Species , Bul l . Mar . Sci . , 27: 595-650 .

1980 , Notes on Some Siboga Expedition Pipefishes Pr ev ious ly Refer r ed to the Genus Syngnathus , B ijd . D ier kunde, 50:

221 -226 .

1 98 1 , Rev iew of the Indo-Pacific P ipefish GenusDoryr hamphu s Kaup (Pi sces : Syngnath idae), with Descr iptionsof a New Species and a New Subspecies , Ichth . Bu l l . ,J. L . B . Sm ith Ins t . Ichth . , 44 : 1 -27 .

Emer y , A . R . , and G . R . A l len , 1 980 , Stegos tes ; A Sen ior

Synonym for the Damselfish Genus Eupomacen tr us ; Os teologi

cal and Other Ev idence,with Comments on Other Genera,

Rec . Wes t . Aus tr a l . Mus . 8 : 1 99-206 .

Eschmeyer , W . N . , and J . E . Randal l , 1 975 , The Scorpaen idFis hes of the H awai ian Is lands , Including New Species and

New Records (Pi sces : Scorpaen idae), Pr oc . Cal if . Acad . Sc i . ,

40 : 265-333 .

ANNOTATED CH ECKLIST OF F ISHES

Fay, R . R . ,J . I . Kendal l , A . N . Popper , and A . L . Tes ter , 1 974,

V ibr ation Detection by the Macula Neglecta of Sharks , Comp .

B iochem . Phys io l . , 47A : 1 235- 1240 .

Fowler H . W . , 1 928 , The Fishes of Ocean ia, Mem . 8 . P . B ishopM us . , 10: i i i 540 .

Fraser , T . H . , 1 972, Comparative Osteo logy of the Shal lowWater Cardinal Fishes (Per ci for rnes : Apogon idae) wi th Reference to the Systematics and Evo lution of the Fam i ly , Ichth .Bu l l . , J . L . B . Sm ith In s t . Ichth, 34: 1 1 05 .

Fr aser , T . H . , 1 974 , Redescr ipt ion of the Ca rdinal Fish Apogonendekataen ia B leeker (Apogon idae), wi th Comments on Pr eviou s Usage of the Name, Proc . B io l . Soc . Was h . , 87: 3— 1 0 .

Fr icke, R . , 1 982, New Species of Cal l ionymus , wi th a Rev i s ion of

the var iegatus Group of that Genus (Teleostei : Cal l ionym idae) ,J . Nat . H is t . , 1 6: 1 27- 1 46 .

Frit zsche, R . A . , 1 976 , A Rev iew of the Com et fi shes , GenusFis tu lar ia (Fis tu lar i idae), with a D i scus s ion of Int r agener ic Relat ionships and Zoogeogr aphy, Bul l . Mar . Sc i . , 26: 1 96-204 .

Gar r ick, J . A . F . , 1982 , Sharks of the Genus Car char h inus ,NOAA Tech . Rep . , NMFS Ci r c. , 445 : v1i 194 .

Ger ber , R . , and N . Mar shal l , l 974a, Reef Pseudoplankton in

Lagoon T roph ic Systems , Pr oceedings of the Second In ter nat ional Co r a l Reef Sympos ium ,

B r isbane, Gr eat Bar r ier ReefComm i tt ee, 1 : 1 05— 1 07 .

1 974b, Inges tion of Detr i tus by the Lagoon Pelag ic Commun ity at En iwetok Ato l l , L im no . Oceanogr . , 1 9: 8 1 5

-824 .

Gibbs , R . H . , Jr . , and B . B . Co l lette, 1 967, Compar ative Anatamy and Systemat ics of the Tunas , Genus Th unnus , F isher yBu l l . , 66: 65- 130 .

Gi lber t , P . W . , 1 963 , Shar ks and Sur viva l, D . C. H eath andCompany , Boston .

Gi lmar t in , M . , 1 958 , Some Obser vations on the Lagoon Planktonof En iwetok Ato l l , Pac . Sci . , 12: 3 13 -3 16 .

Gos l ine,W . A . , 1 951 , The Os teo logy and Class ificat ion of the

Ophichthid Eels of the H awai ian Is lands , Pac . Sci . , 5:

298-320 .

1 958 , Cen tr al Paci fic Eels of the Genus U r op terygius , withDescr ipt ions of Two New Species , Poc . Sci . , 12: 221 -228 .

and D . W . Str asbur g , 1 956 , The H awai ian Fishes of the

Fam i ly Mor inguidae: Another Eel Pr oblem , Cope ia, 1 :

9- 1 8 .

Gr eenfield , D . W . 1 974 , A Rev is ion of the Squi r r elfi sh GenusMyr ip r is t is Cuv ier (P isces : H o locentr idae), Sc i . Bu l l . , Nat . H is t .

M us . Los Ange les Coun ty, 1 9: 1 -54 .

Gr eenwood , P . H . , D . E . Rosen , S. H . Weitzman , and G . S.

Myer s , 1 966 , Phy letic Studies of Teleostean Fishes , wi th aProv is ional Class ification of L iv ing Form s , Bu l l . Amer . M us .

Nat . H is t . , 1 31 : 339-456 .

Gr ey , M . , 1 960 , A Prel im inar y Rev iew of the Fam i ly Gonos tomat idae,wi th a Key to the Gener a and the Descr iption of a New

Species fr om the T r opica l Pacific, Bul l . M u s . Comp . Zoo l . ,1 22: 57- 1 25 .

Gunther , A . C. L . G . , 1 873— 1 91 0 , Andrew Gar rett 's F ische der

Sli dsee, (Jour . Mus . Godeffr oy) , 1 -3 : 1 -51 5 .

H ansen , P . H . , 1 982 , Revis ion of the Threefi n B lenny Genus H elcogr omma, Inc luding Descr ipt ion of Four New Species and a

D iscuss ion of the Fam i ly Tr ip terygi idae, MS thes i s , U n iver s i tyof Mar y land .

H eemstr a,P . C . , 1972, An th ias heraldi , a Synonym of Lutjanus

g ibbus , an Indo Pac if ic Lutian id Fi sh , Cope ia, 3 : 599-601 .

H elfr i ch , P . , and P . M . Al len , 1975 , Obser vat ions on the Spawning of Mullet , Cren im ugi l cren i labus (Fot skal) , at Enewetak ,Mar shal l Is lands , M icr ones ica, 1 1 : 2 19-225 .

H iatt , R . W . , and D . W . Strasburg , 1 960 , Eco log ica l Relation

32 1

sh ips of the Fish Fauna on Coral Reefs of the Mar shallIs lands , Eco l . Monogr . , 30: 65- 1 27 .

H iyama, Y. , 1943 , Report of an Inves tigation of Po i sonous Fishesof South Seas (in Japanese), N issan F ish . Exp . Stn .

Odowora_

B_

r .

H obson , E . S. , 1 963 , Feed ing Behav ior in Thr ee Species of

Sharks , Poc . Sci . , 1 7: 1 71 -1 94 .

and J . R . Chess , 1978 , T roph ic Relationsh ips Among Fishesand P lankton in the Lagoon at Enewetak Ato ll , Mar shal lIs lands , Fis hery Bu l l . , 133- 153 .

F . Mau t in , and E . 5 . Reese, 196 1 , Two Shar k Incidents at

En iwetok Ato l l , Mar shal l Is lands , Pac . Sci . , 1 5: 605-609 .

H oese, D . F . , J . Bowl ing , B . Russel l , and J . E. Randal l , 1 98 1 , APrel im inar y Rev is ion of Austr al ian Species of Coral T rout ofthe Genus P lect ropomu s , Occ . Rep . Aus tr al . Mus . , 2: i i 37 .

H or ch , K . W . , and M . Sa lmon , 1973 , Adaptat ions to the Acous

t ic Env i r onment by the Squi r r elfishes Myr ip r is t is vio laceus and

M . p r al in ius , Mar . Behov . Phys io l . , 2: 1 21 -139 .

H utchins , J . B . , 1986 , Rev iew of the Monacanth id Fish GenusPer vagor , with Descr iptions of Two New Species , Indo -Pocif icFishes , No . 1 2 , pp . 1 -35 .

Inger , R . F . , 1955 , A Rev is ion of the Fishes of the GenusPles iop s Cuv ier , Pac . Sci . , 9: 259-276 .

Johannes , R . E . , and R . Gerber , 1974 , Impor t and Expor t of NetP lankton by an Enewetak Coral Reef Commun ity , Pr oceedingsof the Second In ter nat ional Cor a l Reef Sympos ium , B r isbane,

Gr eat Bar r ier Reef Comm i ttee, 1 : 97-1 04 .

Johnson , R . H . , and D . R . Nelson , 1 973 , Agon is tic D i splay in theGray Reef Shark , Car char h inus men isor rah, and Its Relat ionsh ip to Attacks on Man , Copeia, 1 : 76-84 .

Kanazawa, R . H . , 1 958 , A Rev is ion of the Eels of the GenusConger wi th Descr ipt ions of Four New Species , Proc . U . S.

Nat . M us . , 108 : 2 1 9-267 .

Katayama, M . , 1 960 , F auna Japon ica Ser ran idae (P isces) , TokyoNews Serv ice, L td . , Tokyo .

Kemp , N . E . , 1 974a, Ameloblast ic Secretion in Calcifying Teethof Shar ks , E ighth In ter nat ional Congres s on E lect ron Microscopy, Can ber ra,

2 : 458-459 .

1 974b, U ltr astructure of Ameloblasts and Enamel in SharkTeeth , 32nd Annua l Proceedings of the E lect ron M icroscopySoc . Amer ica, C . J . A rceneaux Clai tor

’

s Publ . D iv . ,

Baton Rouge, pp . 76-77 .

and J . H . Park , 1974 , U ltr astructur e of the Enamel Layer inDeveloping Teeth of the Shark , Car char h inus men isor rah,

A r ch . Oral B io l . , 1 9: 633 -644 .

Kendal l , W . C . , and E . L . Go ldsborough , 191 1 , Repor ts on the

Scientific Resu lts of the Expedit ion to the T r opical Pacific, inChar ge of A lexander Agass iz , by the U . S. Fish Comm i ss ionSteamer A lbatr oss ,” fr om August , 1899 , to Mar ch , 1 900 ,

Commander Jeffer son F . Moser , U . S. N . , Commanding ,XIII, the Shor e Fishes , Mem . M us . Comp . Zob i . , 26: 241 -343 .

Lewis , A . G . , 1 968 , Copepod Crus taceans Paras i tic on Fishes of

En iwetok Ato l l , Proc . U . S . Nat . M us . , 125 : 1 -78 .

Losey , G . S. , 1 971 , Commun icat ion Between Fishes in a Cleaning Symbios is , Aspects of the B io logy of Sym bios is , T . C.

Cheng Un iver s i ty Park Pr ess , Baltimore, pp . 45-76 .

1 972 , Predation Protect ion in the Po ison-Fang B lenny ,Meiocon thus at r odor sol is , and Its M im ics , Ecsen ius bico lor and

Runu lo loudondus (B lenm idae), Poc . Sci . , 26: 1 29- 139 .

1 974 , Asp idon tus taen iatus : Effects of Increased Abundanceon Clean ing Symbios i s w ith Notes on Pe lagic D i sper s ion and

A . f i lamen tos us (P isces , B lenn i idae), Z . Tierp sychol . , 34:

430-435 .

322 RANDALL AND RANDALL

1 975 ,Meiocon thus atrodor sol is : Field Ev idence of Pr edation

Protection , Copeia, 3: 574—576 .

Matsuur a, K . , 1 984 , A L is t of the Fishes Co l lected in the T rukIs lands , Ponape , and Majuro Ato l l fr om June thr ough August ,1 982, Proc . Japan . Soc . Sys . Zoo l . , 27: 14 1 - 1 48 .

and T . Sh im i zu , 1 982 ,The Squi r r elfish Genus Adioryx , a

Jun ior Synonym of Sargocen tron , Japan . Jour . Ichth . , 29:93 -94 .

Mccosker , J . E . , 1 970 , A Rev iew of the Eel Gener a Lep tenchelysand M u raen ichthys , with the Descr ipt ion of a New Genus ,Sch ismo r hynchu s , and a New Species , M u raen ichthys

ch i lens is , Poc . Sc i . , 24: 506-516 .

1 977 ,The Osteo logy , Class i fication , and Relationships of the

Eel Fam i ly Ophichthidae, Proc . Ca l if . Acad . Sc i . , 4th Ser . , 41 :

1 - 1 23 .

K . H atooka, K . Sasaki , and J . T . Mayer , 1 984 , JapaneseMoray Eels of the Genus U r op teryg ius , Japan J . Ichthyol , 31 :26 1 -267 .

and R . H . Rosenblatt , 1 975, The Mo ray Eels (P isces :Muraen idae) of the Galapagos Is lands , wi th New Records and

Synonym ies of Extr al im ital Species , Proc . Cal if . Acad . Sci . ,

4th Ser ., 40 : 41 7-427 .

Mees , G . F . , 1962 , A Pr el im inar y Rev is ion of the Be lon idae,

Zoo l . Ver handel ingen , 54 : 1 -96 .

Munz , F . W . , and W . N . McFar land, 1 973 , The Sign ificance of

Spectr al Pos i tion in the Rhodops ins of T ropica l Mar ine Fishes ,Vis ion Res .

, 13: 1 829- 1 874 .

Nelson , D . R . , 1 982, Aggress ion in Sharks : Is the Gr ay ReefShar k D ifferent?, Oceanus , 24: 45-55 .

and R . H . Johnson , 1 972, Acous tic Attr action of PacificReef Sharks : Effects of Pulse Interm i ttency and Var iabi l ity ,Comp . B iochem . Phys io l . , 4ZA : 85 95 .

No lan , R . S. , 1 974, Ar t ificial Reefs as Exper imental Too ls ,Pr oceedings of the In ter nat iona l Conference on A r t if icial Reefs ,Laur a Clounga and R ichard Stone Texas A MU n iver s i ty , TAMU -SG -74- 1 03 .

R . R . McConnaughey, and C. R . Stear ns , 1975 , Fishes Inhabi ting Two Small Nuclear Tes t Cr ater s at Enewetak Atol l ,Mar shal l Is lands , M icr ones ica, 1 1 : 205

-2 1 7 .

Odum , H . T . , and E . P . Odum , 1955 , T r oph ic Structur e and Pr o

duct iv i ty of a W indward Cor al Reef Commun ity on En iwetok,Eco l . Monog r , 25 : 291 -320 .

Par in , N . V . , 1 960 , The F lying Fi shes (Exocoetidae) of the

Nor thwes t Paci fic, (in Russ ian ; T rans lation IPST Cat . No .

6 1 8 , Trudy Ins t . Okeonol . , Akad . Nouk SSSR . , 31

205-285 .

1 967 , Rev iew of Mar ine Needlefi shes of the West Pacificand Indian Oceans (in Rus s ian ; NMFS T rans lation No .

Tr udy Ins t . Okeonol . , Akad . Nouk SSSR ., 84: 3

-83 .

B . B . Co l lette, and Y . N . Schcherbachev , 1980 , Pr el im inaryRev iew of the Mar ine H alfbeaks (H em i ramph idae, Belan iform es ) of the T ropica l IndoWes t Pacific, Trans . P . P .Shi r shov In s t . Oceanol . , 97: 7

- 173 .

Paulson , A . C. , 1 978 , On the Commensa l H abits of Ptereleotri s ,

Acan thurus , and Zebr asomo with Fossor ial Valencienneo and

Amblygobiu s , Cope ia, 1 : 1 68- 1 69 .

Pomeroy , L . R . , and E . J . Kuenz ler , 1 967 ,Phosphorus Tur nover

by Coral Reef An imals , Sympos ium on Radioeco logy , D . J .Nelson and F . C. Evans Ann Ar bor , Mich . , pp.474-482 .

Randal l , J . E . , 1 955a, Fishes of the Gi lber t Is lands , A to l l Res .

Bul l . 47: x i 243 .

1955b, A Rev is ion of the Surgeon Fi sh Genus Ctenochaetus ,

Fam i ly Acanthur idae, with Descr iptions of F ive New Species ,Zoo log ico, 40 : 1 49- 1 66 .

1 955c , A Rev is ion of the Surgeon Fish Genera Zebrasomoand Par ocan thu r us , Poc . Sc i . , 9: 396-41 2 .

1 956 , A Rev is ion of the Surgeon Fish Genus Acan thurus ,Pac . Sci . , 10: 1 59-235 .

1 963a, Rev iew of the H awkfishes (Fam i ly Ci r r hi t idae), Pr oc .

U . 8 . Nat . M us . , 1 14: 389-451 .

1 963b, Notes on the Sys tematics of Par r otf ishes (Scar idae),wi th Emphas i s on Sexual D ich romatism , Copeia, 2: 225-237 .

1 964a, Notes on the Gr ouper s of Tah i ti , wi th Descr iption of

a New Ser r an id Fish Genus , Pac . Sc i . , 1 8: 28 1 -296 .

1964b , A Rev is ion of the Fi lefish Genera Aman ses and

Can ther hines , Copeia, 2: 33 1 -361 .

1 973 , Tah it ian Fish Names and a Prel im inar y Check l i st ofthe Fi shes of the Society Is lands , Occ . Pap . B . P . B ishopMus . , 24: 1 67-2 14 .

1 976 , The Endem ic Shor e Fi shes of the H awai ian Is lands ,Lord H owe Is land and Eas ter Is land , Co l loque Commer son1 973

, Trav . et Doc . , 47: 49-73 .

1 977 , Contr ibution to the B io logy of the Wh i tetip ReefShark (Tr iaenodon obes us ), Pac . Sc i . , 31 : 1 43- 1 64 .

l 978a, Mar ine B io log ical and Ar cheo log ica l Expedition to

Southeast Ocean ia, Nat l . Geo . Soc . Res . Rep ., 1 969 Pr of ,

pp . 473-495 .

1 978b, A Rev is ion of the Indo-Paci fi c Labr id Fi sh GenusMocropharyngodon , with Descr iptions of Five New Species ,Bu l l . Mar . Sc i . , 28 : 742

-770 .

1 980a, A Sur vey of Ciguatera at Enewetak and B ikin i ,Mar shal l Is lands , wi th Notes on the Systematics and FoodH abits of Ciguatox ic Fishes , F ish . Bu l l . , 78: 20 1 -249 .

l 98ob, Two New lndo-Paci fic Labr id Fishes of the GenusH ol ichoeres , wi th Notes on Ot her Species of the Genus , Poc .

Sc i . , 343 415-432 .

1 98 1a, Underwater Guide to H awai ian Reef Fishes ,H ar r owood Books , Newtown Squar e, Pennsy lvan ia, and

T r easur es of Natur e, Kaneohe, H awai i .1 98 1 b, L uzon ichthys ear lei a New Species of Anthi ine Fishfr om the H awai ian Is lands , F r eshw. Mar . Aquar ium ,

4: 1 3- 1 8 .

1

1 981 c, Two New Species and Six New Records of Labr idFishes from the Red Sea , Senckenberg Mar

-i t,1 3: 79- 1 09 .

1 98 1 d, Palett e Surgeonfi sh Parocan thu r us hepotus , F reshw.

MarnAquor ium ,

4: 48-49, 59.

1 982 ,

“

A Rev iew of the Labr id Fi sh Genus H ologymnos u s ,

Rev . fr . Aquar io l . , 9: 13-20 .

1 983 , Rev i s ion of the lndo Pac i fic Labr id Fish GenusWetmorello, Copeia, 4: 875

-

883 .

1 985 , On the Val idity of the Tet r adon t id Fish A rothronman i len s is (Pr océ ), Japan . Jour . Ichth . , 32: 347-354 .

1 986, 1 06 New Records of Fishes fr om the Mar shal l Is lands ,Bu l l . Mar . Sc i . , 38 : 1 70-252 .

M . L . Bauchot , and M . Desou tt er , 1 985 , Chrom is vi r idis

(Cuv ier , the Cor r ect Name for the lndo Pac if ic Damselfi sh Prev ious ly Known as C . caeru lea Cuv ier , 1 830 (P isces :Pomacentr idae) , Cybium , 9: 41 1 -413 .

l

andA. Ben-Tuv ia, 1983:.

A Rev iewof the Grouper s (Pisces :Ser r an idae: Ep inephel inae) of the Red Sea, wi th Descr iptionof a New Species of Cephalophol is , Bul l . Mar . Sci . , 33 :

373 -426 .

and J . E. Bohlke, 1 98 1 , The Status of the Cardinal fi shesApogon evermann i and A . on isolep is (Per ci for rnes : Apogon i

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324 RANDALL AND RANDALL

1961 , Fishes of the Fam i ly Apogon idae of the WesternIndian Ocean and the Red Sea, Ichth . Bu l l . , Depar tm en t ofIch thyo logy , Rhodes Un iver s i ty, No . 22, pp . 373 -4 18 .

Sm ith , R . L . , and A . C . Paulson , 1 974 , Food Trans i t T imes and

Gut pH in Two Pacific Par r otf ishes , Copeia, 3 : 796-799 .

and A . C . Paulson , 1975 , Car bon ic Anhydrase in Some

Coral Reef F ishes : Adaptation to Car bonate Inges tion?,Comp . B iochem . Phys io l . , 50A : 1 3 1 - 1 34 .

Sm i th-Van i z , W . F . , 1976 , The Saber -Too thed B lenn ies , T r ibeNemophin i (Pi sces : B lenn i idae) , Acad . Not . Sc i . Ph i lo.

Monogr . , 1 9: v i i 1 96 .

and J . E. Randal l , 1 973 , B lennech is f i lamen tosu s Valenci

ennes , the Pr ejuven i le of Asp idon tus taen iot u s Quay and

Gaimard (Pi sces : B lenn i idae), Notu lae Natur ae, No . 448 ,

pp . 1 - 1 1

and V . G . Spr inger , 1 97 1 , Synops is of the T r ibe Salar nn i ,with Descr iption of Five New Genera and Three New Species(Pisces : B lenn i idae) , Sm i thson . Con t r . Zoo l . , No . 73 ,

pp . 1 -72 .

and J . C . Staiger , 1973 , Compar at ive Rev i s ion of

Scomber oides , O ligop l ites , Parono, and H ypacan thus withComments on the Phy logenetic Pos i tion of Compogr amma

(Pi sces : Carang idae) , Proc . Cal if . Acad . Sci . , 4th Ser . , 39:

1 85-256 .

Spr inger , V . G . , 1 967 , Rev i s ion of the Ci r cum tropica l ShorefishGenus En tomocrodus (B lenn i idae: Salar i inae) , Pr oc . U . S .

Nat . Mu s . , 1 22: 1 - 1 50 .

1 972, Synops i s of the T r ibe Omobr anchin i with Descr iptionsof Thr ee New Genera and Two New Species (P isces : B lenn iidae) , Sm i thson . Con t r . Zoo l . , No . 1 30 , pp . 1 -3 1 .

1 982 ,Pacific Plate B iogeog r aphy , wi th Special Refer ence to

Shor ef ishes , Sm ithson . Con t r . Zoo l . , No . 367, pp . iv 1 82 .

and W . F . Sm ith-Van i z , 1 968 , Systemat ics and D i s tr ibut ionof the Monotypic Indo -Pacific B lenn i id F ish GenusA t rosalor ias , Proc . U . S. Not . M us . , 1 24: 1 - 1 2 .

Strasburg , D . W . ,1 953 , F ishes of the Souther n Mar shal l Is lands ,

M imeo Repor t to Off ice of Naval Research , U . S. Navy , pp .

i i 267 .

1 966 , Obser vations on the Eco logy of Four Apogon idFishes , Pac . Sc i . , 20: 338 -341 .

1 967 , G unnell ichthys monos t igma and Ecsen ius bico lor , NewFi sh Reco rds fr om the Mar shal l Is lands , COpeio, 4: 839-841 .

and R . W . H iatt , 1957, Sexual D imo rph ism in the Labr idFi sh Genus , Gomphosu s , Pac . Sci . , 1 1 : 1 33- 134 .

Swerdloff , S. N . , 1 970 , Behav ioral Obser vations on En iwetokDam selfi s hes (Pomacentr idae: Chrom is ) wi th Special Reference to Spawn ing of Chr om is caer u leus , Copeia, 2: 37 1 -374 .

Tester , A . H. , 1 963 , O lfaction , Gus tation and the Common

Chem ica l Sense in Sharks , Shar ks and Sur v iva l , P . W . Gi lber tD . C. H eath Co . , Boston , Ch . 8 , pp . 255-282 .

and S. Kato , 1 966 , V isual Target D iscr im ination in B lackt ip Sharks (Car charhinus melanop ter us) and Gr ey Sharks(C . men iso r rah) , Poc . Sc i . , 20: 46 1 -471 .

Van Campen , W . G . , 1 950 , Po i sonous F ishes of the South Seas ,Spec . Sci . Rep . , U . 8 . Dept . In ter ior , Fish Wi ldl . Ser v . ,

Fisher ies , No . 25 , pp . 12 21 (T r ans lated fr om the JapaneseLanguage) .Wass , R . C . , 1984 , An Annotated Check l is t of the Fishes of

Samoa , NOAA Tech . Rep . , Not . Mar . Fisher ies Ser v. , Spec .Rep . Fish . , No . 78 1 , v 43 .

Webb , K . L . , and W . J . W iebe, 1 975, N i tr ification on a CoralReef, Can . J . M icrobio l . , 21 : 1 427- 143 1 .

Wh itehead , P . J . P . , and W . Ivan tsoff , 1 983 , A ther ino lacunosaand the Fishes Descr ibed by J . R . For ster , Japan . Jour .

Ichth . , 29: 355-364 .

W iebe, W . J. , R . E . Johannes , and K . L . Webb, 1 975 , N i trogenFixation in a Co ral Reef Commun ity , Sc ience, 188 : 257-259 .

Wood land , D . J.,Rev is ion of the Rabbi tf i shes , Fam i ly Sigan idae,

Indo -Pacif ic Fishes , in pr ess .

Woods , L . P . , and P . M . Sonoda, 1 973 , Fam i ly H o locentr idae,

Fi shes of the Wester n Nor th At lant ic, Par t 6 , Mem . Sear s

Found . Mar . Res ., No . 1 , pp . 33 1 -386 .

R eptiles of Enewetak Ato ll

JANET O . LAMBERSON

U . 8 . Envi r onmen tal Pr otection AgencyH atf ield Mar ine Sc ience Cen ter , Newpor t, Or egon 97365

Seven species of l i zar ds and one species of bl ind snakecompr ise the known ter r es tr ial her petofauna of EnewetakAto l l , based on specimen s in the U . S. Nat ional Museumof Natur al H is tor y , the B . P . B ishop Museum , the Amer ican Museum of Natu ral H i s tor y , the Museum of Comparat ive Zoo logy , and the M id-Paci fic Resear ch Labor ator y collect ions (Table Spec ies found on near by ato l ls may alsobe pr esen t on Enewetak ; however , no add i t ional specieshave been col lected from ther e. No amph ibia or sea

snakes ar e known fr om Enewetak .

None of the species of r ept i les known fr om Enewetakis endem ic to M icr ones ia. Pr esumably , Gehyr a ocean ica,

L ip in ia noctua, and Emo lo cyanura or iginated in PapuaNew Gu inea, and H em iphyllodoctylus typus and Lep idodacty lus lugubr is evo lved in souther n As ia (Bur t and Bur t ,1 932 ; B r own , All ar e now w idel y d is tr ibutedamong the is lands of the Pac i fic (McCoy,H em idacty lus fr enotus and Rampho typhlop s br am ina are

known fr om tropi cal As ia, Ind ia, and Afr ica and have beenintr oduced to Cen tral Amer ica and the i s lands of the

Pac i fic and Ind ian Oceans , includ ing H awai i (H unsaker ,1 966 ; H unsaker and B r eese, 1 967 ; Bus tar d , 1 970 ;McKeown , 1 978 ; McCoy, Var anus ind icus pr obablyor ig inated in the lndo-Aus tr al ian ar ch ipelago or Papua NewGu inea and has been in ten t ional ly intr oduced to a numberof the Mar iana, Caro l ine, and Mar shal l Is lands (F isher ,

Th is species i s extens ively d is tr ibuted on Pac i ficIs lands , includ ing the eas ter n par t of the Indo-Aus tral ianar ch ipelago , Papua New Gu inea, nor ther n Queens land ,B ismar ck ar ch ipelago , Solomons , T robr iands , Caro l ines ,Mar ianas , and the Mar shalls (R . Cr ombie, USNM ,

per sonalcommun i cat ion ) . Keys for iden t i ficat ion of the r epti les of

Enewetak , d is tr ibut ion r ecor ds , notes on natural h is tor y ,and excel len t co lor i l lus tr at ions may be found in bookletsby McKeown (1 978) and McCoy

The mos t often seen l i zar d on Enewetak is the housegecko , H em idacty lus f r enotus , wh ich fr equents ar eas of

human occupat ion . I t is common on or ins ide bu i ld ings ,especial l y near l igh ts , or on w indows at n ight wher e i t

Chapter 28

s talks in sects attracted by the l ight . It is also found underscr ap metal , plant debr i s , and dr i ftwood or other debr is onthe beach .

H em idacty lus f r enotus i s an aggress ive species thatmay r eplace other res iden t spec ies of geckos when i t is

introduced to a new area. In H awai i i t i s r epor ted to havereplaced H em idacty lus gar not i (McKeown , 1978) and pos s ibly also Lep idodoctylus lugubr is to some extent (H unsakerand B reese, McKeown (1978) r epor ts that L .

lugu br is , which also frequen ts ar eas of human habi tat ion , is

s t i l l qu i te abundan t in H awai i . On Enewetak Ato l l , L .

lugubr is is common on Enjebi Is let but i s in frequentl y collected r elative to the co l lect ion of H . f r enotus on EnewetakIs let (Table H em idacty lus fr enotus is h ighly voca l , and inencoun ter s w i th other geckos , i t fr equen tl y em i ts a ser ies

of five to s ix ch i rp ing ca l l notes . If attacked , i t em i ts a

squeak . An extens ive di scuss ion of vocal i zat ion in th i s spe

cies i s given by Mar cell in i The female H . f renotuslays two r ound wh i te eggs in cr ev ices , and the eggs or

adults may be in troduced to new areas as s towawaysamong mater ials or equ ipmen t moved among i s lands(McKeown ,

The mour n ing gecko , Lep idodoctylus lugubr is , is foundon bu i ld ings ; tr ees and bushes ; among coconut , pandanus ,and other plan t debr is in ar eas occup ied by people; inopen for es t ; and among beach dr i ftwood . Th is species ispar thenogen ic (McKeown , and i t s h ighly adhes iveeggs are eas i ly tr anspor ted among goods car r ied by boat .Th i s gecko has been w idely d is tr ibuted ar ound the Paci ficand is known fr om several is lets on Enewetak Ato ll (Table

It s cal l has been compared to the sound of two pebblesbeing hi t together (Mar shal l , I t somet imes em i ts a

squeak when attacked by another gecko or captu red by aco llector .

The tr ee gecko , H em iphyllodoctylus typus , is repr e

sen ted fr om Enewetak by on l y two spec imens in the

USNM col lect ion . Th is gecko , however , i s r epor ted to havea qu ick escape react ion (O l iver and Shaw , 1 953) and to bewar y and ag i le (H unsaker and B reese, Therefo re, i t

may be more abundan t on Enewetak than ind icated by thefr equency of co l lect ion . In H awai i i t i s found in low den

s tty in fo r es ted areas , r ock and wood pi les , occas ional ly onthe s ides of bu i ld ings and on tr ee tr unks at n igh t , and

under the bases of coconut palm fr onds . I t is not gener all y

326 LAMBERSON

TABLE 1

Checkl i s t of Ter r es tr i al Rept i les of Enewetak A tol l‘

Phylum CH ORDATAClas s REPT IL IAOrder SQUAMATASubo rder SAUR IA (L i zards )Fam i ly GEKKON IDAE

“

gecko”

BPBM— Bokombako Is let : Dec. 30 , 1 950 , under loose canvas , Y . Osh i r o , 1 spec . , 0695 .

Gehyr a ocean ica (Les son)USNM— B iken Is let :USNM — Elugelab Is let :USNM— l kur en Is let :

BPBM — Ananij Is let :BPBM— lku ren Is let :

MPRL — Japtan Is let :

H em idacty lus fr enotu sDum é r i l and B ibran

USNM — Enewetak Is let :

USNM— Medr en Is let :USNM - Run i t Is let :

BPBM — Enewetak Is let'BPBM — Japtan Is let :BPBM — Medren Is let :

BPBM — Run i t Is let :

MPRL — Enewetak Is let :H em iphyllodoctylus typus B leekerUSNM — Alembel Is let : Apr i l 1 7 , 1 977 ,

in r oof, P . B . Lamber son and J . Lamber son , 2 spec . , 205534— 35 .

Lep idodoctylus lugubr is

(Dumé r i l and B ibran )USNM — A lembel Is let : July 30 , 1 966 , W . B . Jackson , 1 spec . , 1 97844 ;USNM— B iken Is let : May 25 , 1 946 , J . P . E . Mor r ison , 2 spec . , 124078 — 79 ;

Aug . 1 , 1 966 , W . B . Jackson , 1 spec . , 197843 ;USNM — B i llae Is let : Aug . 24 , 1 968 , L . N . H uber , 3 spec . , 197876 ;USNM — Boken (l r ene) Is let : Aug . 24 , 1968 , L . N . H uber , 2 spec. with 1 7 eggs , 197875 ;

Feb. 28 , 1 977, on bunker , J . L am ber son , 3 spec. , 205538 — 40 ;

‘W i th co l lection data for specimens r ecorded in the co l lections of the U . 5 . Nat ional Museum of Natural H istory (USNM) , theB . P . B i shop Museum (BPBM ), the Amer ican Museum of Natur al H i stor y (AMNH ) , and the M id-Pacific Resear ch Labo r atory (MPRL ).

(Th is table cont inued on next page. )

May 25 , 1 946 , J . P . E . Mor r ison , 2 spec . , 1 24076— 77;

1948 , D raeger , 7 eggs , 1 28000 ;

Aug . 21 , 1 976 , in Pison ia forest at n ight , P . B . Lamber son , 1 spec . , 205527 ;Aug . 21 , 1976 , E . Reese, 1 spec . , 205528 .

Dec. 25 , 1 965 , under loose bar k of fal len coconut tree, K . J . Frogner , 1 spec. , 4033 ;June 28 , 1 964 , in chela of crab , Geogr aps us cr in ipes (sti l l al ive), in coconut grove,

C. Ber r y , 1 spec . , 4028 ;Dec . 29 , 1 965 , K . J . Frogner , 1 spec . , 4034 ;Jan . 25, 1967 , K . J . Frogner , 20 spec. , 4273 , 4279

— 97 ;

July 1 5 , 1967 , on Pison ia at n ight , E . 5 . Reese, 1 spec . , 400 1 .

Jan . 1 1 , 1976 , J . Lamber son , 2 eggs ;

March 7 , 1 976 , around bui ldings near lagoon , F . C. Rabelais , 1 spec .

March 5 , 1 977 , trai ler park area, J . Lamber son , 1 spec . , 205529 ;Mar ch 1 2 , 1 977 , on lawn fur n iture, tr ai ler park area,

P . B . Lamber son and J . Lamber son , 2 spec . , 205530— 3 1 ;

March 1 3 , 1977, on lawn under Cocco loba tr ee,trai ler park area,

P . B . Lamber son , 1 spec . , 205532 ;Mar ch 1 7 , 1 977 , under piece of cor r ugated metal , J . Lamber son , 1 spec . , 205533 ;Aug . 1 964 , W . B . Jackson , 3 spec . , 1 97841 ;July 24 , 1966 , W . B . Jackson , 1 spec . , 197842 ;Sept . 1 , 1 968 , L . N . H uber , 2 spec . , 1 97874 .

Jan . 22, 1 967 , K . J . Frogner , 1 spec . , 4247;Feb. 2, 1 967 , K . J . Frogner , l spec . , 4253 ;June 22, 1964 , K . J . Frogner , 2 spec . , 4046 — 47 ;

Aug . 8 , 1 964 , K . J . Frogner , 2 spec . , 403 1 and 4078 ;Aug . 8 , 1964 , in 10 ft coconut trees at edge of lagoon beach , K . J . Frogner ,3 spec . , 4048— 50 ;

Dec . 30 , 1 965 , large crumpled tarp on seaward beach , K . J . Frogner , 1 spec . , 4064 ;Dec . 30 , 1965 , K . J . Frogner , 27 spec . , 4065— 67 , 4079— 4 100 , 4200— 0 1 ;

Dec . 30 , 1 965 , under pi le of sheet t in on cement tent base, K . J . Frogner , 3 eggs , 6948 ;

Jan . 22 , 1 967, K . J . Frogner , 21 spec . , 4202 and 4227— 46 ;

July 5 , 1964 , in smal l pi le of rusted sheet metal , K . J . Frogner , 1 spec . , 4021 ;Aug 29, 1964 , in wood and tarp pi le on seaward beach , K . J . Frogner , 1 spec . , 4044 ;Aug . 29 , 1964 , in dr i ftwood on seaward beach , K . J . Frogner , 1 spec . , 4045 .

March 7, 1976 , ar ound occup ied bui ldings , F . C . Rabelais and J . Lamber son , 2 spec .

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328 LAMBERSON

TABLE 1 (con t'd)Fam i ly VARAN IDAEVaranus indicus (Daudin)USNM— Japtan Is let : June 7, 1947, J . P . E. Mor r i son , 2 spec . , 1 241 1 2— 13 .

AMNH — Japtan Is let : A . A . V incenzi , 1 spec . , 78994 .

MPRL — Japtan Is let : Sept . 15 , 1975, in open fi eld, P . B . Lamber son and J . Lamber son , 1 spec .Suborder SERPENTES (Snakes )Fam i ly TYPH LOPIDAERamphotyphlop s br am ino (Daudin)BPBM— Enewetak Is let : Sept . 1 3 , 1 980 , MPRL lanai at night , S. lge, 1 spec . , 6953 .

MCZ— Medren Is let : June 1954 , C. Ok ino and A . Souza , 1 spec . , MCZ 53782 .

Order TESTUD IN ES (=CH ELON IA)(Tur t les )

Suborder CRYPTOD IRAFam i ly CH ELON IDAEChelon ia mydas(L innaeus , 1 758)— Green tur t le.

Eretmochelys im br icoto(L innaeus , 1 766)- H awksbi ll tur t le.

found in assoc iat ion w i th man (H unsaker and B reese,

1 967 ; Oliver and Shaw , 1 953 ; McKeown , The

female produces two eggs wh ich adher e to sur faces(McKeown ,

The large Po l ynes ian gecko , Gehyr a ocean ica, is foundin the fores t on the leaves or tr unks of coconut , P ison ia, or

Pandanus tr ees or under fal len logs or other debr i s on the

gr ound (H eatwo le, I t can be spo tted at n ight by theway i t s eyes sh ine red when touched by l ight . It s call i s a

loud “

K r aaaaaaa”

(Mar shal l ,The blue-tai led sk ink , Emo ia cyanura, is common under

coconut or other plan t debr is , under dr i ftwood , and amongscr ub vegetation at the top of is land beaches , especiallyamong the v ines of the beach mor n ing g lor y Ipomoea pescap r ae (Sachet , These sk inks are r epor tedly poo rcl imber s and ar e act ive pr imar i ly dur ing the day (Moul ,1 954 ; Mar shal l ,The moth sk ink , L ip in ia (L ygosoma or Leio lop isma)

noctua, is fai r ly common on Enewetak Ato l l among leafl i tter and coconut debr i s . It is essent iall y d iur nal and maybe par t ly ar bo real (McCoy, Oliver and Shaw( 1 953 ) repor ted that they had co l lected L . noctua on Run i t

Is let on Enewetak Ato l l on the g r ound among “ her baceousvegetat ion along the beach dunes ”

and that specimensthey co l lected had “

two or more incomplete d igi ts on the

feet . They speculated that the m is s ing d ig i ts m ight havebeen sheared off by ter res tr ial herm i t crabs wh ich are com

mon on the i s let or by other l i zar ds du r ing fights . Rats arealso common on Run i t and may have been r espons ible forsome of the m i ss ing d ig i ts . M iss ing toes among specimen sof L . noctua in H awai i have been attr i buted to the l i zar ds ’

char acter is t i c r o ll ing escape behav ior (McKeown ,

L ip in ia noctua br ings fo r th one to fou r young al ive fr omeggs r etained in the body (McKeown ,

The larges t l i zar d on Enewetak Ato l l is the mon i torl i zar d , Varanus indicus , a nat ive of Aus tral ia and PapuaNew Gu inea wh ich was in tr oduced to Guam , the Mar iana,

Carol ine, and Mar shal l Is lands by the Japanese for r at con

tr o l or as a food sou r ce (F i sher , 1 948 ; Mar shal l ,

Th is species has been col lected on Enewetak Ato l l on lyfrom Japtan Is let , and i t appar ent ly feeds on land cr absand some mar ine l i fe. Spec imens fr om Enewetak Ato l l arecommon l y about 1 m in lengt h .

The B r ahm iny bl ind snake, Ramphotyphlops bram ino(Daud in) (Typhlops or Typhl ina br am inus ), is a smal lcm) , secr et ive bur row ing snake nat ive to the Ph i lip

pines and southeas t As ia. The spec ies has been introducedin to new ar eas in the d i r t accompanying transpor ted plan tsand equ ipmen t and is now w idel y d is tr ibuted among majoris land groups of the Pac i fic and Ind ian Oceans . It is foundin Afr ica, Aus tr al ia, Ind ia, and Mexico (Brown , 1 957;McCoy, In M icr ones ia i t is found on Guam , T in ian ,

Saipan , and Enewetak (Bur t and Bur t , 1 932; Cagle, 1 946 ;Kn ight , It may have been in troduced to Enewetakw i th plan tings fr om H awai i , where i t was in itial ly obser vedin 1 930 (Slev in , 1930 ; Fisher , 1 948 ; O l iver and Shaw ,1 953 ; H unsaker and Br eese,

B l ind snakes ar e secr et ive, noctur nal , and general lyseen abr oad on l y fo l low ing a heavy r ain . They are found inloose, mo is t so i l in or under r o tt ing logs and other debr i sand in gar dens under s tones , plas t ic cover , and po ttedplan ts (Cagle, 1 946 ; McKeown , 1978 ; McCoy, Two

specimens have been co l lected on Enewetak Atol l : one on

Medr en in 1 954 (Kn ight , 1984) and one in 1 980 on

Enewetak Is let on the MPRL lanai at n ight . I t r emains tobe seen whether any mor e specimens w i l l be found on theato l l or whether the species can sur v ive in the is land ’

s relat ively dry, sandy so i l . B l ind snakes ar e par thenogen ic , andthe s ingle female par en t lays two to eight elongate eggs in

mo is t so i l (McKeown , A populat ion may thus bees tabl ished fr om a s ing le in troduced ind iv idual . B linds nakes are benefic ial to man because they eat term i tes and

other soft -bod ied insects and insect lar vae (McKeown ,

1 978 ; Mccoy ,The r elat ively dry cl imate at Enewetak makes i t a

rather har sh env i ronment for ter res tr ial r ept i les . Some of

the spec ies found ther e are adapted to liv ing wherehumans l ive and may have been transpor ted there by man

REPTILES 329

acc idental ly or intentional ly (H em idactylus , Lep idodoctylus ,Var anus , Ramphotyphlops) . Some species inhabi t beachareas and are found among dr i ftwood (L ip in ia, Emo ia,

H em idacty lus , Lep idodoctylus) . They may have beenwashed ashor e on the atol l fr om other ar eas on float ingdebr is or come as s towaways on the boats of ear l y interis land tr aveler s . Gehyr o and H em iphyllodoctylus ar e pr i

mar i ly res tr icted to fores t areas and may have been r aftedto the ato l l fr om other is lands by

.s to rm waves wh ich some

times wash over low - ly ing ar eas and can d i s lodge largetr ees and fal len debr is .

Enewetak Ato l l has few spec ies of r ept i les compared tolar ge, high is lands such as H awai i . Due to i ts dry cl imate,

lack of extens ive fores t ar ea , and relat ive isolat ion , i t alsohas fewer species than some other ato l ls . Sys temat icrepor ts on the ter r es tr ial r ept i les of ato l ls ar e rar e and

often ar e based on chance obser vat ion Mar shal l ,1 95 1 ; Moul , 1 954 ; Sachet , Spec imens d is cus sed inthese r epo r ts and many of the spec imens co l lected fr omEnewetak were found by chance by people wor k ing on

other pr ojects . K . J . Frogner took advan tage of l im i tedoppor tun i t ies to v i s i t the var ious is lands of Enewetak Ato l lto co llect as many spec imens as poss ible, and much of

what is known about the habi tat of the l i zar ds on the ato l li s based on hi s wor k . H owever , a more thor ough sear ch ,especial l y on the for es ted is lets , could r eveal mor e speciespr esent . Based on spec imens in the USNM col lect ion fromnear by Pac i fic is lands and other atol ls in the Mar shal lIs lands , the fo l low ing spec ies of l i zar ds poss ibly could befound on Enewetak Ato l l (R . Cr omb ie, USNM ,

per sonalcommun icat ion) :Gehyra mut i lata w idespr ead in Ocean iaLamp r olep is smar agdina w idespread in Ocean ia

(found on Ai luk , Ar no , Jalu i t , L ikiep , Taka,

Ujae, Ujelang , U t i r ik , and Wotho Ato l ls in theMar shal l Is lands)

Cryp toblephor us bouton i w idespr ead in Ocean ia(Pokak and Taong i in the Mar shal ls)

Emo ia baudin i group spott y in Ocean ia,

iden t i ficat ion doubtfulE . ar noens is Mar shal ls and pos s ibly Car ol ineIs lands (Ar no , B ik in i ? , Lae, Jalu i t , Ronger ik?)

E . boettger i or ien tal is spo tty (Ar no , B ik in i ? ,Ronger ik?)

E . m ivor t i spo tt y (Lae, Ujae,Wotho)

Cyr todoctylus pelag icus spo tty (Ar no Ato l l)Per ochi r us oteles spott y in Ocean ia but on Ar no Ato l lAccor d ing to George Balazs (per sonal commun icat ion) ,

on l y two species of sea tur t les — the green tur t le, Chelon iamydas , and the hawksb i l l tur tle, E r etmoche lysimbr icoto— are known to Enewetak Ato ll . Too few specimens have been examined to determ ine whether these twospecies belong to the subspecies C . mydas japon ica of the

wes ter n Paci fic or E . im br icoto squamoto of the IndoPaci fic . At leas t one incidence of a poss ible sea tu r tle nes ting on lku ren Is let has been obser ved (P . Lam ber son , per

sonal commun i cat ion and photo) .

REFERENCES

B r own , W . C . , 1 957, The D is tr ibut ion of Ter r es tr ial Repti les inthe Is lands of the Paci fic Bas in , in Pr oceedings of the E ighthPacific Science Congr es s , Vo l . IIIA , Un iver s i ty of the Ph i l ippines , Quezon City , Nov . 16— 28 , 1 953 , pp . 1479- 1 491 ,

Pacific Science Association , Quezon Ci ty , Ph i l ippines .

Bur t , C. E . , and M . D . Bur t , 1 932 , H erpeto logical Results of the

Wh itney South Sea Expedi tion , VI , Pacifi c Is land Amph ibiansand Rept i les in the Co l lection of the Amer ican Museum of

Natur al H i stor y , Bu l l . Amer . Mus . Nat . H is t . ,_

63 : 46 1 -585 .

Bus tard, H . R . , 1 970 , Activ ity Cycle of the T r opica l H ouseGecko , H em idacty lus fr enotus , Copeia, 173- 176 .

Cag le, F . R . , 1 946 , Typh lops bram inus in the Mar ianas Is lands ,Copeia, 10 1 .

Fisher , H . l . , 1 948 , Loca l i ty Records of Pacific Is land Repti lesand Amph ibian s , Copeia, 69 .

H eatwo le, H . , 1 975 , B iogeogr aphy of Repti les on Some of the

Is lands and Cays of Easter n Papua — New Guinea , A to l lResear ch Bu l l . No . 1 80, Paci fic Science Board , Wash ington ,

D . C.

H unsaker , D . , 1966 , Notes on the Population Expans ion of the

H ouse Gecko , H em idacty lus f renotus , Ph i lipp ine J . Sci . , 95:1 21 - 1 22 .

and P . B r eese, 1 967, H er petofauna of the H awai ian Is lands ,Pacif ic Sci . , 21 : 423-428 .

Kn ight , J . L . , 1984 , Typhl ina bram ino, H erp Review, 1 5: 1 1 5 .

Mar cell in i , Dale L . , 1 974 , Acoustic Behav ior of the Gekkon idL i zard, H em idacty lus fr enotus , H erpetolog ico, 30 : 44-52 .

Mar shal l , J . T . , 1 951 , Ver tebr ate Eco logy of Ar no Atol l , Mar shallIs lands , Ato l l Resear ch Bul l . N o . 3 , Paci fic Science Board,Washington , D . C.

Mar shal l , M . , 1975, The Natural H istor y of Namoluk Ato l l ,Easter n Car o l ine Is lands , A to l l Research Bu l l . No . 1 89, PacificScience Board , Wash ington , D . C.

McCoy, M . , 1980 , Rept i les of the So lomon Is lands , Wau Eco logyIns ti tute H andbook No . 7 , Papua New Guinea .

McKeown , S. , 1 978 , H awai ian Rept i les and Amph ibian s , The

Or iental Publ ish ing Company , H ono lu lu , H awai i .Mou l , E . T . , 1 954 , Prel im inar y Repo rt on Land An imals at Ono

toa Ato l l , Gi lber t Is lands , A to l l Resear ch Bul l . No . 28 ,Pacifi c

Science Board , Wash ington , D . C.

O l iver , J . A . , and C. E. Shaw, 1 953 , The Amphibians and Repti les of the H awai ian Is lands , Zoo logico, 65-95 .

Sachet, M . H . , 1962 , Geogr aphy and Land Eco logy of Cl ippertonIs land , A to l l Resear ch Bu l l . No . 86 , Pacific Science Board,Wash ingt on , D . C .

Slev in , J . R . , 1 930 , A Note on the Di scovery of the GenusTyph lops in the H awai ian Is lands , Copeia, 1 58 .

ACKNOWLEDGMENTSTh is r epo r t i s based in par t “

on obser vations made bythe author dur ing employmen t at the M id-Paci fic Resear chLabor ator y , wh ich is suppor ted by the D iv is ion of B iomedi

cal and Env i r onmen tal Resear ch of the Depar tmen t ofEnergy under con tract No . AT (26-1 )-628 to the Un iver s i tyof H awai i . Appr eciat ion is extended to R . Crom bie for hisin ter es t and help in ident i ficat ion of specimens and for

his cr i t ique of the manuscr ipt . Thanks ar e also extended toS. McKeown for many helpful sugges t ions and to

G . Balazs for information on sea tur tles .

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332 BERGER

TABLE 1 (con t’d)

Order ANSER IFORMESFam i ly ANAT IDAEAnas acuta L innaeus : Temme, MS; H ai lman , 1 979 .

Anos quer quedu lo L innaeus : H ai lman , 1979.

Or der GALL IFORMESFam i ly PH ASIAN IDAEGollus ga l lus (L innaeus ) : Amer son , 1969 .

Order CH ARADRIIFORMESFam i ly SCOLOPACIDAEL imoso lappon ica bauer i Naumann : Pear son and Knudsen , 1968 ; Carpenter et al . , 1 968 .

Numen ius phaeop us (Scopo l i ) : Woodbury , 1 962 ; Pear son and Knudsen , 1967.

Numen ius tohitens is (Gmel in ) : Woodbury , 1962 ; Pear son and Knudsen , 1967 .

‘Tr inga g lareo la L innaeus : H ai lman , 1979.

H eteros celus brevipes (V iei l lot) : Pear son and Knudsen , 1 967 ; Carpenter et ai . , 1 968 .

H eter osce lus incanus (Gmel in ) : Woodbury , 1962; Pear son and Knudsen , 1 967 .

Ar enar ia in terp res (L innaeus ) : Woodbury , 1962 ; Pear son and Knudsen , 1 967 .

Gal linago har dwicki i (Gr ay) : H ai lman , 1979 .

Calidr i s olbo (Pal las ) : Woodbur y , 1 962; Pear son and Knudsen , 1 967 .

Calidr is acum inata (H or s field) : Woodbur y , 1 962; Pear son and Knudsen , 1 967 .

Calidr is r uficoll is (Pal las ) : Johnson and Kienho lz , 1 975 .

Tryngi tes s ubr uficoll is (V iei l lot) : Pear son and Knudsen , 1 967 .

‘Ph i lomachus pugnax (L innaeus ) : Temme photogr aph ; H ai lman , 1 979.

Fam i ly CH ARADRI IDAECharadr ius dubius cu ron icus Gmel in : Baker , 1 951 ; Pear son and Knudsen , 1 967 .

Char adr ius mongolus Stresemann : H ai lman , 1979 .

P luvia lis dom in ica fu lva (Gmel in) : Woodbur y , 1962 ; Pear son and Knudsen , 1 967 .

P luvialis squatar ola (L innaeus ) : Baker , 1 951 ; Pear son and Knudsen , 1 967 .

Fam i ly LAR IDAEStem a poradisaea Pontoppidan : Woodbur y , 1962 .

Stem a s umot rana Raffles : Woodbur y , 1962; Pear son and Knudsen , 1 967 .

Ster na lunata Pea le: Woodbury , 1 962; Amer son , 1 969 .

Stem a fu scata L innaeus : Woodbury , 1962 ; Pear son and Knudsen , 1 967 .

Tholas seu s bergi i (L ichtenstein) : Woodbury , 1 962; Pear son and Knudsen , 1 967 .

P rocels tem o ceru lea (Bennett ) : Woodbur y , 1962; Pear son and Knudsen , 1 967 .

Anous s to l idus (L innaeus ) : Woodbur y , 1962 ; Pear son and Knudsen , 1 967 .

Anous tenu i r os tr i s (Br yan) : Woodbur y , 1 962 ; Pear son and Knudsen , 1 967 .

Gyg is olbo (Spar rman ) : Woodbur y , 1 962 ; Pear son and Knudsen , 1 967 .

Or der CUCUL IFORMESFam i ly CUCUL IDAEEudynam is taiten s is (Spar rman) : Woodbury , 1962 .

Or der STRIGIFORMESFam i ly STRIGIDAEAs io flommeus (Pontoppidan) : John son and K ienho lz , 1 975 .

‘New r ecord for Enewetak and nor thern Mar shal l Is lands .

REF ERENCES

Amer son , A . B . , Jr . , 1 969 , Orn i tho logy of the Mar shall and G i lber t Is lands . Ato l l Res . Bul l . No . 127, Sm i thson ian Ins t i tution , Wash ington , 348 pp .

Baker , R . H . , 1 951 , The Av ifauna of Micr ones ia, i ts Or igi n , Eva

lut ian and Di s tr ibution , U n iver s i ty of Kansas Pub . , M u s . ofNat . H is t .

, 3 : 1 -359 .

Carpenter , M . L . , W . B . Jackson , and M . W . Fall , 1 968 , B i rdPopulations at Enewetok Atoll , M icrones ica, 4: 295-307 .

H ai lman , J . P . , 1979 , Notes on B i rds of Enewetak Ato l l , Mar shal lIs lands , E lepaio , 40 : 879 0 .

Johnson , 0 . W . , and R . J . Kienho lz , 1 975 ,New Av i fauna]

Records fr om Enewetak , Auk , 92: 592-594 .

Pear son , D . L . , and J . W . Knudsen , 1 967, Av i fauna] Recordsfr om Enewetak Ato l l , Mar shal l Is lands , Condor , 69: 201 -203 .

Woodbur y , A . M . , 1 962 , A Rev iew of the Eco logy of EnewetakAto l l , Pacific Ocean , Un iver s i ty of U tah Press Ins t . of B io l .Res . , 123 pp .

M amma ls of Enewetak A to ll

ERNST S. REESE

Depar tmen t of Zoo logyUn iver s ity of H awai iH onolulu

,H awa i i 968 1 7

Man i s the dom inan t mammal of Enewetak Ato l l .H umans ar r ived at leas t 2000 year s ago (Cr aib , In

all pr obabi l i ty the Po l ynes ian rat , Rattus exu lans , and the

domes t ic p ig , Sus s crofa, accompan ied him . Domes t icdogs , Can is fam i l iar is , and cats , Fe l ix catus , mos t l ikel y didnot ar r ive un t i l Eu ropean contact in the 1 9th centu r y .

The ubiqu i tous common house mouse, M us m us culus ,and the roof rat , Ratt us r attus , apparen t ly ar r ived ear ly inthe 20th centu r y when foreign contacts wer e mor e fr e

quent . Ther e i s no ev idence of the Norway r at , Rattus nor

veg icus , on Enewetak , although i t occur s elsewher e in theMar shall Is lands (Jackson , Vessey , and Bast ian , Chapter 1 2of Vo lume I ) . A ll of these spec ies of rats and m ice are

cons idered as human commensals and have been spreadw idely thr oughout the wor ld thr ough the activ i t ies of man

(Eisenberg , 1 98 1 ; Nowak and Parad iso ,

Domes t ic cats , pigs , and goats , Cap r a h ir cus , wer e lefton Otdia

_(Wo tji i ) A to l l of the Ratak , or sun r ise chain , by

Otto von Kot zebue in 1 81 6 and 18 1 7 . There is , however ,no ev idence that goats ever l ived on Enewetak Ato l l ,wh ich l ies far to the nor thwes t in the Ral ik , or sunsetchain of .ato l ls . Kotzebue did no t v is i t th is chain unt i l hissecond voyage in 1 824 (Kot zebue,

Mar ine mammals are occas ional v is i tor s to EnewetakAto ll . U n for tunatel y there ar e on ly two pos i t ive ident i ficat ion s based on skeletal r emains , beached specimen s ,and photog raphs co l lected by Ph i l ip Lamber son , formerlabor ator y manager of the M id-Paci fic Mar ine Laborator y ,1 974- 1 977 , and sent to W i l l iam F . Per r in ,

Nat ionalMar ine F isher ies Ser v ice,

La Jo l la, Cal i for n ia,for

iden t i ficat ion .

The sp inner do lph in , Stenella long i r os tr is , i s iden t i f iedfrom photog raphs (F ig . 1 ) and by obser vat ion s of i t s di s

t inct ive spinn ing br each ing behav ior (Per r in ,wh i le

the s tr iped do lph in , Stenella coe r u leoo lbo , i s iden t i fied fr oma skeleton . The occur r ence of th i s spec ies at Enewetakr epresents a new recor d for the central Pac i fic (H ubbs , Perr in , and Ba lcomb , 1 973 ; W . F . Per r in , per sonal comm un icat ion to P . Lamber son ,

Chapter 3 0

A checkl i s t of mammals of Enewetak A tol l is pr ov idedin Table 1 .

Twenty-seven species of whales , dolph ins , and por

po ises may be expected to occur at Enewetak Ato l l (W . FPer r in ,

per sonal comm un icat ion) . These ar e l is ted below inthe or der they are presen ted in Leatherwood et ai .wh ich pr ov ides a gu ide to thei r iden ti fi cation .

Large wha les wi th a dor sa l f in :B lue whale, Ba laenopter a mus cu lusF in whale, Balaenopter a p hysc iasSei whale, Ba laenoptera bo r ea lisB ryde

’

s whale, Ba laenopter a eden i

H umpback whale, Megapter a novoeang l iae

Sperm whale, Physeter macr ocepha lusMedium -s iz ed who les with a dor sa l f in :M inke whale, Balaenopter a acu tor os tr ato

(Souther n bottlenose whale, H yper oodon sp7

Cuv ier ’

s beaked whale, Z iph ius covi ros tr is

Beaked whales of the genus Mesop lodonStejneger

’

s beaked whale,M . s tejneger i

B lainv i lle’

s beaked whale, M . dens i r os t r is

G inkgo-toothed beaked whale, M . g inkgoden s

K i ller whale, Or cinus o r coFalse k i l ler whale,

Pseudo r co cros s idens

Shor t-finned p i lot whale, Globicephala macror hynchusR isso

’

s do lph in , G rampus gr iseusSmal l wholes , do lph ins , and por po ises with a dor sal f in :Spo tted do lph in , Stenella attenuataSp inner do lph in , Stenel la longi r os t r is

Str iped do lph in , Stenella coer u leoolba

Common do lph in , De lph inus de lph isFr aser ’

s do lph in , Logenodelphis hosei

Bott lenose do lph in , Tur s iops tr uncatu sRough-toothed do lph in , Steno bredanens isPygmy k i ller whale , Fereso attenuataMelon -headed whale, Peponocephala electr aPygmy sperm whale,

Kog ia brev icep s

Dwar f sperm whale, Kog ia s im us

Other mar ine mammals , such as seals and sea l ions ,ar e not known to occur in the Mar shal l Is lands .

Accor d ing to Lamber son ,the Mar s hal lese r ecogn i ze

two k inds of porpo i ses . One is smal l and travels in lar ge

REESE

F i g . 1 Two v iews of the spinner dolph i n , Stenel la longi r os t r is . in the lagoon of Enewetak A tol l .[Photos by (a) Paul M . A l len and (b) Ph i l ip Lam ber son .]

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M is cellaneous Species R ecords ofEnewetak A to ll

RICH ARD H . TITGEN and BEATRICE L . BURCH

Ber n ice P . B is hop MuseumH ono lu lu , H awai i 968 1 7

IN TRODUCT IONThe pr eced ing chapter s in Volume I I ar e comp i lat ions

of spec ies recor ds fr om Enewetak Ato l l . For var iousr easons all s ys temat ic groups could not be covered in a

separ ate chapter . Consequen tly , th is final chapter includesm iscellaneous recor ds fr om the M id-Paci fic Resear chLabor ator y at Enewetak Ato l l and from the li terature.

TABLE 1

Check l i s t of M i s cel laneous Spec ies of Enewetak A tol l

K INGDOM PLANTAED iv is ion PYRRH OPH YTA

Class D INOPH YCEAEOrder PER ID IN IALES

Fam i ly H ETERAULACACEAEGambierdiscus tox icus Adach i and F ukuyo , 1979 : see Co l in Vol . I .

D iv i s ion CH LOROPH YTAClass CH LOROPH YCEAE

Order CLADOPH ORALESFam i ly CLADOPH ORACEAEC ladopho ra sp .

C ladophor a hem isphaer ica: Bai ley-B r ock et al . , 1980 .

O rder SIPH ONOCLADALESFam i ly VALON lACEAEValon ia t r abecu lato Egerod, 1 952 : Bai ley-B rock et al . , 1980 .

Valon ia ven t r icosa J . Agardh , 1 887 .

Valon ia ven t r icavo: Bai ley-B r ock et al . , 1 980 .

O rder DASYCLADALESFam i ly DASYCLADACEAEAcetabu lar ia clovoto Yamada, 1 934 : Bai ley-B rock et al . , 1 980 .

D iv is ion PH AEOPH YTAClas s PH AEOPH YCEAE

Order D lCTYOTALESFam i ly D lCTYOTACEAEPadina japon ica Yamada , 193 1 : Bai ley-B r ock et al . , 1980 .

Chapter 3 1

(Th is table cont inued on next page. )

Ideal ly , all Enewetak spec ies recor ds not con tained inthe pr ev ious chapter s would be pr esen ted her e. U nfor

tunately, the vas tnes s of the bio log ical l i teratur e and the

t ime avai lable would not perm i t such an under tak ing .

Ther efore, the sear ch was con fined to the more obv iousr eference mater ials that m ight con tain these recor ds . We

hope th is smal l checkl i s t (Table 1 ) w i l l be of some help ,and we apo log i ze to those autho r s whose recor ds we havem issed .

Some of the pr ev ious chapter s l is ted on ly geo log i ca ll yr ecen t mater ial . For th is r eason some foss i l mater ialand/or mater ial fr om dr i l l cor es is contained herein , even

though the s ys tematic group may have been mor e com

p letely d iscus sed in pr ev ious chapter s of th is volume.

338 TITGEN AND BURCH

TABLE 1 (con t ’d)

D iv i s ion RH ODOPH YTAClass FLORIDEOPH YCEAE

Order RH ODYMEN IALESFam i ly RH ODYMEN IACEAEChyloc ladio sp . : Bai ley-B r ock et al . , 1 980 .

KINGDOM AN IMAL IAPhy lum SARCOMAST IGOPH ORASubphy lum SARCOD INAClass GRANULORETICULOSA

O rder FORAMINIFERIDASuborder ROTAL IINA

Super fam i ly GLOBIGERINOIDEAFam i ly GLOB IGER IN IDAE

‘G lobiger ina sp . aff . G . aper tur a Cushman , 1 9 1 8 : Todd , 1 964 .

‘G lobiger ina cong lomer oto Schwager , 1 866 : Todd , 1 964 .

‘G lobiger ina (Beella) dig itato B rady , 1 884 : Todd , 1 964 .

‘G lobiger ina (G lobor otalo ides) hexagono Natland , 1 938 : Todd , 1 964 .

‘G lobiger ina obes a (Bo l l i , Todd , 1 964 .

‘G lobiger ina rubescen s H ofker , 1 956 : Todd , 1 964 .

‘G lobiger inella adams i (Banner and B low, Todd , 1 964 .

‘G lobiger inello aequ i lateral is (B r ady , Todd , 1 964 .

‘G lobiger in ita g lut inata (Egger , Todd , 1 964 .

‘G lobiger in ito hum i l is (B r ady , Todd , 1964 .

‘G lobiger ino ides e longatus (d’

Or bigny, Todd , 1 964 .

‘G lobiger ino ides soccu l ifer f is tu los a (Schuber t , Todd , 1 964 .

‘G loboquodr ino olt isp i ro (Cushman and Jar v is , Todd , 1 964 .

‘G loboquodr ino alt isp i ra g lobosa 8 0 111, 1 957 : Todd , 1 964 .

‘Or bu l ina bi loboto (d’

Or bigny, Todd , 1964 .

‘Or bu l ina s u tu ral is B r onn imann , 1 95 1 : Todd , 1 964 .

‘Sphaero idinello disjuncta Fin lay , 1 940 : Todd , 1 964 .

‘Sphaer o idinello koch i (Caudr i , Todd , 1 964 .

Fam i ly GLOBOROTAL IIDAE‘G lobo r otol ia (Tu r bo rotal io) acos toens is B low, 1 959 : Todd , 1 964 .

‘G lobo r otal io fohs i r obus ta 80 111, 1950 : Todd , 1964 .

‘G lobor otal ia (Tr uncorotal io) punctu lata (d’

Or bigny, Todd , 1964 .

Fam i ly H AST IGERIN IDAE‘H as t iger ino pelag ico (d ’

or bigny , Todd , 1 964 .

Phy lum CN IDAR IAClass SCYPH OZOA

Order CORONATAEFam i ly L INUCH IDAETL in uche sp .

H YDROZOAOrder H YDRO IDASuborder ANTH OMEDUSAE

Fam i ly CLAVIDAE?Rh izogeton sp . : Cooke, 1975 .

Fam i ly EUDENDRIIDAEEudendr ium ?br eve Fraser , 1 938 : Cooke, 1 975 .

Eudendr ium cap i l lar e A lder , 1 856 : Cooke, 1 975 .

Fam i ly H ALOCORDYL IDAEH olocordyle dis t icha (Go ldfuss , Cooke , 1 975 .

Subo rder L EPTOMEDUSAEFam i ly CAMPANULARIIDAEC lyt ia hem isphoer ico Cooke, 1975 .

Fam i ly H ALECIIDAEH alecium beon i (Johnson , Cooke, 1 975 .

H alec ium sp . : Cooke , 1 975 .

‘Foss i l mater ial and/or mater ial fr om cor es .

(“Records fr om the Enewetak La bo rator y Card Catalogue . (Th is table continued on next page.

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340 TITGEN AND BURCH

TABLE 1 (con t’d)

Fam i ly FUNG IIDAE‘D is cot rochu s sp . cf . D . or bignyanus M i lne Edwards and H aime, 1 848 : Wel ls , 1 964 .

Fam i ly POR IT IDAE‘AlveOpo ra po lyacan tha Reuss , 1 867 : Wel ls , 1 964 .

‘D ictyaraea m icran tha Reus s , 1 867 : Wel ls , 1 964 .

‘Por ites sp . cf . P . capr icor n is Rehber g , 1 892 : Wel ls , 1 964 .

Order ZOANTH IDEAFam i ly ZOANTH IDAETGemmar ia sp .

Tlsau r u s duchasso ing i Andr es , 1 883 .

TPalythoa tuber cu losa (Esper ,TZoon thus sp .

Subclass CERIANT IPATH ARIAOrder CERIANTH ARIA

PEN ICILLARIA

¢Fam i ly ARACH NACTIDAE‘

l'A rachnan thus ous t ral iae Car lgr en , 1937 .

Phy lum PLATYH ELMINTH ESClass CESTODASubclass EUCESTODA

Order CYCLOPH YLL IDEAFam i ly H YMENOLEPID IDAEH ymeno lep is dim inuta (Rudo lphi , see Jackson et al . , Vol . I . , Chapter 1 2 .

Phylum NEMATODAC lass SECERNENTEASubclass SPIRURIA

Order SPIRUR IDASuborder SPIRUR INA

Super fam i ly SPIRUROIDEAFam i ly GONGYLONEMATIDAEGongy lonema neop las t icum F ibiger and D i t levsen , 1 9 1 4: see Jackson et al . , Vol . I , Chapter 1 2 .

Phy lum SIPUNCULAFam i ly PH ASCOLOSOMATIDAE§Phosco los oma olbo l ineotum Bai rd , 1 868 .

§Phas co losoma pacif icum Kefer s tein , 1 866 .

Fam i ly SIPUNCULIDAE§Siphonosoma r otuman um (Sh ip ley ,

Phy lum CH AETOGNATH AClass SAG ITTOIDEA

Order PH RAGMOPH ORAFam i ly SPADELL IDAESpodella legazp ichess i A lvar ino , 1981 .

Phylum ARTH ROPODASubphy lum CRUSTACEAClass MALACOSTRACASubclass EUMALACOSTRACASuperorder PERACARIDAOrder ISOPODASuborder ON ISCO IDEAInfraorder L IG IAMORPH A

Sect ion CRINOCH ETASuper fam i ly ATRACH EATAFam i ly ON ISCIDAE§A llon iscu s oahuens is Budde-Lund , 1 885 .

‘Fos s i l mater ial and/o r mater ial fr om cor es .

TReco rds from the Enewetak Labor ator y Card Catalogue .

rt See den H ar tog ( 1 977) on the systemat ic clas s ificat ion of the genus A r ochnan thus .

§New Enewetak reco rds . (Th is table cont inued on next page. )

MISCELLANEOUS SPECIES RECORDS 34 1

TABLE 1 (con t’d)

Super fam i ly PSEUDOTRACH EATAFam i ly PORCELL ION IDAEPor cell ion ides p r u inosus (Br andt ,Metapor nor thr us p r u inos us : Woodbur y , 1962 .

Suborder EPICAR IDEAInfraorder BOPYRINA

Fam i ly BOPYRIDAEBopyrello thom son i Bonn ier , 1 900 ; Bourdon , 1 980 .

Bopyrello thom son i m u iens is Danfor th , 1 970 .

Cancr icepon gar th i Danfor th , 1 970 .

Concr icepon? knudsen i (Danfor th ,

Merocepon knudsen i Danfor th , 1 970 .

G igon t ione p r ott i Danfor th , 1 967 : Danfor th , 1 970 .

G raps icepon sp . : Danfor th , 1 970 .

Merocepon knudsen i Danfor th , 1970 .

Scyr ocepon hawai iens is R ichardson , 191 1 : Danfor th , 1 970 .

Order AMPH IPODASuborder GAMMARIDEA

Super fam i ly GAMMAROIDEAFam i ly GAMMAR IDAEBeaudett io pa lmer i Bar nard , 1 965 .

Elasm op us pecten icr us (Bate, Bar nard , 1 965 .

ElasmOp us p seudaff in is Schel lenber g , 1 938 : Bar nard 1 965 .

Elasmop us r apox Costa, 1 853 : Bar nard , 1 965 .

Elasmopu s sp in idactylus Chevreux , 1 907 : Bar nard , 1 965 .

Elasmop us sp . : Croker , 1 971 b .

Jer bar n ia mecoch i r a Croker , 1971 a.

Maera ham igera H aswel l , 1 879 : Bar nard , 1 965 .

Maera inoequ ipes (Costa, Bar nard , 1 965 .

Moera othonop s is Schel lenberg , 1 938 : Bar nard , 1965 .

Maera pocif ica Schel lenberg , 1 938 : Bar nard , 1965 .

Maera quodr imana (Dana, Bar nard , 1 965 .

Maero ser rata Schel lenberg , 1 938 : Bar nard, 1965 .

Ma l lacoo ta ins ign is (Chev reux ,

Maer a ins ign is : Bar nard , 1965 .

Me l i to celer icu lo Croker , 1 971 b .

Super fam i ly LEUCOTH OIDEAFam i ly LEUCOTHOIDAELeuco thoe hyhelia Bar nard , 1 965 .

Super fam i ly TAL ITROIDEAFam i ly H YAL IDAEH yale chevreux i K . H . Bar nard , 1 916 : Bar nard , 1965 .

H yale den t ifere Chevr eux , 1 907 : Bar nard, 1 965 .

H yale hono lu luens is Schel lenberg , 1 938 : Bar nard , 1 965 .

H yale media (Dana, Bar nard , 1 965 .

Talor ches tia sp in ipolmo (Dana, Bar nard , 1960 ; Woodbury , 1 962 .

Super fam i ly COROPH IOIDEAFam i ly AMPITH OIDAEAmp i thoe romandi Audou in , 1826 : Bar nard , 1 965 .

Cymadu so br ev idactyla (Chevr eux , Croker , 197 1 b .

Cymaduso f i loso Sav igny , 1 8 1 6 : Bar nard, 1 965 .

Parag r ubia vorax Chevreux , 1 90 1 : Bar nard , 1965 .

Fam i ly AORIDAELembos oequ imon us Schellenber g , 1938 : Bar nard , 1 965 .

Lembos bryop s is Bar nard , 1 965 .

Lem bos sp . cf . L . f r oncan n i Reid , 1 95 1 : Bar nard , 1 965 .

Lem bas sp . cf . L . in termediu s Schel lenberg , 1 938 : Bar nard , 1 965 .

M icrodeu topus t r idens Schel lenberg , 1 938 : Bar nard , 1 965 .

Fam i ly ISAEIDAEGammar op s is at lan t icus (Stebbing ,

Eu r ys theus at lan t icu s : Bar nard , 1 965 .

(Th i s table con tinued on next page. )

342 TITGEN AND BURCH

TABLE 1 (con t'd)

Fam i ly ISAEIDAE (cont ’d)Gammarop s is dig itatus Schel lenberg , 1 938 .

Eu r ys theus dig itatu s : Bar nard , 1 965 .

Gammarop s is pacif icus Schel lenberg , 1 938 .

Eu r ys theus ?pocif icu s : Bar nard , 1 965 .

Megomphop us abbot t i Bar nard, 1 965 .

Subo rder CAPRELL IDEAFam i ly CAPRELL IDAE§Metap r o tella sandalens is Mayer , 1903 .

Phy lum TARD IGRADAOrder EUTARDIGRADA

Fam i ly MACROBIOTIDAEH yp s ibius (Isohyp s ibius) augus t i Mur r ay , 1 907 .

H yp s ibiu s (Isohyp s ibius) angus t i : Mehlen , 1 972 .

H yp s ibius (Colohyp s ibius) t r uncatu s Thu l in , 1 928 : Mehlen , 1 972 .

Macr obiotus al lan i Mur ray , 1 9 1 3 : Mehlen , 1 972 .

Macr obiotus harmswor thi Mur r ay , 1 907 : Mehlen , 1 972 .

Macr obiotus r ichter s i Mur ray , 1 9 1 1 : Mehlen , 1 972 .

Fam i ly MILNESIIDAEM i lnes ium tardig r adum Doyer e, 1 840 : Mehlen , 1972 .

Phy lum ECH INODERMATASubphy lum ECH INOZOAClass ECH INO IDEASubclass EUECH INOIDEASuperorder GNATH OSTOMATAOrder CLYPEASTEROIDASuborder LAGANINA

Fam i ly F lBULARlIDAE‘Echinocyomus par vipor u s Kier , 1 964 .

‘Echinocyamus cf . par v ipor u s K ier , 1964 .

‘Echinocyom us petolus K ier , 1964 .

Phy lum H EMICH ORDATAClass ENTEROPNEUSTA

Fam i ly PTYCH ODERIDAETPtychoder a flava Eschscholtz , 1 825 .

‘Foss i l mater ial and/or mater ial fr om cores .

“

( Reco rds fr om the Enewetak Labo r ato ry Ca rd Catalogue .

§New Enewetak r ecords .

ACKNO“ &EDGMEN TS

The autho r s would l ike to thank those who helped suppl y species r ecor ds used in th is chapter . J . L . Bar nar dsuppl ied r ecor ds fo r per acar id Cr us tacea, G . A . Samuelson helped w i th the Tar d igr ada, and J . H . Bai ley-B r ockforwar ded reco r ds of Sipuncula that had been iden t i fied byE . B . Cut ler . J . Mar kham helped us update the bopyr idi sopod r ecor ds . I . Abbo t helped w i th the algae, and

A . A lvar ino helped w i th the Chaetognatha .

REF ERENCES

A lvar ino , A . , 198 1 , Spodella legoz ip iches s i , A New Benth ic Chaetognath from Enewetak , Mar shal l Is lands , Pr oc . B io l . Soc .

Wash . , 94 : 1 07- 1 21 .

Bai ley-B rock , J . H . , J . K . Wh ite, and L. A . Ward, 1980 , Effectsof A lgal Tur f and Depr ess ions as Refuges on Po lychaete

Assemblages of a W indward Reef Bench at Enewetak Ato l l ,M icrones ica, 1 6 : 43 -58 .

Bar nard , J . L . , 1 960 , Insects of M icr ones ia, Crustacea: Amphipoda (Str and and Ter r estr ial Tal i t r idae) , Insects M icrones ia, 4:

1 3 -30 .

1 965 , Mar ine Amph ipoda of Ato l ls in M icr ones ia, Pr oc .

U . S . Nat l . M us . , 459-552 .

Bourdon , R . , 1 980 , Les especes du gen re Bopyr ello, J . Bonn ier(Crus tacea,

Isopoda, Bopyr idae), Bu l l . M us . Natn . H is t . Not .

(Par i s ) , (4) 2(A ) : 1 85-236 .

Ciereszko , L . S., M . A . Johnson , R . W . Schm idt , and C. B .

Koons , 1 968 , Chem i str y o f Coe lenterates — VI, Occur r ence of

Gorgos ter o l , a C30 Ster o l , in Coe lenterates and Thei r Zooxanthellae, Comp . B iochem . Phys io l . , 24: 899-904 .

Cooke , W . J. , 1 975 , Shal low Water H ydro ids fr om EnewetakAto l l , Mar shal l Is lands , M icr ones ica, 1 1 : 85

- 108 .

Cr oker , R . A . , 1971a, A Remar kable New Amph ipod Genus(Crustacea, Gammar idae) fr om En iwetok Ato l l Lagoon , Poc .

Sci 25 : 382-386.

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Author Index

Ar neson ,A . C . , 53

Bai ley-B r ock , J . H . , 97

Ber ger , A . J., 33 1

Br uce , A . J. ,221

Bur ch , B . L . , 337

Chave,E . H . , 37

Ch i ld , C . A . , 1 79

Cox , R . S. , 85

Cuffey , R . J. , 85

Cu t ress , B . M . , 263

Cu t ress , C. E . , 53

Devaney , D . M ., 37 , 49 , 63 ,

95 , 97 , 1 97 , 221 , 277

Dunn , P . H . , 1 1

Eldr edge, L . G . , 287

Gar th , J . S. , 235

Ger ber , R . P . , 203

G ran t , R . E . , 77

H aig , J. , 235

Johnson , S. , 105

Kay, E . A . , 1 05

Kor n icker , L . S. , 21 7

Knudsen , J . W . , 235

Lamber son , J . O . , 1 7 , 325

Lang , J . C ., 67

Mann ing ,R . B .

, 1 8 1

N ish ida, G . M . , 1 47

Randal l , H . A . , 289

Randal l , J . E . , 289

Reaka, M . L . , 1 81

Reese, E . S. , 333

Reyno lds , D . , 1 1

Rowe , F . W . E . , 263

Samuelson , G . A . , 1 47

T i tgen , R . H . , 191 , 337

Tsuda , R . T . , l

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348

Mammals , 333Man , 333

Mangr ove swamps , 1 7Manti s shr imps , 18 1 , 1 82, 1 87

- 1 89

Mar iana Is lands , 1 05 , 1 34 , 1 41 , 1 59,

1 65 , 1 67- 1 69, 289 , 3 14 , 325 , 328

Mar quesas Is lands , 33 1Meiobenthos , po lychaetes , 1 03Melanes ia, 33 1

M icrones ia, 21 7, 33 1

Mo lucca Is lands , 228Mo l lusks , 1 05 , 1 32, 133 , 1 34

bivalves , 1 33 , 1 34

cephalopods , 1 05 , 1 34

gas tr opods , 1 32 , 133 , 1 34 , 1 39

nonmar ine, 1 33

opi sthobranchs , 1 32

Mon itor l i zard, 328Moar ea, 1 5

Mor tal ity , of s tomatopods , 1 88Mouse, 333

Natur al products chem is tr y , of sponges ,63

Nematodes , 95 , 96

New Caledon ia, 1 5 , 1 94 , 288

New Guinea, 1 5 , 225 , 228

New H ebr ides , 1 42New Zealand, 1 5, 1 80 , 33 1

Nonmar ine Mo l lusks , 1 33

Ocean ia, 1 81

Ok inawa 1 33 , 158 , 1 94

Oper ation Cr oss roads , 1 , 37 , 97 , 105

235 , 289

Ostr acods , 21 7

Palmyr a, L ine Is lands , 288Palau, 1 58 , 1 59 , 160 , 1 62, 1 67, 1 94 ,

237, 280

Papua New Guinea, 325 , 328

Paras i tesCi r r ipedia, 1 93

copepods , 1 97, 1 99cr abs , 236 , 237 , 248

ho lothur ia, 264

SUBJECT iNDEX

insects , 1 59 , 1 67 , 1 68

Pycnogon ida, 1 79

worm s , 95

Ph i l ippines , 14 1 , 1 62 , 1 63 , 167 , 1 69 ,

1 94 , 225 , 237 , 264 , 283 , 287 , 288 ,

328

Phoen ix Is lands , 1 61 , 1 63Phytoplankton , 203

Pigs , 333P ison ia for est , 23 , 28 , 30 , 34

P lankton , see ZooplanktonP lants , 1 7 , 23 , 24 , 25 , 27, 28 , 30 , 34

br eadfru it , 24 , 25

coconut , 24 , 25 , 30 , 34

pandanus , 1 7 , 24 , 30 , 34

p ison ia, 1 7 , 23 , 27 , 28 , 29, 30 , 34

Po lychaetes , 97 , 1 02 , 1 03

Po lynes ia, 228 , 33 1

Porpo ise, 333

Predationfishes on for am in ifera , 37

nudibranchs on sponges , 49s tomatopods , 1 86 , 1 87

Product iv i tyalgae, 1

cor als , 73phytoplankton , 203

Pr otozoans , 37 , 46

Samoa Is lands , 1 59 , 1 60 , 1 61 , 1 64 , 1 67,

1 68 , 1 8 1 ,

Scorp ions , 1 58Sea anemones , 53 , 54

Sea bi rds , 33 1Sea cucumber s , 263 , 264

Sea grass , 8 , 1 4 , 1 34 , 1 41

of scleract in ia

Ujelang Ato l l , 8 , 25 , 235

Vanuatu , 1 42

Whales , 333

Zanzibar , 288Zooplankton , 203 , 21 1 -21 5

Zooxanthel lae, cor als , 63

Seychel les Is lands , 264Shearwater s , 30Shoreb i rds , 33 1Shr imps , 221 , 225 , 226

Singapo re, 1 80

Sk inks , 328Snakes , 325 , 328

Soc iety Is lands , 1 5 , 264 , 3 1 4

So lomon Is lands , 37, 1 02, 325

South China Sea, 1 8 1

Spider s , 1 58Sponges , 49 , 51

Sr i Lanka, 1 64 , 1 65

Starf ish , 277Sumatra, 1 02

Symbios isbar nacles , 1 94 , 1 95

copepods , 1 97

corals , 63 , 73

cr abs , 236 , 237

foram in ifera, 46

po lychaetes , 97pycnogon ida, 1 79

shr imps , 225-228