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Page 1: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon
Page 2: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon

NOAA Techn ica l Report NMFS C ircu lar438

or“ # Co

# , Marine F ora and Fauna of

5$33; The Northeastern Un ted States .

g Sc eract in iasu res OT

93“

S tephen D. Ca i rns

J u ly 1981

Marine Biological Laboratory7

l SQAFIV

US . DEPARTMENT OF COMMERCE

Nat i ona l Ocean i c and Atmospher i c Adm i n is t rat i onN at i ona l Mar i ne F isher i es Se rvi ce

Page 3: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon

N OAA TECHN ICAL REPORTS

N at ional M ar i ne F i sher ies Servi ce, Ci rcu l ars

The major reSpon si b i l i t ies o f t he N a t iona l Mari n e F is heri es Serv ice (N M FS) are t o mon i t o r and assess t he abun dan ce and geograph ic d is t r ibu t ion o ffishe ry resou rces , t o understand and pred ic t fluc tuat i on s i n t he quan t i t y and d is t r ibu t ion of t hese resources , and to estab l i sh leve ls for opt imum use of t heresou rces . N M FS is a lso charged wi t h t he developmen t and implemen ta t ion o f po li c ies for managin g na t iona l fis h ing groun ds , deve lopmen t and en forcemen t o f domest ic fisher ies regu lat ion s , su rvei l lan ce o f foreign fish ing o ff Un i ted Stat es coasta l wa t e rs , and t he developmen t and en forcemen t o f in t ernat ional fis hery agreemen ts and po l i c ie s . N M FS a ls o ass is ts t he fish in g in dus t ry t h rough mark e t in g se rv ice and econom ic ana ly s is p rograms , and mort gagein su rance and vesse l con s t ruct ion subs id i es . l rco l lect s , ana lyzes , and pub l i shes s ta t i s t ics on var iou s phases o f t he indus t ry.

The NOAA Techn ica l Repo rt NM FSC i rcu la r se rie s con t in ues a seri es t hat has been i n ex is tence s in ce 194 1 . The C i rc u lars are techn ica l p ub l i cat ion so f gen era l in te res t in tended to a i d con servat ion and managemen t . Pub l i cat ion s t ha t rev iew i n con s iderab le deta i l and at a h igh techn i ca l leve l certa inb road areas o f researc h appear i n t h is se ries . Techn ica l papers ori g in at ing i n econom ic s s tu d ies and from managemen t in vest iga t ion s appear i n t he C i rc u la rser ies .

NOAA Techn ica l Repo rt NM FS C i rc u la rs are ava il ab le free i n l im i t ed n umbers to governmen ta l agenc ies , both Fed era l and Sta te . They are a lsoava i lab le i n exchange for o t her sc i en t i fic and t echn i ca l pub l i cat i on s i n t he mar in e sc ien ces . I nd iv idua l copies may be ob ta in ed (un less o t he rw i se not ed )from 0822. U se r Serv ic es B ranch , En v i ronmen ta l Sc ien ce I n fo rma t ion Cen te r, NOAA , Rockv i l le , M D 20852 . Recen t C i rc u lars are:

4 18. A nnota ted b ib l i ography of four A t lan t ic scombri ds : Scomberomoms

bras i l i en s is , S. cava l la, S. macu l arus, and S. regal is . By Char les S. M anooch 11 1 ,

E ugen e L . N akamura , and A n n Bowman Ha ll . D ecember 1978, i i i + 166 p .

4 19. Marin e f lora and fauna of t he n ort h eas te rn U n i t ed Sta tes . P rotozoa : Sarcod ina : Amoebae . By E ugen e C . Bovee and Thomas K . Sawyer. Jan uary 1979,

i i i 56 p . , 77 f igs . For sa le by t he Superin tenden t o f Documen ts , U .S. Governmen t P rin t ing O ffice , Wash in gton , D .C . 20402 . St ock N o . 003-017 -00433-3.

420. Pre l i m in a ry k eys to o to l i t h s o f some adu lt fis hes o f t he Gu lf o f A la ska,

Ber in g Sea , and Beau fo rt Sea . By James E . Morrow . Feb rua ry 1979. i i i + 32 p9 p l .

42 1 . La rva l deve lo pmen t o f sha l low wate r bam ac les o f t he Caro li nas (C irrip ed ia : Thorac ica) wit h keys to naup l iar stages . By W i l l iam H . Lang . February1979, i v + 39 p . , 36 f igs , 1 7 tab les .422 . A rev is ion of t he catsharks, fam i ly Scy l ioh in idae. By St ewart Springer.A pr i l 1979,

v + 142 p . , 97 figs . For sa l e by t he Supe rin t en den t o f Documen t s ,U .S . Governmen t Prin t in g O ffice , Wash in gton , D .C . 20402 , Stock No .

003-020-00147 -5 .

423 . Ma rin e flora and fauna of t he n ort heas te rn U n i ted Sta te s . C ru stacea:Cumacea . By Les Wa i l i ng . A pr i l 1979, i i i + 23 p . , 35 figs . For sa le by t heSu pe r in t e n d en t o f D oc umen t s , Wa s h in g to n , D .C . 20402 . S t o c k N o .

003-017-00446-5 .

424 . Guide to t he leptocepha l i (E 10p i formes. A ngu i l l i fo rm es , and N o tacan th ifo rmes) . By Dav id G . Sm i t h . J u ly 1979. i v + 39 p . , 54 figs .

425 . Marin e f lo ra and fauna of t he n ort h eas t e rn U n i t ed Sta te s . A rt h ropoda :C i r r i ped ia . By V ic tor A . Zu l lo . A pri l 1979, i i i + 29 p . , 40 figs . For sa le by t heSu pe r in t en d e n t o f D ocumen t s , Wa s h in g t o n , D .C . 20402 , St o c k N o .

003-017 -00453-8 .

426. Synopsis o f b io logica l data on t he rock crab , Can cer irrorarus Say. By

Thomas E . B igford . M ay 1979, v + 26 p . , 1 1 figs . , 2 1 tab l es .427 . Ocean variab il i t y i n t h e U S . F is he ry Con se rvat ion Zon e , 1976 . By J u l i enR . Gou let , J r. , and E l i zabe t h D . Hayn es , ed i to rs . J u ly 1979, i i i + 362 p .

In t roduct ion . By J u li en R . Gou le t , J r. J u ly 1979, p . 1-2 .

Summa ry. By J u l i en R . Gou le t , J r. J u ly 1979, p . 3— 10.

A tmospheri c c i rcu la t ion i n 1976. By E li zabet h D . Hayn es . J u ly 1979. p .

1 1 — 18 , 2 figs .

A tmospheri c c l imato logy and i t s e ffec t on sea su r face tempera tu re1976. By Robe rt R . D i ck son and J erome Namias . J u ly 1979, p . 19— 33, 6 figs .

Eas te rn Pac i f ic sea su r face cond i t ion s i n 1976. By E l i zabet h D . Hayn es .

J u ly 1979, p . 35—4 2 .

Sea su rface cond it ion s i n t he weste rn N ort h A tl an t i c i n 1976. By J u li enR . Gou let , Jr. , and E l izabet h D . Hayn es . J u ly 1976, p . 43— 50.

A noma li es o fmon th ly mean sea level a lon g t he west coa ts o f N ort h andSou t h Ameri ca . By Da le E . Bretschnei derand Douglas R . M cLai n . J u ly 1979, p .

5 1 — 64 . 6 f igs .

Coas ta l u pwe ll in g o ff weste rn N ort h Ameri ca , 1976. By C ra ig S. N e ls on .

J u ly 1979,p . 65— 75 , 2 figs , 2 tab les .

Ocean ic cond i t ion s d urin g 1976 between San F ran c is co and Hono lu lu asobse rved from Sh i ps o f O pport un i t y. By J . F. T. Saur and D . R . M cLa i n . J u ly1979. p . 77 —92. 5 figs , 2 tab le s .

The 1976 E l N in o and recen t p rogress i n mon i t o rin g and p red ic t ion . By

W i l l iam H . Qu in n . J u ly 1979, p . 93— 1 10. 7 figs. , 4 tab les .Sea su r face tempera t u re anoma l i es . By Douglas R . M cLain . J u ly 1979,

p . 1 1 1 — 149, app . 1 .

F l uct uat ion s o f sea su r face tempera tu re and densi t y at coas ta l s ta t ionsdu r ing 1976. By Douglas R . M cLai n . J u ly 1979, p . 15 1- 166 , 7 figs. , app . 1 .

Da ta on cold weat her cond it ion s a long the A tlan t i c and Gulf coas t s duri ng t he fa l l and win t er o f 1976— 77. By J . Lockwood C hamber l in and Reed 5 .

A rmst rong . J u ly 1979. p . 167— 174 , 1 fig . , 1 tab le .

W in d dri ven t ran sport A t lan t i c coas t and Gulf o f M ex ico . By M erton C .

l ngraham. J u ly 1979. p . 175-208. 4 figs , 1 tab le , app . 1 .

Sea su r face t emperat u re d is t r ib ut ion from Cape Cod , Massachuset t s , toM iam i , F lori da 1976. By Joseph W. Deaver 1 11 . J u ly 1979, p . 209—229, 3 figs , 2

tab les , app . 1 .

Water co lumn t herma l s tr uc tu re across t he she l f and s lope sou t heast o fSan dy Hoo k , N ew J ersey, i n 1976. By Steven K . Cook . J u ly 1979, p . 231-257,

19 figs .

A n t i cyc lon ic Gu l f Stream edd ies o ff t he N ort heastern U n i t ed Sta tes d uring 1976. By Dav id M i zen ko and J . Lockwood C hamberl in . J u ly 1979, p .

259—280, 16 figs. , 2 tab les .R iv e r run o ff a lon g the M id d le A t lan ti c coas t in 1976. By E li zabet h D .

Haynes . J u ly 1979, p . 281 - 287 , 1 fig . , 1 tab le , app . 1 .

Climat ic cond it ion s re la ted to t he fish k il l and anox ia off New J erseydu ring t he summer o f 1976. By Reed S. A rms t ron g . J uly 1979, p . 289-300,

5 figs .

Varia t ions i n t he posi t ion o f t he shel f wate r fron t o ff t he A t lan t ic coastbe tween Georges Ban k and Cape Roma in in 1976. By John T. Gun n . J u ly 1979,

p . 301 1 tab le .

Tempe ra tu re s t ructu re on t he con ti n en ta l she lf and s lo pe sou th o f N ewE n gland du ri n g 1976 . By R . Wy l ie C r is t and J . Lockwood Chamber l in . J u ly1979, p . 3 15— 335 , 2 figs. , app . 1 .

Con t in uous p lan k ton record s: Zoop la n k ton and net phyt oplan k ton i nt he M id-A t lan t ic B igh t , 1976 . By Dan i e l E . Sm i t h and Jack W. Jossi . J u ly 1979,

p . 337 — 348 ,7 figs .

Si p honophore l i po swarm in g i n N ew E nglan d coasta l wate rsupdate , 1976. By Caro lyn A . Rogers . J u ly 1979, p . 349—352 , 1 fig .

Bot tom-wate r tempera t u res i n t h e Gu lf o f Main e and on Georges Ban kduring spr ing and au t umn , 1976. By C laren ce W. Dav is . J u ly 1979, p . 353— 362 , 6

figs. , 3 tab les .

Page 4: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon

NOAA Techn ica l Report NMFS C ircu lar438

Marine Flora and Fauna of

the Northeastern Un ited States .

Sc leract in ia

Stephen D. Ca i rns

J u ly 1981

US . DEPARTMENT OF COMMERCEMa lcolm Ba ld rige , Sec re ta ry

N at i ona l Ocean i c and Atmospher i c Adm i n ist rat i on

N at i ona l Mar i ne F isher i es Serv i ceTe rry L. Le itze l l , Assistan t Admi nist ra tor for Fishe ries

Page 5: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon

FOREWORD

Th is N M FSC i rcu lar i s part o f t he s ubser ie s Mar in e F lo ra an d Fauna o f t he N ort heas te rn Un i t ed Stat e s ,wh ich con s i s t s o f o r igina l , i l l us t ra t ed , modern man ua l s on t he iden t ifica t i on , c lass ificat ion , and gen era l b i o logyo f th e est uar in e and coas ta l ma r in e plan t s and an ima l s o f t h e N ort heas t e rn Un i t ed Sta te s . The man ual s arepub l i shed at i r regu lar in t erva l s on as many taxa of t he reg ion as t h ere are spec ia l i s t s ava i lab le t o co l labora te i nt h e i r p reparat ion .

Geograph ic coverage o f t he Mar in e F lo ra and Fauna of t he N o rt heast e rn Un i t ed Stat es i s p lan n ed toin c lude o rgan isms from t he headwat e rs o f est uar ie s seaward to approximat e ly t h e 200m dep th on t h e con t in en ta ls hel f from Ma ine to V i rg in ia, bu t may vary somewha t w i t h each major taxon and t he in t eres t s o f co l laborat o rs .W henever poss ib le represen tat iv e spec imen s dea l t w i t h i n t he man ua l s are depos i t ed i n t h e re fe rence co ll ect ion s o fmajor museums o f t he reg ion .

The Ma r in e F lora and Fauna o f t he N o rt heaste rn Un it ed Stat e s i s bein g p repared i n co l laborat ion wi t hsyst ema t ic spec ia l i s t s i n t he Un i t ed Sta t es and ab road . Eac h man ua l is based pr imar i l y on recen t and ongo ingrev is ionary systemat i c research and a fresh exam inat ion o f t he p lan t s and an ima l s . Each major taxon , t reat ed i n asepara te man ual , in c l udes an in t roduct ion , i l l ust rat ed g lossary , un i fo rm or igina l l y i l l ust rat ed k eys , annotat edc heck li s t w it h in fo rmat ion when ava i lab le on d is t r ibut ion , hab i ta t , l i fe h is t ory , and re la t ed b io logy . re ferences tot he major l i t erat u re o f t he group , and a sys t emat ic index .

These man ua ls are in tended fo r use by b io logy s t uden ts , b io logis t s . b io logical oceanographers , in formedlaymen , and ot hers w ish ing to iden t i fy coas ta l o rgan i sms for t h i s reg ion . O ften t hey can serve as gu ides toadd it iona l in fo rmat ion about spec ies o r groups .The man ua l s are an outgrow th o f t he wide ly used Keys to Mar ine 1nvertebrates o f t he Woods Hole

R eg ion? ed it ed by R . 1 . Sm i t h i n 1964 , and p roduced under t he ausp ices o f t he Syst ema t i c s E co logy Program ,

Ma r in e B i o log ica l Labora t o ry , Woods Ho le , Mass . A ft er a su ffi ci en t n umber o f man ua ls o f re la t ed taxonomi cg roups have been pub l i sh ed , t he man ua l s w i l l be rev ised , grouped , and i ss ued as spec ia l vo lumes , wh ic h wi l lcon si s t of comp i lat ion s fo r phy la or groups o f phy la .

Page 6: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon

I n troduct i o n

M orpho logy

Keys to the Scleract i n ia o f th e northeastern coast o f the U n i ted States

D i chotomou s key

Tabu lar k eys

A nnotated systemat i c l i st

Se lected b ib l i ography

Systemat i c i ndex

A ck now ledgment s

Coord inat i ng Ed i to r ’ s comments

Page 7: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon
Page 8: €¦ · N OA A TECHN ICAL RE PORTS N ation al M arin e Fisheries Servi ce, Circu lars The major reSpon sibil ities of the Nation al Mari n e Fsi heri es Service (N M FS) are to mon

M ari ne F l ora and Fau na o f the N ortheastern

U n i ted States. Scleracti ni a

STEPHEN D . CA I RN S'

A BSTRACT

Th i s man ua l d iscusses the 14 spec ies o f sc lerae t i n ian co ra ls k nown from t he n o rt heas te rn l i n i ted S lat e s coas tfroth \ i rg i n ia to N ova Sco t ia . Fo l low in g a b r ie f in t rod uc t ion to t i l e gen era l b io logy and morpho logy o f

Sc leracti n ia . an i l l us t ra ted d ic ho tomous km an d two tabu lar keys are gi \ en fo r t h ese spec ies . A n an nota tedsss te l n a t i c l is t in c l udes comp let e geograph ic an d ha th u n c tri c ran ges , refe ren ces to pe rt in en t l i t e ra t u re , and , fo r

some spec ies . eco log ica l and taxonom i c n o tes . Zoo gco graph ic aff in i t ies o f t h e fau na are b r ie fly d iscussed . A

se lect ed b ib l iography is p rov ided .

I N TRODU CT I ON

Fou rt een species o f stony coral s (order Scleract in ia) are

k nown from o ff the northeaste rn coast from Chesapeake Bay t o

s o u t h e r n N ov a Sc o t i a o f w h i ch on l y o n e , A s trang i a

asrre if orm is , occu rs a t depth s shal l ow enough to be co l l ected

rou t i n ely by snorke l i ng or scuba d i v ing . The remai n ing spec ies

are u sua l l y co ll ected by ben th ic t rawls or d redges and occu r as

d eep as m o ff t he no rt heast coas t . M uch research h as been

done on the easi l y accessi b le A . asrreif orm is , bu t li t t le more than

physic al descr ip t ions and d i st r ibu t ions are known for t he deeper

wate r species . A l l 14 species are i nc luded i n th i s report .

Of th ese 14 species , 13 are ahermatyp ic; i .e t hey do no t

possess symb i ot i c zooxanthe l lae (a un ice l l u la r di no f l age ll ate) i n

t he i r endodermal t i s sue . I nd iv id u al co lon ies o f A . asrreif orm is

may or may not have zooxanthe ll ae , depend ing perhaps on

w ate r t emperatu re o r l igh t i n ten si t y ; more frequen t ly t h i s species

l a c k s t h e symb i on t s . The t e rm ahermatyp i c i s t h u s a

p hys io logical cond i t ion det erm i n ed by ecological factors an d

there fore i s not a characte r o f great val ue in c lassificat ion .

A hermatyp ic co ral s have been equated wi t h deepwater corals,so l i t a ry co ra l s , o r n onreef bu i ld i ng co ra l s . Th i s i s an

oversimp l i ficat ion ; i n fact , many species occur i n shal l ow water ,and some large , co lon ia l deepwate r (500-800 m) ahermatypes

Lap hel ia p ro l ifera ,E na l lop samm ia p rof una

a) fo rm the

framework for reefl i ke s t ru ctu res . I t i s t rue , however , that a l l

deepwater coral s are ah ermatyp ic because be low t he euphot ic

zone the zooxan thel lae , bei ng p lan t s , cannot photosyn thesize .

N o t re s t r i c t ed b y t h e g en e ra l l y h i gh e r (o ft e n t rop i ca l )temperatu re and l igh t requ i rements that zooxanthe l lae impose

on h ermatyp ic co ral s , the ahermatypes i n habi t a more extensive

geograph ic range . They are found in a l l oceans from the

N o rwegian Sea ( la t .70°30'

N ) to th e Ross Sea , A nt arct ica

( lat . 78°29'

S) , from m depth (Kel le r and in

temperatu res o f 1 . 1 °C to o v er 29°C .

Scleract i n ia are monoec ious or d ioec iou s ; i n e i the r case thei r

mot i le larval fo rm i s a pl an u la capab le o f remai n i ng p l an k ton ic

for w ee k s . I n add i t ion to sexual rep roduct i on,some species

p ropagate by asexual budd ing and others have remarkable

po w e r s o f regen era t i on . For i n s ta nce,t h e co ra l l um o f

D asmosm i l ia Iyman i usual l y sp li t s longi t ud inal l y in to several

Depan n ten t o l'

In sertebra i e Zoo logy . N a t i onal M useum o f Nat u ra l H iston .

Sm i t hso n ian I n st i t ut ion . ash ington , D .C . 20560.

fragmen t s , each wedge-shaped p iece subsequen t ly p roduc ing

1 -30 smal l buds growing di rect ly from the mesen ter ia l t i s sue . I t

i s rare to find a spec imen o fD . lymam'

that grew d irect l y from a

p lanu la , i .e . , not at tached to an in s ide fragment o f a paren t

spec imen .

A l l p lanu lae need a hard subst rate on wh ich to set t le .

Fo l l ow i ng set t lemen t and subsequen t g rowt h , coral s may e i t her

rem ai n a t t ach ed t o th e subst rat e o r become free , ly i ng

unat tached on the subst rate . The at tached cora ll um usual l y

rein forces i t s base o f at tachmen t by var ious mean s , whereas t he

su bsequen t ly unattached cora l l um becomes free by lack i ng

r e i n fo rcemen t o f an or ig i n a l l y weak at t achmen t o r b y

comp lete ly overgrowi ng the subst rate , as i n the case o f a sand

p art i c le . Some unatt ached species Caryop hy /I ia ambrosia)become top-heavy , fal l to one s ide , bu t su bsequen t l y reorient

t he i r cal i ces toward an upr igh t posi t ion , produc ing a horn

shaped coral l um .

The scleract i n ian fau na o f the northeaste rn coas t o f the

U n i ted States i s re lat i v e l y low i n d i vers i ty when compared to

o ther areas i n the western A t lant i c , even when the ree f co rals

(hermatypes) are exc luded from considerat ion . For example ,t h ere are app rox imate ly 160 specie s o f Scleract i n ia in t he

Car ibbean : 76 deepwate r (o ve r 200 m) ahermatypes (Cai rns

about 30 exc lu si v ely sha l l ow-wate r ahermatypes, and

abou t 54 hermatypic speci es . I n the region between north

F lo r ida and Cape H atteras there a re about 40species k no w n : 28

deepwater ahermatypes (Cai rns abou t 8-10 exc lu sive ly

s ha l l ow-wate r ahermatypes, and severa l hardy he rmat ypic

spec ies,i .e . , So l en oasrrea , Siderastrea , Ocu /i n a . I n the weste rn

A t lant i c no rth o f th e N orth Caro l i na-V irgin ia border there are

1 7 spec i e s k now n , 16 deepwa te r aherma types and one

exc lu si v e ly sha llo w-water species , A . asrreif orm tls. Fou rteen o f

t hese 1 7 species are t reated i n th i s wor k , the ot her th ree :

Vaughan el la margari ta ta ( Jou rdan ,F u ng iacyarh us d u rus

Kel l er, 1976; and F . maren ze/leri (Vaughan ,1906) are k now n

from l oca li t ie s north o f M aine . O ther species occu rring south o f

the area o f considerat i on fo r th is manua l that may subsequen t ly

be found there i nc lude : D en drop hy /I ia gad i tan a (Duncan ,Po u rra /osm i / i a conf erra C a i r n s , 1978 ; Con cen trorh eca

Iaevigata (Pou rta les, and Po lymy ces fragi l is (Pou rta les,The 14 spec i e s o f Sc l eract i n i a k n own from o ff t h e

northeaste rn coast o f the U n i ted States are , i n general , wide ly

d is t r ib uted species . Six are cosmopo li t an an d seven are amph i

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A t lan t ic . Of t he lat ter , five are p rima ri l y cold temperate , one

(A . astreif ormts) i s found bot h i n t emperate and t rop ical waters ,and one (D el tocyath us i ta l icus) i s primari l y a t rop ical species

w it h north ern lim i t s in t he co ld temperate region . O nl y one

spec ies, E na l lop samm ia p rofu nda , i s endemi c to the western

A t lan t ic an d i s most common in the warm t emperate region .

M ORPHOLOGY

The cal careous s ke leton o f a scleract in ian co ral , t he co ral l um ,

may be composed o f numerous ind iv idual u n i ts p roduc ing a

colon ia l coral l um o r i t may be a so li tary coral wi th on ly one un i t

p roduced by one po lyp . The shape o f the cora ll um o f so l i tary

corals i s importan t at the generic and specific levels an d i s

commonl y desc ribed in geometri c term s coni ca l , t rochoid ,

cyli ndri cal ) ; shapes o f co lon ial coral s are descr ibed by b ranch in g

pat te rn .

Coral s may be ei t her at tached to the subst rate o r unat tached

( free) , t h i s d i fference usua ll y be ing consi sten t at the species

leve l . I f at tached , t he base o f th e coral i s firm ly cemented to a

hard su r face . A sol i tary at tached coral usual l y has a steml i ke

pedi cel d i r ect ly above t he base (F ig . wh ich suppo rt s th e

ca l i ce , the round to e ll i p t i cal oral su r face o f the cora l l um . The

so l i tary at tached coral o ften re in forces i t s at tachment by

PALUS

l UM ELLA

CO STA

PEDICEL

F ig u re l .— C u taway d rawin g o f hypo t h e t i ca l s o l i tary co ra l i l l u s t ra t in g

mo rpho logi cal fea t ures .

th i ckeni ng i t s ped ice l , expand ing i t s base , an d , in some spec ies ,p roduc i ng an chor in g root le t s o r add i ng successive r ings o f

compa rtmental i zed , concen t ri c cham bers around the base and

ped ice l . A so l i t ar y unat t ached coral usual l y has a flat or bowl

shaped base and lack s the ped ice l . The sides or wa ll s o f sol i t ary

coral s and coral l i t es o f co lon ies are t ermed t he theca . Th is t heca

may be granu lar or porce laneous in textu re and o ften bears

longi t udi nal r idges cal l ed costae correspond ing to the larger

septa . Ske letal deposi t s formed between ind iv idua l coral l i tes o f

a co lony are cal l ed coenosteum .

The ca l i ce o f most spec ies i s regu lar l y and hexarn eral l y

s ubdiv ided by rad ial par t i t io ns , cal l ed sep ta . The six largest

septa,d iv ide the cal i ce i n to six equal areas or systems (F ig .

and comprise t h e fir s t cycle o f septa . The second cyc le al so con

si st s o f six septa wh ich are general l y sma ll er and occu r hal fway

between t he fir st cyc le septa . The 1 2 t h ir d cyc le septa are formed

in each space c reated by the prev ious 12 septa . Ca li ces w i th 6-7

cyc le s o f septa ( 192-384 septa) are k nown . Septa wi t h the upper

edges extend ing above the theca are term ed exsert septa .

Somet im es one or several adjacent septa o f a cal i ce are ext reme

l y exse rt and ben t over the ca l i ce , fo rm ing a hood li ke st ructu re

cal l ed a rostrum see figu re o f E n al lop samm ia rostrata) .

Sma ll accesso ry lobes are somet imes present on the i nne r edges

o f th e septa o f ce rt ain cyc les . These are cal l ed pal i (s i ngul ar:

palus) , or pa li form lobes , and are often u sed as a generi c level

character . D irect ly in the cent er o f the cal i ce t here is o ften a

st ructu re ca l l ed a colume l l a , which may appear as a Iame l l a

( lame l l ar) , a spongy mass (trabecu la r) , a s in gle rod (sty l i fon n ) , a

fie ld o f simp le or tw i sted rods (pap i l lose , fasc i cu l ar) , or s imply a

fusion o f th e in ner edges o f the larger septa (ru dim en tary) . The

type of colume l l a i s o ften used to di st i ngu i sh genera an d

subgenera .

SYSTEM

F lgure 2.— Di agram of rel ati ve slzec an d posi tion of sep ta In a cal i ce

con tai n ing th ree cyc les (24 sep ta ) . N umbers refer to th e cyc le to wh ichthe sep tum bel ongs ; onl y the upper righ t system ls comple tel yn umbered .

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KEYS TO THE SCLERACTI N IA OF THE NORTHEASTER N COAST OF THE U N I TED STATES

The ident ificat i on an d c lassificat ion o f Scleract in ia depend sto red d ry .

en t i re ly on characteri st i cs o f the cora l l um ; there fore , i t i s A n i l l u s t rated , d ichotomous key i s presented fi rs t , fo l lowed

general l y necessary t o remove t he t i s sue from the co ral l um by two t abu lar k eys to t he same species : one keyi ng t he co lon ia l

be fo re u si ng the keys . Thi s i s eas i l y accomp l i shed by soak ing the spec ies and the othe r , the so l i tary ones . A blan k space i n the

coral in fu ll -st rengt h commercial b leach fo r several hou rs t abu lar key in dicates that t h i s character does no t app ly to th i s

fo l lowed by thorough ri ns ing . The specimens shou ld then be species .

DICHOTOM OU S KEY

Coral l um so l i t ary (see t abu lar key 1 )

Coral l um colon ial (see tabu l ar key 2)

2 ( I ) Corall um fi rm ly at tached t o subst rate

2 ( I ) Coral l um unat tached o r at tached to smal l fragmen t o f p aren t spec imen 4

3 (2 ) F iv e o r more cyc les o f septa (2 96 septa) ; septa an d t heca th ic k , robust ; th eca gr anu lar and costate ; i n ner

sept al edges strai gh t (F ig . 3) Desmop hy l l um cristaga l l i

F i g u re 3 .— D esm op hy l l um cri sraga l l i : s i d e v i e w .

Scale : 3 cm . F igure 4 .—

'

Javan lu cal l l erI : A . si de vi ew of spec lmen wi t hdamaged uppe r sep ta#edges ; B , s ide v lew of spec imen w i th In tactsep ta. Il l u stra t in g b road . en crustl n g at tac hmen t . Scal e s : 1 cm.

3 (2) U sual l y fou r cyc les o f septa (48 septa) , occas ion al l y 64 septa , septa and upper theca very de l i cate ; theca

smooth and porce laneous ; i n ner septal edges s in uou s (F ig . 4A , B) J avam'

a ca i /Ieri

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5 (4) Corall um at tached to fragment o f paren t coral l um or has broken base ; co ral l um fragil e , eas il y fragmented ;pa li form lobes o ften mu l t i lobate an d i nd is t i ngu i shab le from the co lume l l a (F ig . 5A , B) D asmosmi l ia lymam

'

F igure S.— Da.rmosm i l ia lymam

'

: A , fragmen t of larger paren t coral l um wi t h t hre e sma ll coral l n asexual l y budd in g from paren t ; B . s id e v ie w of in ta c tcora l l um . Sca le : 2 cm.

5 (4) Coral l um free,base in tact ; coral l um s t rong , not eas i l y fragmented ; pal i s ing le-lobed and di s t in ct from co l

umel l a

6 (5) Pal i ( 12— 18) o f un i form s ize occur on in ner edges o f th ird cyc le septa ; co ral l um a curved cone (horn

shaped) ; ca l i cu lar d iameter up to 40mm (F ig . 6) Caryop hy l l ia ambrosia ambrosia

F igu re 6 .— Caryop hy l l ia am b ros ia amb ros ia : s ide v ie w

showin g crown of pal i an d severa l cen tral co lume l larri bbon s. Scal e : 2 cm .

6 (5 ) Pal i o f d i fferent s izes occur on first , second (somet im es m i ss in g) , and th i rd cyc le septa ; coral l um con ical but

not cu rved ; ca l i cu lar d i amete r rare ly over 15 mm (F ig . 7A , B)

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7 (4) Coral l um d i scoidal ; bas'

e flat (F ig . 8)F u ng iacyathusfrag i l is

F ig ure 8 .— Fu ngr

'

a cy arhus f rag i l i s : Ca l i c ula rvi ew of sli gh tl y damaged spec imen . Scal e : 1 cm.

7 (4) Coral lum con ical , compressed-coni

cal , or fragmen ted ; base not f l at 8

F ig ure 7 .— Delrocyathus i ta l i cus: A , cal i cu lar vi ew showin g

sep ta# and pal e r arrangemen t and co l ume l l a; large r pal i beforet h ird cyc l e septa ,

smal l e r before firs t cyc le septa ; pal ibefore secon d cycle ; B , s ide vi ew . Scal es : 5 mm.

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8 (7) Rudimentary co lumell a formed by fusion o f lower in ner edges o f larger septa ; coral l um usual l y i n tac t ; 9

8 ( 7) N o co l umel la ; coral l um i nvari ab ly fragment ed

(F ig . 9) . F label l um macandrewi

F ig ure 9.— F la bel l um macan drewi

si d e vi ew . Scal e: 1 cm.

9 (8) Ca l i ce compressed or constri c t ed in cen ter (Fig 10A , B)

F ig ure 1 0.— Flabeflum ala basrrum : A ,

cal i c u la r v iew showin g constric ted cal icef B , sid e v ie w . Scale: 2 cm.

9 (8) Cali ce a regul ar e ll i pse (F ig . 1 1 )F label l um angu l are

F ig ure l l .— Flabel l um an gu lare

s ide vi ew . Scale: 2 cm.

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10 ( I ) Corall um encrust ing , rare ly p roducing sho rt , st ubby branches ; i n ner edges o f septa den tate ; coral l i t es d i rect l y

adjacen t (Fig . 12) A strang ia astreif orm is

F igure 1 2 .— A strangia arrreif orm i r: an average-s i zed

co lony en crus t in g a roc k . Scale : 2 cm .

10 (1 ) Coral l um b ranch i ng , arborescen t ; i n ner edges o f septa smooth ; co ral l i tes not d i r ec t ly adjacen t

l l ( 10) Colon ies un ip lanar , wi t h cal i ces arranged on on ly one s ide ; each cal i ce bordered proximal l y by a rost rum

(hood ) (F ig . 13A , B) E na l /op samm ia rostrata

F lgure l 3.— Enaflop sammla rostrum: A , a bran ch from part o f a larger co lo

ny ; note trian gu lar swe l l lngs (ros tra) benea t h each ca li ce . Sca le: 2 cm . B , en

largemen t o f ro strum of one cal lee . Scale : 3 mm.

1 1 ( 10) Co lon ie s b ushy , wi t h cali ces random ly a rran ged on b ranches ; no cal i cu lar rost ra

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1 2 ( I I ) Co lumel l a p resen t ; septa not exsert ; coenosteum fine ly

porous,especia l ly nea r ca l i ces ; usua l ly on ly 24 septa per

cal i ce (F ig . 14 ) E n a l /op sanmu'

u p rofu n da

F ig u re 1 4 .— E n al l op samm ia p ro/ u n do : a bran ch f rom

pa rt of a large r co lon y . Sca le : 5 cm .

12 ( 1 1 ) Co lumel la ab sen t ; septa exsert ; coenosteum not porous ;

usua l ly more than 24 (up to 60) septa pe r ca l ice .

13 (12) Ca l i ces in th e p rocess o f i n traten tacu lar di vi s ion

common (vari ous stages o f sp li t t i ng usual l y p re

sen t) ; f i r st th ree cyc les o f septa hexameral l y

arran ged , septa o f fou rt h and fi ft h cyc les added

i rregul arl y ; nonsp l i tt in g cal i ces rarely exceed 5 mm

i n cal i cu lar d iamete r ; coenos tea l cost ae o ften

ext end up to 10mm from cal i cu l ar edge

(F ig . 1 5) So lenosm i l ia variab i l is

F i g u re 1 5 .— So len osm i l i a vari a b i l i s : a d i sta lbran ch of a large r co lon y . Sca le : 3 cm .

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13 (12) Cal i ces i n p rocess o f i n traten tacul ar di vi s i on rare (most cal i ces , even

temi i nal ones, appear d i scre te) ; septa not hexameral l y arranged ;

cal i ces up to 15 mm in cal i cul ar di ameter ; coenosteal costae rarely

extend more than 5 mm from ca l i cu lar edge (F ig. 16)

Tab le l .— Tabu la r k ey for so l i tary spec ies.

Cora

ll

umat

t

achment

Septa

lhe

ight2

Theca

l Pali

,

pali

f

o

rm

I

Asexual

budding

5

Cora

llumfragi

lity6

Colume

l

la

7

1A tt achmen t o f coral l um: a— at tach ed ; f— unat tach ed , flat bn e; c— un attached , con i cal base .

2Heigh t o f septa : e— exs ert ; n— not exsert .

3Text ure o f t h eca : p— po rce lan eo us ; g— gran u lar .4Pre sen ce o f pal i or pal i form lobes : O— ab sen t ; l — on t h ir d cyc le se pta ;

2 —o n al l bu t last cyc le sep ta .

5A se xual budd in g : — pre se n t ; O— ab sen t .6F ragi l i t y o f co ral l um : r— robust , usual l y in ta ct ; f— frag il e , ea s i l y b roken ;

v f— v ery fragi l e. usual l y foun d i n fragmen t s .7Co lume ll a : O — ab sen t : r— rud imen tary ; f— fasc ic u lar ( rods , tw is ted ri bbo ns , pap il l ose ) ; s— t rabecu lar ( spon gy) .8F la bel/um a labarl rum and F . an gu lare d i f fe r mai n ly in t he shape o f t he ir co

ral la , which is no t a cod ed charac te r i n t h is k ey .

L op hel iap ro/(fem

F ig ure l 6.— Lop hel i a p ro l if era : a bran ch

from part of a large r co lony . Scal e : 5 cm.

Tab le 2 .— Tabul ar key for co lon ia l s pec ie s.

1Shape o f coral l um : a— arborescen t ; e— en cru st in g .

zl n n er edges o f septa : d— den tat e ; s— smoot h .

3Ori en ta ti on o f cal i ces on b ran ch : l — cal i ces occ ur on on ly one s ide o fb ran ch ; co lon y un i p lan ar; 2— cal i ces occur on al l s ides o f b ran ch ; co lon y bushy .

4Cal i c u lar r ost rum : r— presen t ; O— absen t .SBudd in g: e— extraten tacu l ar; is— in traten tacu lar, sympod ia l ; id— in traten

tacu lar, d icho tomous .6Co lume ll a : s— spon gy ; p— pap i l lo se ; O— absen t .7Heig ht o f septa : e— exse rt ; n— n o t exse rt .

8Textu re o f coenost eum n ear cal i c e : p— porous ; s— so lid .

9A rrangemen t o f septa : h— hexam eral ; i — i r regu lar .

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AN NOTATED SYSTEM AT IC L IST

Th is l i st fo l lo w s the c lassi ficat ion proposed by W e l l s

Genera a re a rranged a lphabe t i ca l l y w i t h i n fam i l ies . Geograph ic

ranges i n c lude al l pub l i shed reco rds and data from add i t io nal

specimens exam i ned at the N at iona l M useum o f N atu ra l H i s

to ry, Sm i t h son ian ; Ya le Peabody M useum ; and M useum o f

Compara t ive Zoo logy , Harva rd . R e ferences to s ign i fi can t

papers are c it ed i n bracket s at t he end o f each accoun t .

Suborder FU NG l l NA

Super fam i ly FU NGl l CA E

Fam i l y FU NG l l DAE

F u ng iacyath usf rag i /is Sars , 1872 . D is t r ibut ion p robab ly wor ld

w ide . l n western A t lant ic , k nown from cont inen ta l s lope

south o f Cape Cod , M ass . (4 12—460 m) , A lso

k nown from eastern A t lant i c , o ff Hawai i , and sou th o f N ew

Zealand (285 m ) . Because o f i t s fragi l i t y i t is u sua l l y

damaged or in fragment s w hen co l lected . I t i s d i st i ngu i shed

from other c lo se ly re lated species by i t s possession o f fi v e

cyc l es o f sept a and sep t a w i t h s i n uo u s u ppe r ed ges .

[Z i browiu s 1980 ]

Suborder FAVI I N ASuper fami l y FAVI ICAEFami l y RH IZANGII DAE

A strang ia astrerf orm is M i ln e E dwards and Haime, 1849 ( =A s

trangia dance A gass iz , 1850; not Astrangia danae M il n e

E dwards an d Haime, St ar coral , N orthern coral .

M assachuset ts to Texas from low t ide to 35 m 1 -22°C) .

Th i s i s t he onl y scl eract in ian l i kel y to be seen by snorkel in g or

wi th scuba off the no rtheaste rn coas t o f the U n i ted States .

A lso k nown from Puert o R i co and o ff t rop ical western

A fri ca . Common on any hard subst rate , i .e. , s tones , she l l s ,

p i l i ngs . M ay or may not have zooxan the ll ae . O ften u sed as an

expe r i men t a l l abo ra to ry an ima l . P h ys i o l og i ca l s t u d i es

i nc lude : Cummin gs Jacques et al . Szman t

Froe l i ch an d P i l son an d Jacques N at ural h is

t ory accounts in c l ude : A gass i z Fewkes an d

Bachand

M i l ne E dwa rds and Ha ime ( 1849) desc ribed both A strang ia

astreif orn u'

s and A . dan ae in th e same paper ; A . dan ae has

page priori t y bu t , because the type i s lost , the descri p t ion i s

poor , and th e type-local i t y is un known , i t i s conside red a

n omen dub i um . The type o f A . astreif orn u'

s i s a lso lost , bu t

because M i l n e Ed wards and Ha ime at least designated the

type- loca l i t y o f U n i ted States , i t i s chosen as th e va l id name

fo r th e common shal lo w -water A strang ia o f the eastern and

Gu l f coast s . Lou i s A gassi z , i n a paper read on 15 A ugust

1849, i ndependent ly descr ibed the same species and cal led i t

A strang ia danae , bu t because h i s accoun t was not pub l i shed

unt i l 1850, i t i s a jun io r synonym o f A . as l l'

eif orm is .

Suborder CA RYOPHYLL I I NASuper fam i l y CA RYOPHYLLI ICA EFam i l y CARYOPHYLL I I DA E

Caryop hy I/ia ambrosia ambrosia A lcock , 1898 . Cont in en tal

s lope o ff Georges Ban k m) , A l so

k nown from eastern A t lan t ic and I nd ian O ceans

10

Super fam i l y FLABELL ICAE

Fam i l y FLABELLIDAE

F label l um alabas l rum M oseley , 1873 ( = F label l um goodei Ver

ri l l , 1878 . i n pa rt ) . Common on con t i nenta l s lope from

Georgia to Dav i s St rai t , i nc lud in g Gu l f o f M aine (357 - l ,977

m ) ,°

C . A l so k no w n from easte rn A t lant ic from o ff

H eb r id es t o Gu l f o f Guin ea m ) . Coral lum some

t imes redd i sh b rown . [Z i browi us

m) . The other subspec ies , C . ambrosia cari bbeana Cai rns ,

1979, i s found in more south ern , shal lower waters . I t d i ffers

i n hav i ng a b roader , more open cora l lum and a rougher theca l

t extu re . [Z ibrow i u s

D asmosn u’

h'

a lymam'

(Pou rta les, Common on oute r

edge o f con t i nen ta l she l f from A labama to south o f Cape

Cod , M ass . (48— 366 m ) , 7°—ZI

°C . A l so k nown from o ff

Venezuela , southeastern B raz i l , and , in the eastern A t lan t ic ,i n area bordered by Portuga l , the A zores , and Spani sh Sahara

(85- 316 m ) . Th is spec ies most frequen t ly p ropagates by asex

ua l budd i ng from longi tud inal fractu res o f the frag i le cora l

l um . F iv e to ten smal l bud s o rig inat i ng from one wedge

shap#

ed fragmen t a re n ot uncommon . Th is is the most com

mon ly co l lected coral from t he s t udy area an d i s not l i k e ly to

be con fused w i th any othe r spec ies . [Cai rn s

D el l ocy arh us i ta l i cu s (M i ch e lo t t i , C ommon from

F lo rida to sou thern B razi l , i nc lud i ng Gu l f o f M exico and

C aribbean ; Bermuda (403 m ) , 3°

— 7°

C . One d i sjunct

record on con t inen ta l s lope o ff N ew J ersey (A l batross stat i on

4°C . A l so k nown from east ern A t lant ic and A zores

( l , 500 m) . Some deepwater t raw ls resu l t i n hund reds o f

spec imen s . D is t ingu ish ed from ot her species i n th is genus by

i t s d i st i nct i ve con ica l base and frequent absence o f pa l i on the

second cyc le septa . Cora l la o f some specimen s have a pin k

pigmenta t ion . [Ca i rns

Desmop hy /Ium cris i aga/I i M i l n e E d wards and Halme , 1848 .

Cosmopo l i tan : widespread in A t lan t ic , Paci fic , and I nd ian

O cean s ; Subantarct ic ; o ff Georges , Sab le , and Grand Ban ks

(o ff N ew E ng lan d coast ) ; M u i r Seamount and seamounts

between San Pab lo and Kel v in Seamounts (o ff N ew E ng land

coast ) . W o rldw ide depth range : 35 m . Thi s species has

no co lume l la o r pal i . Polyp l igh t o range . Found i n g reat n um

bers on u nders ides o f ledges in Lydon ia Canyon , o ff M assa

chusetts. [Cai rns

Lop /tel ia p ro l ifera (Pa l las , ?L . p ertu sa (L i nnaeus ,1758) n omen dub i um ) . Common in west ern A t lan t ic from

N o va Scot ia to sou theaste rn Brazi l (95 m) , 3°- 12

°C .

A lso kno w n from eastern A t lan t ic , I nd ian , and Pac i fic

O cean s (60— 2 , l 70 m) . A bundant on B lake P lateau and i n

St ra it s o f F lo r ida as a major con st i t uen t o f deepwater coral

bank s . G rowth rate 6— 8 mm / yr. Systemat i cs [Cai rns

eco logy [W i l son l 979a , b ] .

So len osm i /ia variab i l is D uncan , 1873. Known from on ly two

reco rds o ff northeastern U n i ted States : Lydon ia Canyon , o ff

Cape Cod , M ass . , and sou th o f San Pablo Seamount . A l so

k n own from M u i r Seamoun t ; cont in en ta l s lo pes from

Georg ia to sou theastern B razi l (exc lud ing Gu l f o f M exico ) ;

easte rn A t lan t ic ; south o f G reen land and I ce land ; I nd ian

O cean ; o ff sou theaste rn A ust ra l ia (280- 2 , l 65 m) . Sim i la r to

L . p ro /[f em but eas i l y d is t in guished by i t s equal , i n traten tacu

la r budd ing , w h ich a l ways produces some term i na l ca l i ces

t hat a re i n the process o f sp l i t t ing i n t wo . Polyps l igh t o range .

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1885. N ot ice o f t h e remar kab le marin e fauna occupy in g the ou te r ban k so ff t he sout hern coas t o f N ew En glan d , no . 1 1 . Am . J . Sci . , Ser. 3,

- 1 57 .

WE LLS, J . W .

1956 . Scleract i n ia . I n R . C . Moore (ed i t o r) , Treat i se on in vert eb ra tepal eon to logy . Part F , Coelen t era ta , p . 328-444 , figs . 222-339. Un i v .

Kan sas Press , Lawren ce .

W I LSON , J . B .

1979a . The d is t r ib ut ion o f t he cora l Lop hel i a p er tu sa (L .) [L . p ro lrf era

12

(Pa l las )1 in t he n ort h-east A t lan t i c .- 164 .

J . M ar. B io l . A ssoc . U .K .

l 979h .

# Pa t ch # developmen t o f t he deep-wat er coral Lap /rel ic p e rtu sa(L .) on Rockal l Ban k . J . M ar. B io l . A ssoc . U .K .

-177 .

Z I BROW I U S, H .

1980 Les Sc leract i n ia ires de la Me d i t e rranee et de I ’A t lan t ique n ordo ri en ta l . Mem . I n st . Ocean ogr. , Monaco p . , 107 p ls .

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SYSTEM A T IC I N DE #

A strangia

asrretf orm is

d an ae

Cary op hy I/ia

ambrosia amb ros ia

ambrosi a cart’

bbean a

Caryophy l l i i dae

Caryophy l l i i n a

Con cemrotheca

Iaev i gata

D asmosm i l ia

Iymam'

1 , 4 , 9, 10

D el l ocyarh us

i ta l icus 2 ,4

, 5 , 9, 10

D en drop hy /I ia

gadnan a

D en drophyl l i idae

Den drophy l l i i na

D esmop hy I/um

cris l aga/I i

ebum eum

mob i le

E n a/Iop samm ia

amp he/i odes

p rofu n darostra ta

Favnn a

13

a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

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ACKNOWLEDGM ENTS

Preparat ion of t h e Marin e F lora an d Faun a of t h e Nort heas tern Rolan d L . W ig ley , N a t ional M arin e F i sheries Serv i ce ,U n i ted States # is being coord inated by t h e fol lowi n g Board : N ort h east F is h eries Cen ter , NOAA , Woods Hole , M A

02543.

Coord in at ing E d i tor R obert T . W i l ce, Departmen t of Bo tan y , U n iversi t y of

M e l bourne R . Car ri ker , Col lege of Marine Stud ies , Massach u se t ts , Amherst , M A 02 1 16.

U n iv ersit y of Delaware , Lewes , DE 19958.

T h e Board , w h ic h es tab l i shed t he format for t h e Marin e Flora an d

Ed i torial A d v i sers Fauna of t he N ort h ea stern U n i ted States , in v i tes systemat i s t s to

Mari e B . A bbot t , 259 Hig h St reet , Coven t ry , CT 06238. co l laborate i n t h e p reparat ion of man ual s , rev iews man u scrip ts , and

A rt hur G . H umes , Boston U n i versi t y Marin e Program , ad v i ses t he Scien t i f i c E d i tor of t he N at ional Marine F isheries Serv ice .

Marin e B io logical Laboratory , Woo ds Ho le , MA 02543. T han ks are expressed to Duane Hope , Departmen t of I n vertebra teWes ley N . T i f fney , Professor Emeri tus , Bos ton U n i versit y , Zoo logy , Smi t h son ian In s t i t u t ion , for perm i tt in g t he au t hor to work226 E d ge H il l R oad , Sharon , M A 02067 . at t h e N at ional M use um of N at u ral H is tory d u ring t h i s study .

R u t h D . Turner , M u seum of Comparat ive Zoology , Preparat ion of t h e i l lus t rat ion s by C harles G . Messin g i s al so

Harvar d U n iversit y , Cambridge , M A 02 138. gratefu ll y ac know ledged .

14

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COORDI NAT I NG ED I TOR’

S COM M ENTS

P ub l i cation of t he Marine F lora and Fauna of t he N ort heas ternU n i ted Sta tes is mos t t imely i n v iew of t he grow ing un i versa l

emphas is on en v i ronmen tal work and t he u rgen t need for more preciseand comp lete i den t ifica t ion of coas ta l organ i sms t han has beenavai lable . I t is man datory , wh ere possib le , t hat organ isms be ident i fied accu ra t e ly to species . A ccurate scien t ific names un lock t he greatq uan t i t ies o f b io logical in format ion stored i n l ibrar ies , obv iate

d upl icat ion of research a l read y done , and often make possib le predict ion o f a t t ributes o f organ i sms t hat h ave been in adeq uate ly studied .

Stephen Cai rn s s tarted h i s research on Scl eract i n ia at t he Rosen st ie lSchool o f Marin e an d A tmosph eric Scien ce , U n iversity of M iam i ,w here he stud ied t he deepwater cora ls of t he western A t lan t ic . He hascon t in ued h is stud ies as a Research A ssociate at t he Sm i thson ianI n s t i tu t ion , w i t h an emp hasis on western A t lan t ic , Hawa i ian , and

A n tarctic Scleract i n ia an d A n tarct ic Sty lasterina.

Preparation of t h i s manual was supported i n part by a gran t fromt h e E n v ironmen tal Protect ion Agency to t he Editoria l Board of t h e“ M arine Flora and Fauna of t he N ort heastern U n i ted States . Workon t h e Marin e F lora an d Fa una of t h e N ort h eastern U n i ted States #

M arine F lora and Fauna of t he N or t h eas tern U n i ted States

by t he Coord inat in g E d i tor is supported by t he Col lege of MarineStud ies , U n iversity of Delaware .

M an uals are avai lab le from t he fol low ing

Super i n t en den t o f Documen t s , U S . Go v ernmen t P r i n t i n gO ffice , Wash in g ton , D .C . 20402 , at a few cen ts a page .

U ser Serv i ce Branch , L ibrary and I n format ion Serv ices , D i v i sion0822 , Was h ington Scien ce Cen ter , Bui l d ing #4 , Rock v i l le , M D20852 , at no charge as long as t h e supply lasts .

N at ional Techn ical I n formation Serv ice , U S D epartmen t o f

Commerce , 5285 Port Royal Road , Springfield , VA 22 161 ei t heras paper copy or m icrofiche , for a charge .

M an uals are not copyr igh ted , and so may be p hotocopied from t he

NOAA Techn ical Report NM FS C i rcu lars avai lab le i n most majorl ibrar ies .

The man ua l s so far pub l i s hed i n t he NOAA Tech n ical ReportN M FS C i rcular Series are l i sted below by au thor , t i t le, circular

number , and N T IS accession number :

N T IS No .

COOK , DAV ID G and RALPH O . BR INKHU RST . A n nel ida : O l igochaeta .

BO R RO R , A R TH U R C . Protozoa : C i l iop hora

MO U L , EDW IN T . H ig her P lan t s o f t he Marin e F ringe .

M cCLOSKEY , LAW RENCE R . Pycnogon ida .

MAN N ING , RAYMON D B . C rustacea: Stoma topoda .

W ILL IA MS, A USTON B . C rus tacea: Decapoda .

POLLOC K , LELAND W . Tard i g rada .

LA RSON , RON A LD J . Cn idaria: Scyphozoa .

CAVA L IE RE , A . R . H igher Fun g i : A scom yce tes , Deuterom ycetes , and Bas idiom ycetes .

CO U L L , BR UCE C . C0pepoda : Harpact ico i da .

CU T LE R , EDWA RD B . Sip uncu la .

PAWSON , DAV ID L . Ech inodermata: Holot huroidea .

HO , J U O SHEY . C0pepoda : Lernaeopodidae and Sphyrndae .

HO , J U -SHEY . Copepoda : Cyc10po i ds Parasi t ic on F ish es .

C RESSEY , ROGE R F . C rustacea: Branch i ura .

BOVEE , E UGE N E C . , and THO MAS K . SAWYE R . Protozoa : Sarcodina ; Amoebae .

WA T L ING , LES. C rus tacea : Cumacea .

ZU LLO ,VICTOR A . Art h ropoda : C i rripedia .

TODD , R UTH , and DO R IS LOW . Protozoa: Sarcod ina : Ben t h ic Foram in i feraBU SH , LOU ISE F . T u rbel laria : A coela and N emertodermat ida .

CA I RN S, STEPHEN D . Cn idaria : Scleract i n ia .

15

t GPO 1 98 1 — 798-545

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N OAA TECHN ICAL REPORTS

NM FS C i rcu lar and Special Sci en t i f ic Report— F i sher i es

Gu idel i nes for Con tr i bu tors

CONTEN TS OF M AN U SCR I PT

F irs t page . Give the t it le (as conc ise as possib le) o f the

paper and the au thor ’s name , and footnote the au thor’

s

a ffil ia t ion , mai l ing address , and ZI P code .

C onten ts . Contain s th e t ext h ead in gs and abbreviated

figu re legends and tab le head ings . Dots shou ld fol low each

en t ry and page numbers shou ld be om it ted .

A bs t ract . N ot to exceed one doub le-spaced page . Foot

notes and l i te ratu re ci tat ions do not be long in the abst ract .

Text . See a lso Form o f th e M anuscr ip t be low. Fo ll ow t he

U . S. Governmen t Pri n t i ng Offi ce Sty le M an ua l , 1973 ed i

t io n . F ish nam es , fo l low the American F i sh eries Society

Specia l P ub l i cat i on N o . 6, A L i st of Common and Scien t if i c

N ames of F i shesfrom the U n i ted Sta tes an d Can ada, th i rd

ed i t ion, 1970. U se sho rt , bri ef in format ive head i ngs i n p lace

o f M ate ria ls and M ethods .

Text footnotes . Type on a separate sheet from the t ext . For

unpub l ished or some processed materia l , give au tho r, yea r,

t i t le o f manuscr ip t , number o f pages , and where it i s fil ed

agency and i t s loca t i on .

Personal commun icat ions . C i t e name i n text and footnote .

C i te i n footnote : John J . Jones , F i shery B i o logi st , Sc r ipps

I nst i t u t ion o f O ceanography, La J o ll a , CA 92037, pers . com

mu n . 2 1 M ay 1977 .

F igu res . Shou ld be se l f-exp lanato ry, not requ i r ing re fer

ence to the tex t . A l l figu res shou ld be ci t ed consecu t ive ly i n

the text an d t he ir p lacemen t , where fir st men t ioned,i nd i

ca ted i n the le ft-hand margi n o f the manusc ri p t page . Photo

graph s and l in e d rawings shou ld be o f “ pro fessio nal # qua l i t y— c lea r and ba lanced , and can be reduced to 42 picas for

page w id th o r to 20 p icas fo r a s ingle-co lumn w id th,bu t no

more than 57 pi cas h igh . Photograph s and l i ne d raw ings

shou ld be prin ted on glo ssy paper— sharp ly focused,good

cont rast . Label each figu re . DO NOT SEN D orig i na l figu res

t o the Scient i fic E d i tor ; N M FS Scien t i fic P ub l i cat io ns O ffice

w i l l request t hese i f t hey are needed .

Tab le s . Each tab le shou ld start on a separate page and

shou ld be se l f-exp lanato ry, not requ i ri ng re ference to the

t ext . Head in gs shou ld be short bu t amply descrip t ive . U se

only horizonta l ru les . N umber tab le footnot es consecu t ive ly

ac ross t he page from l e ft t o r igh t i n A rab ic n umerals ; and to

a v oid con fu sion wi t h powers , p lace them t o the left o f then umerals . I f t h e orig inal tab le s are typed in ou r fo rmat and

are c lean and legib le , these tab les w i l l be rep roduced as they

a re . I n the text a l l tab l es shou ld be c i t ed consecut ive ly and

thei r placemen t , where fi rs t men t i oned ,i nd icated i n the le ft

hand margin o f the manu scr ip t page .

A c knowledgmen ts . P lace at t he end o f text . Gi v e cred i t

on ly to those w ho gave except i ona l cont r i bu t i on s and n o t to

those whose cont r ibu t ions are par t o f the ir normal dut ies .

L i tera tu re c ited . I n text as : Sm i t h and Jones ( 1977 ) or

(Sm i th and Jones i f more than one au tho r, l i s t accord

i ng to years Sm i t h 1936; Jones et al . 197 5 ; Doc 197 7 )A l l papers referred to i n t he tex t shou ld be l i sted a lphabet i

cal ly by th e sen io r author ’s su rname under the head ing“L i te ratu re C i ted on ly the au thor #s su rname and in i t ia ls

a re requ ired i n the au thor l in e . The au tho r i s respons i b le fo r

the accu racy of the l i teratu re ci ta t ions . A bb reviat i ons o f

names o f per iod ica ls and seria ls shou ld con form t o B io logi

ca l A bstracts L ist of Seria ls wi th Ti tle A bbrev ia t ions. For

mat , see recen t SSRF or C i rcu la r.

A bb rev i at i ons and symbo l s . Common ones , su ch as mm ,

m , g , m l , mg,

° C ( fo r C els iu s) , 070 , etc . , shou ld be u sed .

A bb rev iate un i ts o f measu res on ly when used w i th numerals ;periods are rare ly used in these abbrevia t ions . Bu t periods

a re used in et a l . , vs. , e .g . , i .e. , Wash . (WA i s used on ly wi th

Z I P code) , et c . A bb rev iat io ns are acceptab le in tab le s and

figu res where there i s lack o f space .

M easu remen ts. Shou ld be given in met r ic un i t s . O ther

equ ivalen t u n i ts may be given in paren theses .

FORM OF THE M AN U SCR I PT

O r ig inal o f t he manusc rip t shou ld be t yped double-spaced on

wh i te bond pape r. Trip le space above head i ngs . Send good

d up li cated cop ie s o f manuscrip t rather t han carbon cop ie s .

The sequence o f the materia l shou ld be :

A DD IT IONA L IN FORM ATION

Send ribbon copy and two dup l i cated cop ies o f the manu

scr ip t to :

D r. Car l J . Sin derman n , Scient i fi c Ed it or

N orth eas t F i sheries Cen ter Sandy H ook Laboratory

N at io nal M arine F isheries Service , NOAA

H igh lands , N J 07732

Copies . F i fty cop ies wi l l be supp l ied to the sen io r au thor

and 100 to h is organ i za t ion free o f charge .

F IRST PAGE

CON TEN TS

ABSTRACT

TE# T

L ITERATU RE C ITED

TE# T FOOTNOTES

A P PEN D I#

TA BLES (prov i de head i ngs , i nc lud i ng Tab le and A rab i c

numeral , e .g . , Tab le 1 . Table 2 . etc . )

L IST OF F I GU RE LEGEN DS (en t i re legend , i nc lud i ng

F igu re and A rab ic numera l , e.g . ,F igu re 1 . F igu re

2 . e tc . )F I GU RES

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UN ITED STATES

DEPARTMENT OF COMMERCENATIONAL OCEAN IC AND ATMOSPHER IC ADM IN ISTRATION

NATIONAL MARINE F ISHER IES SERVICESCIENTIF IC PUBLICATIONS STAFF

ROOM 3361 700WESTLAKE AVENUE N .

SEATTLE , WA 98 109

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WHSE 00528

NOAA SC IENTIF IC AN D TECHN ICAL PU BL ICAT ION S

The N a t ion al Ocean i c and A tmo sph eri c A dm i n i s t ra t i on was e stab l i shed as part of the D epa rtmen t ofComme rce on Oc tober 3. 1970. T he m i ss ion respon s ib i l i t ies of NOAA are to assess t he soc ioeconom ic impac tof nat u ra l and techno logical c hanges i n the en v i ronmen t and to mon i tor and pred ic t the state of t he sol id Eart h ,t he oceans and the i r l i v ing resources . t he atmosphe re . and t he space env i ronmen t of t he E art h .

The major compon en t s of NOA A regu larl y produce var ious ty pes of sc ien t ific and techn i cal i n fonn at ion i n t he fo l lowi ng k inds of pub l i cat ion sPROFESSION A L PA PERS Importan t defin i t i veresearch resu l t s . major techn iques . and s pec ia l invest igat ions.

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out look pe r iod ica l s : techn i ca l man ua l s , t rain ing pape rs . plan n in g re port s . and in format ion se r ia l s : andm i sce l laneous techn i cal pub l i cat i on s .TECHN ICA L RE PORTS Jou rna l qual i t y wi t he x ten s i ve de ta i l s , mathemat i cal developmen t s , or datal i st in gs .

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