The Northern Bay of Safaga (Red Sea, Egypt): An actuopalaeontological approach. III Distribution of echinoids

. . . . . . . . . . . . . . . . . . . . . . I . ln,,od,,itio,, 6 . . . . . . . . . . . . . . . . . . . . 2. Mcil,odr 10

3. HesuiLr . . . . . . . . . . . . . . . . . . . . . . . . . . . I 4 3 I DisLrib.ti"s olcchmoidr and

. . . . . . . . . . . . . . . . . . . frlgmmts 14 . . . . . . . . . . . . 3.1 1. i ieguiai rclii#?oids 14 . . . . . . . . . . 3 1 . 2 irregtilnr eibinoidr 24

. . . . . . . . . . . . . . 3 2 . Corrclnlio~, of erliinoidr 3 0 3 3. hfsltivariale r lat i r t ica l analysis . 3 0

3 3 .1 . Analysis o f n l l echinoid irtagmenl . . . . . . . . . . . . . . . . . . . . . 3 0

.... 3 . 3 2 Analysis of reguiri echinoids 3 2 3 . 3 3 , Analysis o l irregular erhinoids . 32 3 1 . 4 . Anniysis 01 irregular cchi,,oidr

. . . . . . . . . . ~ i l i , , ~ ~ t , Clype,wter sp. 3 2 :! 4 iu lnyar i~on ~ r ~ ~ h i ~ ~ i d fragmenls lo

. . . . . . . . . hotton, mi sedimentary h c i n 36 : I 5 Coirelnlion lo graili size imrt io l ,~, sedimeit-

. . . . . . . . . . . . my pamn,etrr~ and depth 1 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. u.~arion 3 8

. . . . . . . . . . . . . . . . . . . . . . 4 . 1 'rnphonomir hia 38 . . . . . . . . . . . . . . . . . . 4 . 2 Taxon d$sti ibstions 38

4 3 D i i t r i b u t i ~ n of t ixa with r n p c c l Lo ~ n v i r o n ~ . . . . . . . . . . . . . . . . . . . . mental p a r n m c ~ c n 40

4 4 . Relationship to f a r i n dsl r ihal ions . . 4 3 . . . . . . . . 4 . 5 Ap!,licnbilily to pnlnmn,.olol;y 4 4

.inriltilr G i Pniiandusie dor ilnil=nital Wirn, Usiiurr .,ti Ls,~.,, SC,, *.,or0 wen (Mr. addina: ,nr,i,utrir Ceo- logic ,and Pal.%ntologiv dri Univenitit 'Tilbingm. SisrarG drur 10. 0-7400,'ICbinscn I . HRI1.1

The Northern Bay of Safaga (Red Sea, Egypt):

An Actuopalaeontological Appro 111. Distribution of Echinoids

ach

Dic NGrdliche Bucht von Safaga (Rates Meer, gypt ten): ein aktuopaliontolugischcs Beispiel

111. Vcrteitong van Echiniden

by

James H. NEBELSICK'

. . . . . . . . . Contents 5 6. G o ~ x l ~ ~ s i o n s A ~ L ~ , ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ , L I . . . . . . . . . . . . . . . 4 8 4 . 5

. . . Ahstra i l . . . . . . . . . . . . . . . . . 5 7 . Ldrrenres . . . . . . . . . . . A8 . . . . . . . . . . . . . . . . . ~ ~ ~ ~ ~ ~ , % ~ f ~ , , ~ ~ 6

Abstract

Acliiopalnmnlulo~iiai iiiverfigallolis lbsvr h e m cork- <!,,<td on Llie r rh inoid ln i lnafro ln the Norlhern Hay dSsfnga, Red Sen, Egypt. This area was chosen lor stlllly hs i t rcpresrn~s a 11ighiy strscisied. shallow water, tropical, carbonate cnvironmenl. Echinoidr frrgrncnls were common i. 6 7 balk sediment r a m p l a which werc ~r r l Ie r l ,~d i n n w i e l y f lbollom and srd- i~nen la iy facie8 down to n depth o f 57 n,. The l r a g rnentr car, be iden,iSed duc to the numero,is c h n i ~ acterr p rexn , "I, ,he edrinoid lest and spines ,upon comnnri~on to zcbwchins round willsin l h r

r,.,,dy nien and muscsm c.ollccLion~. The l i s g m n l zmateiial of 18 l a x a wsr d isLw

grirhcd, with tho exceylion C ( y p e ~ L r r sp. and

Sdicnsle~idae, aL ihespeciw level. Ho l l i regular and irregular echinoid fragmenls were prcsenl i n rixrillar amo,,ntr.

MullivarinLe staliilncal analysn (durter and p r l l i ~

r i p d oinponenU) and ro,,elalion metllods are used to nnalyre the dislr ibulion off iagmont weight8 of ai l ecl,inoidr, regular ccbinnids only, iiregulnr echinoidr only, and irregular ecliinoidr r i l h o u l CIypeartei sp. 'The m u l U of IIx~sc annlyser were ~o lnpnred l o the rlirlribuLion of bo t tom facics (P ILLEP 61 PEU- VI:SLEIC, 1989), sedirnanlnry l a c i a ar l grain

sire dislr ihulions (I ' ILLER & M ' 1.90)

'Vlh di,er, rolrlpniison olechi- < l h

delaiied rediilientologicrl

....~ llcslc "Oil rrg,i1arrn iiliil ,rregaiire,, sec,gein

.,,id 11, c , w r gie,rlir,, Mc,,grii ar,grlrolfen n,orde. oil. I d e i i ~ d i z i r ~ i g dcr P Y I I ~ I ~ ~ ~ I ~ ~ erlolgi.r anhand lcr mlrli<:iclien hlrrki,,.ilc dct i~cl,,,,idencclrale und slnchrlii. I hc frngmcslr wiirilen n,it konlpletl e i ~ ~,sli.erien I:xrrnplarm, die im Arbeltsgebiet gesarii- ,i~rll wurden ode, sic17 irn hliiseesbrsl,nr,d befnden. v < ~ ~ ~ I ~ c I > " , ,

h l~ i l t i ra i in t r Cluster- uiid l Iai iptko~sposen1en~~~a-

an , grsamtes itagnmlmaleriai. sunder" nuch gdrennl fiii die rcgiiliien, i i r*guI i ien aowie i i ~ rcgulir~:n Sccigeil, ohnc CJypeirsLcr sp. d i i r ihgel3~r t . IOir I:igci~i,irrc~ dirsei Unl.eisucl,ungen mnd m i l den Vertriliroges von I i udcn tind Sedimes~tlaxirstypen suwic clrn niitc,lc~res Korngrhllen verglirhen wor- CIW Dirckfe Vrrglrirlir dci Echiii idenvcrteilu~~g mi t drtnilliri~.rs srdio~eiitologisrlieil Paranielern lind ~ ~ ~ ~ ~ ~ g ~ i i ~ i c i , ~ ~ ~ ~ ~ i l ~ ~ t ~ g ~ , , (I'II.l.Ell k I'ER\ll:Sl.tR. 198s: I'll,l.cll. k MANSOUR, 19!10) r ind i n .15 I w l e ~ i tnbglicll g w m s l l ic Verteilung dei iicll~~,~~lcnirag,,~colc wsrd<: cuch is i~r rha lb dei jewri- tige. ~ ~ , ~ ~ ~ c ~ ~ i ~ ~ ,I~.I st:diillcsirnrics u ~ ~ ~ ~ s ~ I c ~ I . . lh lirgcbnisse r l r i Aen1yri.1~ icigen. dnU die

t~ci~riliiiig von i:rliinidcn I dereii Frag- I I W ~ ~ I ~ ~ l a t i s t i ~ ~ l ~ I I I I ~ P ~ ~ I ~ I w ~ r d ~ n k l n n obwoiil VcrinIschiiogcn d w c h tapiiosoeiisclir Vorgisge rmil ~ ~ ~ ~ t l ~ o d i s d ~ hc&ngI,e k ; i n s c l ~ r ~ r ~ k t ~ ~ ~ g c ~ ~ l~ez i i g l k l~ dm !,,lt<!,s,t<l,t~" l+&g">cntgr"h (> 2 m m j aUftml,">~. 11,~. vmci~iing dci liiilinides i nm S~CII

hcv:ils publ8zierlrri i(us1ilaliveii UiiLeisucluiigeii rou Echis~Ici , a m dern Rolea Maei bind drin 1,idu \vesl.gszi(ik verglciciirl, 1:s q i b t sic11 einr cngc Herirl~ung iwirr i ien 1ic l i i i i ideevrrtei l t t~~g w d Sediment sod Bodesfuies sowlc I(orsgr68r oiid i\'asseihcwegung. Die Mclhodcii, welclke iii diesri akluopaliuritologiscl~e~~ Unlersiicbung arigcweridet sordeo sirid, rolll.en auch fur dnr Erkmi i rn dci Vrrlriliing iossilrr iic1,isidrnfauiien Verweidiiiig h d e n

A ~ 1 ~ ~ ~ ~ ~ 1 ~ ~ ~ o ~ ~ l o 1 o g y has i i ~ c i i d ~ s i i i b ~ t l hy I I O l i l l fi. SrANTON (1990) a? the study oforganisms in their ~,,<><Ic,,, e,,",rOr~m<m~t rrom a geol"glc/pal~o~l l~" l"gi< ~ x : ~ ~ p ~ c I i v ~ a d w e h p ~ i by RICHTER (1929). AIJCI. (lS15) reid SCllAl ' I i l t (1962). Thc Nor~l iere 13~) - or sacagga, s i~sn~ed os the I:~YIILC~C, WCS~CU~LIC,

or tsc Red sea (Fig. I ) , w w chosce by resenic11i:rs at the I,,rliti,l,e of I ' a l x .~~ , l .~ l~gy of the Ulii"eisity or V i c ~ a LS the object o f ~ u i h ashidy a il. ieprrresls a lligl,ly stracturt:d, slrallar w a t ~ r , ~kopii-nl, ~a ibona t r esvironinmt. 'The broad scope o f research imludcs the ge,ierhl i,i"rsligallun 0s eliviroririlelital paisin- eters, topogrrphy, and bot1onr lacirs (PII.LEIt & i ~ t r t v ~ s ~ . ~ ; ~ . 1 ~ ) . redio,c.my rsciw ( I~~LLEI~, 1990. I'Il,I.ER k MANSOUR. 19RO). a wcll rrr a wide rang,? of detsilcd s,,,dics on idevast orsanis,,,s or pa~aeosto~ogic interesi inci.disg rornminihs, red a lga . biyoronns. molluscs (GOI.I33IOWSKI, 1930). r o ~ r l s and boiiug bivalves (KLEEMANN. 1990A, 1990B. 1992). echinoidr (NEHEI.SICK, 1!190, 1992).

lebensspuien ( I~WORSCI IAK & PI<ItVESI.IIR, 1988; PERVESLI2R, 1990).

BOGGILD k ROSE (1984) showed t l i r i r i ipar~ ~ m r e or drawing spot, reccrit asnlogiies i s disl.iit~ gilishing biohcies and l~alnroei~vironn,eiiIr irjr ec l i i~ noid assemblages i s tl,e Mid-Terliaiy carbonate s i i c ~ cession or the Maltese Islands. There are. I m w ~ ever, nimerous problen~s i n awessing recent echi- ,,"id distrii,.~.ions and their inipiications ror tological research. T l lae include tlipir cryptic and pitchy dirtriholionr (BUCllANAN, 1966; I M F N I & TOBOI,. I986), as weli as the intricate tapho- soinic, pallways of 1.llesc muili-element skrleloss (SCIIAF1:H.. 1962; SI<Il,ACIIER, 1979; S'I'KA'~l1~ MAN. 1981: SMITI I , 1984; GR.EL<NSTEIN, 1989, 1990, 1991; GREENS'PEIN k hll?YI<A.. 1990; K I D WEI.1. k RAUMII.1.13R. 1990: DONOVAN, 1991). Research on the clistributimc n d controlling envi- ronnirntal ini torsofseb iwcliins lisvi: h m i i e r l i i i l ~ d l o largescaled drrr i ipl ive s t i id in (KIl<R. & GIMWI', 1965; I3RNST I?!' AT.., 197:3 KlER. 1975). as well as qaaslilative studies of regular ta rn over siriailcr

Fig. 1. Location atid geneial topogiapl,y o l t he study area; modified aftter PILLER k MANSOUR 1990.

SCI IUI~MACHER, 1974: CAMPDELL 8z M O R R l ~ SON, 1988; nr: BEER, 1 9 s ~ ; now^. 1990~) . One method i n overcoming the dificulties of ss-

sesring eclri,mid dirtribirtiunr over larger areas and srross facies boundaries l ie+ in the identification and

the distribul.ion of boll, regular and irregular echi- noidr with the Eacirs. grain sire parameters and en-

vironmental factors d the Northern Bay o f Safaga. I'lw knowledge gainrd simsld be aschi1 i n arriving at a more complete ~canst ruc l ion of ancient ecllihmid biococnoses from 'he available skcietal rer,,ai,,s.

'Two studies serve as a basis for this siudy: ,lie topography and hotton, racies nre presented in

P1LI.ER k PEHVESLER (1989) and the isvrsl iga tions ar sedimentary iacies mid grain size parame- ters i n FILLER k MANSOUR (1990). 'The study area, a shallow water carbonate environment, shows a wgged bottom morphology cl~arnrterized by N-S mnning topographic highs which are mani- fested either -3 islands or submarine ridges (Fig. 1). The topography of the bay was %-red using echo

sounding along 55 transects with a compiled lengtir of 125.8 km. The shnliow "Southwrsl channel" is bordered by the mainland and GaziraL Safngalsland. I t reaches a width of 1 km and depths which deep^ ens from 5 m i n the south l o 30 rn i n the north. The pmlected "West arc*" Ills n basinal character and reaches depths of 38 m i n a Rat muddy plain. 'The "North area'' is generslly shallow with a mcau

Rcili Palaonl 6r lc ir . . 17, \\.,en IY'J? NCBI~1,SICK. J . 11 , S h e Northern Ray iii Sniagn

Kcgular echinoidr Cidaroida

Emidmi= meiuiow (iamarck) Pl4yiiacmfhw impmidis (Lamarck) Prioaocidari.r haacuioso (Lamarck)

Echinolhurioida Aliiienor",,'" voriun, Gmbe

Dlademaloidn Diademn ieimutn (1.cske) EchiiioihrLx coinnin,i.r (Pallas)

se"m"plurioi& M i m o ~ p h u r roiisseoui L. Agassri Ternnorre",o sp. Niviechinur s p ~ TripneuJrer p v i N n (Lmnaeus)

Echinoidn Eciiinorrirrro mnrhnei (de Blainville)

n,hirh serves M the bus oi roral reds extending Lo 11n suriace (e. p., ' h h y a AlLKohir).

'The Norlhern Rqu of S a i a g a ir chnrnctpiined by a inre ~ ~ ~ i ~ h i l i t ~ and mpid ~ranr i t ion of f a c i o within a relstiwly m a l l are, (ca. 7 x 10 km). Twelve bot- ~ o r n inrirr were dislinguished hy FILLER & F E R ~ VESI.ER (1989) nr reen in Figures 19. 23 and 28. ' R a e represent the observed ihaiacleristics of llie rebfloor and nrc based enrllisiveiy on mnp-

Fig. 2. Lo.:rtion of sample pomin ( A m = Aerisl mmi. H = "Saisgr hotd")

'L'lii. i,,,<>sl,gnlio,i of *he sedimentary f a r i n il'il.l.I:11 & MANSOUR, 1990) revealed s mined ..rl,",!aL<:/Lerrig6"6"6 enviranmenl due to the high ~ u , c c n l r a l i ~ n of non-rarbonale minernir along llir ioasiiinc. Of inlerest i m the interpretation of echi- ,noid fragmmnt distribution is the irnmedinte neigll- iborhood o i contras~ing sedimentary environments, crprrinliy that of ''Coral carpet" and muddy sedi- nmen~s. l:igiil sedirnenlary fxcies (see Fig. 20. 24 and 2!Ji rrx: dirtinni~ished. T l m c renrescnl a svnthe - a ~ s of gmlu si re , carboime constituent. rnineralogib i d and 1rrre rie,nent charac1eristica of a Lots1 of 122 .~;mpIcs colicrlpd t b roag l i o~ l the study area. The ,r~:f.&d docsmentation o i grain size paramotors (see Pig. 21, 2 i rnd 10) is o i special importance for lhe nealjsis oiechinoid dislribalions.

2. Methods

T h e analyzed echinoid iragmcnls ( I 8 kxa ) originate from 67 standardized bulk sedinienl samples (Fig 2) takrn by pushing a sled cylinder with a dinanr~ ter of10 cm into lhe sediment and removing the top 20 cm with a m e l d scoop into a net with I m m mesli ~ i d l l l . The chrrncterislics of the rsmpln pui,,ts arc shown in A p p 1 Echinnid fragments larger tlian 2 m m were removed. identiced and weighed to a pre. &ion o f000 lg . Large r s m p l n were ~ p l i l t e d using 1 modified sample splilter ns devcrihed by KCNNARI) br SMITI I (1961). The weigh1 percentage? o i lhe split samples were iecnlculaled to the original sample size. The o f echinoid fragment praencr i n each sampln are shown i n App. 2.

Rhinoids were discov.red a. l iving rpeciincns.

dead pmpty l n l s w i lh and without ptesrrvatios of l l l e apical system, and ar diverse frag,nentr of the lest and +es ( ~ ~ m p a r e ' l s h . I ) . Theobseived p i es~ ente of echinoids within the Northern 13ay o f Saings is rompiled from numerous dives throegh thc atilllor and other memhe,* "i 1he working group.

ldeiwiicalion ofec1,inuid iragoet\i.s iailowed "pun corspaiwm to complete speclnrenr rollecled m lire field ( idml i i i ra l ion foilowing MORTENSEN ( l a 8 1951). CLARK k ROWE (1971) and DOLLFUS k ROMAN (1981)) as a 4 a\ colicctions of Red Sen echinoids i n the "Naturh idor i rc l~n Museum. Zool- "girche Ahleilung" i n Vienna. Tliere are numer- o m rharnclem allowing the identdcation orechinoid i~agmenrr N summarized i n MORTENSEN (1928 51). I IURBAM CT Ai.. (1966) and SMITH (1979, 1980A. 1980B. 198oC. 1981) and docszmcnlcd i n PI i to 8 ' i ' l ~ s l j e include:

I . general tw1 sire *nd oalrphoiogJ

6. sine and eunrhei oigeni la i pores;

7. shape, diflcrenl,atl". and d,s'rlbuli". o f tuber clcs;

11 internal support syr1em or clypeaslcroids;

14. and hne slructsie (?idges, wharlr, barbs and llrorns) of spines.

Most. irngn,ents could bc assigned to the species level. Idenli6cation of Clypcaster was restric1ed to Lhc level, that of schirasterids to the i a m ~ dy level dthough larger fragments of Dipioporasler ~ ~ " i ~ " ~ i were encnun1ered

echmoid tnxa nciors the Nortliern Bay o f Safaga w e dmwn ,a Figs. 3 to 16. The r~~rnu ia l i ve weight per-

offragmcnls t!,? - = percclrtngr o f test fragn,eets and spines within l l le inxa are shown i n T tb . 1.

The Pear,soni prodiiiIGinomen1 correlation c o d ficient r, niesruring l l ic linear relationship bctween two variables, is used to mirelale between rbaslule ecltinoid weights (l'nb. 2). grain sizr parnmelers a;. well - dep1.h (Tab. 5).

C l i i r~er analysis i r n i ru l l~vnr in lc rlatislical te&

,,,,ue wl,irh can bc iisrd Lo identify n smaller "vim.. her g'oup" SUCI, that eicmeutr ,=idin& i n a pnr- t,c,,Iar group are more simiiar to each o1her t!,nn a,,, clemenls belonging LO otlier gmups (DILLON k GOI.DS'TEIN. 19R4) 'The relnlioiiships of the s a m ~ pier are displayrd in dcndrograrnr whereby the clor. csi r~ la t ioss l~ ips are s l ~ o r n bctween objects nearat ~ ~ g ~ t l ~ e r . four h i r r a~ i i i i i a l Q-mode cliisler analy- sea were calculated !using the the cosine cocfli i irnl as the similarity iuclcx and t h iinwei~lxted pair-

" . = the classification procedure: total eclninoid pres- ence (Fig. I?), regular ecihinoids only (Fig 22). i r ~ regtzlar ~ d ~ i n o i d s O Z ~ ! ~ (Fig. 26) and ii iegular cchi- inoids excluding C!ypralei s y (Fig. 27). Samples i n which the tax& do no1 include more than I % o f the w ~ a l echinoid weigh% percmLagc were cncluded from the later tl,ree analyses. lnleresling results are given by the r lvslei arialysis of total, regular and i r ~

erhinoids without Clvpmasler rp. Slatiaticnl parameters for ihese i l u r l e i analyses srr picroi led ~. in A p p a, 6 and '1.

The diat i ibst io i~ o f the irsuil ir ig cliirLeir with re^

t o ~ O ~ I O ~ fnrics, srdim~n~.ary raries n,ean

grainsize aresliown i n Figs. 13-21.23~25, and 28~30 . The faries and mean grain size d is t rhmons are s i m pliAed replications o f the maps published in PlLLI:R & PERVES1,ER (1989) and PILLER br hlANSOIJR. (1990). 11, is imporlnnl lo note that t l w e maps rep^

rrsest a generalization olfacies and grain size dirtrib I>.li""i.

A Qhnode principal m ~ i p o s c s t s a d y s i s was ion^

Conipsrisun t o g r a i n size pnrametcrs

duired on the total echmoid frrglnent s,ater~al of67 The cordat ion l o batlorn fac ie (Tab. 3) were made

~ w n p l e r 'This ir a niet i~od for the overall following Tab. 2 i n PILLER h MANSOUK (1990);

culnplclity of mul~ivaristc data bytaking aduni,tago l l le position o f the ssrnples i n bot lom facia map and

of inhercnl inler-dfpendencin (JORESKOG e l al., selected profiles a m = the Northern Bay of Safaga

1976; SCNUUO k UEIILINGER, 1986) by ertmct i n PILI,ER h 1'ERVESI.ER (1989); t i le descriotion

loadlngs on m n ~ ~ i g ~ i l i ~ ~ ~ t l y ronlribuling uniinblea. The factor scores ertiinnt,e the ronrrihuiiun of t lw factors to each obseivntian (the rampla). The factac loading of the individual unr iabla (the rarnlmnenls) are weighled with ieapect l o the con^

trihutioii of lhat varinhle to the ractnr. Tl ie factor loadings are given i n App. 4; the focLor scorer and

of variance arc shorn i n App. 5; fnclor loadillg plots are shown i n Fig 18. A l l inull ivarirte rlalislics were computed on the ma inh ree Sl'SSx r taf i r l i rn i package nl tlie University of Vienna.

~ n n l ~ l ~ ~ and sediment ramples taken for grain size analysis (see F ILLER h MANSOIIR (1990) for s a m ~

methods) miginating from the same locatioll. A n ,igmnent ro ~edimentnry fac ia (Tab. 4) was possible for 39 out of 67 cases (= 58% o( the ram^

pies) The direct carrelation oiechinaid prerence to grai" size fraclions and parametcn (mean grain size; and sorting) i n 45 out of 67 cases (= 67% of the mmples) 'Tab. 5). The distribufion of echinoid rragnizlll weights wi lh in discie~e mcnn grain size val- uer are sLown i n App. 9. The s~d imen l description

a ~ c d ~ ~ l h hard lubsrialcr ~ ~ ~ I ~ ~ d i s g the rota1 r e d cornl carpel bottom racier. Modcrale porcent-

q e s are iound i n sands associated with coral patches and seagr-. They a m most common i n the coralgal r e d i n m ~ n r y racies

Eucidaris meldaria (24.58% of total fmgmenl weighls)

Observed clistribution and mode of life: Found living on a w i t t y of hard suhrirates, often i n i r ige nurnberr belvee,, the hrallchm or corals. rd&ific.tion: Small cidaroids (< 3 m ) ; wil l? d i r $. iec~ly sinuatc amhularrn (PI I , Fig. 2. 3) wi lh nun^

~ ~ ~ j t i g a t r pores by n raiinded prominence; IrLmlry teheriles ~ i t h snlw~li gl-y knobs on the adriledim side the bolr mil pai l ly also a l the 1nnmc1on (PI. I. Fig. 4 ) ~

lnteiarnbiilscia have son-cienalatc, ~ ~ d o r a l e d h r - bmles wi lh hio~l-sunken areoles (PI 1 . Fir, 4). ~ ~ ~ ~ l , i c u I a r ~ ~ b ~ ~ c l e s with semi~ell iplicrl bms and mamelon prolonged toward the pr i~nnry tubeirle and cyliildricsl primary spines w t h ahiaptly lruncnted

~ l d t kmlirbating :" ~ o w o with a central promi-

nence (PI. 1, Figs. 1, 5, 6). Low rounded waits arc disposed i n regular, longitudinal serin. ~ ~ ~ , h ~ ~ ~ ~ ~ : = io.xn%; = 8920%). Spinn or living sPccimenr are commonly incrusted by a number a[ diverse including coralline .Ige, forarnini!ers, bryonoans, and bivalves (PI I. Fig. 6). Clean as ell as incrlisted ~omplete, empty test are (PI. 1, Fig. 2). %L !ragment show mostly i n t q h t e h r t s r i n g (PI. I. Fig. 3). nlssment distribution (fig. 5): F O W ~ i n mud- erntely coarrc and gravels i n s wide disti ihulion of rllhstrater -cisted with hard sub- rates including coral reel. cornl carpel and rand ~ i t h prtclles ~ O L L O ~ I lacie, and c o d g a l sedi- men1nry rncirs.

Plryiiacaethos inrperkiis (1.58% of total f~ngrneol ceighls)

Liv iog were not observed in the atbid? area.

~dest i f iaut ion: ~ a r g c c i i lsmid warla widely e m i t oc- ,,Iats in the sys tem S isua~e ambalacrr w i th cu,,jug*le an,belnrrnI pores l,rve an i.terporiferour zone shout t i le same width 01 the pore zonr T h e i i i lcrarnhulncrn show well repninlcd. sunken nrmles wi th a prominent. "on-crcniilale, pr in inry luberr ie n,d a large b o w (PI. I , F ig 7) f'romincnl se in i l i i ~ nar sciol,iculnr lubercler, w i th n d i r l i nc l oleualion on llh side lowards the nreolc, a x larger Lhnn the s e r ~ osdaiy lubercles. T h c median aim is very narrow 'She Inrge. robur l , fus ih rm primary spines rhow d i s ~ l i l rc t fine giniiules arranged i n 16-20 regular longlb ~ i i c l i nn l m i e s (1'1. i . Fig. 8) T a p L o n o m y : = 27.88%; r p i i m = 72 12%) A ~ ~ ~ ~ ~ ~ L ~ ~ spines, shuwi t~g a wide varicly 01 p i e r r - ~ ~ t i ~ ~ , wrre round i n sand porkcis 11,~ rrer. Fr"f""',i d i s t r i b u t i o l l : M o ~ l common i n mode.- aldy sorted, coarse sands i n l l i e coral i re1 and sand d l c o r d carpet b o l l o m i m e s and i n lhe r o d g a l sriflmestary fiarirr.

I.wing specimens w e i r no1 observed i n l h r d s d y ...... I d e n t i t i c s t i o n : Large i i d r i o i d wit11 iriserl ocsiair in the apical ryslenl. The ronjugr!r ae~b i i l acml oores lhrve an inlcruoiifeious zone about twice the

well 8s <",,fluent am&s (PI. 2, pig. I ) . The median area can reach s wid lh greater Lhan

that he 'Ttie cylindrical or tapering prib rnary spine lhnve aspol led collar, alongiludinalsericr o l l homc nrrnnged i n d i r l i n r l whoi ls and r terminal mown ( I 2, Figs. 2 ~ 5 ) T a p h o a u m y : (plato r 1256%; spine. -- 87.44%) hlostly round N ~ i n g l c large inleiambu1;icral plates ur~eii w i l h ar i icu la lcd nmhulncml piales (Pi. 2, Fig. I). The spines rhow n wide v r r ia l i on o f p rev rva l ion dm being lhiokea (1'1. 2, Pig. 4, 5) and i n m s l e d . F r a g m e n t d i s t r i b u t i o n (F ig 6): Wide rangc ol dir t r ibui ios i n niadcraleiy sorted coarv rands and

NEBELSICK. J. 11 , T h e Northern Bay ofSaiagn

grmcls o i the coral caipel and sand w i th coral coral reds and c o r d cnrpetr. pn~cha bottom lacia and the corilgnl u d i m e n t a i y Identification: Ainbulacra, which are iontipic.

iacies. I t is also present, ar the regs- uously ~ i d e n e d orally, contain very narrow pore lar eclrinoid. in the m u d ho&tom iacies wi th a carre- forming lleaily a ~ t m i g h t l ine. 'bigerni-

ipondlng high presence in the msd, rnol~escan, and omp pound plates lrnve conspicuous, crenulaled,

operdna sedimentsry iacies. piilnnry tubcrcies ~ i t h wry ~ ~ n i i wc- ~ n d n r v tuberc les l.niqe, inteiamlrulacral tiihercles (PI. 2, Fig. 6) are well separated f rom one another wit11 a naked median space ociining adapically (PI. 2. F i r . 7). The very long. (> 20 m ) , slender, hollow,' ~.

Observd dirtribetion and of life: C o t , spines are conrpiciiourly vertici l lale (PI.

man i n groups o i up t o 20 individuais of different 2, F ig. 8, 9). sizes ~l 111c I ~ Y OE oral T ~ C ~ d r o Tnphormrr,y: (platcs rr 58 74%; +as = 41.26%). ubrrrved on the reef (In1 m well rr. hetween corals i n N e w found hr compiete emply t a t . Fingmentr

i,rmt.nt IS mla te i i ~ ' l r l c s and i w s l i y short scgr,icn&s (< I m,) of I h h s p m s (1'1 2 , 1:~g 8, 9) Emg,,",'t ' l i s t i i b n t i o n 7 ) \v idr ly i l ls- tril~ii~c,l, $most f r q u e n t $,, n m d m ~ t c l y s m t m l , ,,,,.<I ,,a,,, Smds x,, LIP< samd wit18 c o r d ,pd"I!es I>& torn (n&s r,,d trrng'."""s rrrliriieiiLaiy fncirs

, lri,"'"".slr:s , g i - s t i l i a

( 2 48% 01 iota1 f m g n w n ~ wcngl$l8)

Ohsr tv r :d d i s t r i b ~ ~ t h , ~ and mode o f l i fe : i ' h ~ ~ i ~ g ~ t e d tcsi o f l iv ing r p r r i i n e ~ s show a wide color iirra,ioi, l'l>ey sic iiiiiimoi, iii rhali"w wn1er. es-

ixxhalli wzlI,in rmagihrs imrrdowr of lhe "SuslhwrsL C I , ~ , ~ W ~ , w h m l h c sr.niiichins c u v c m l thro,srIutl

. . . rys l r rn, r,Yr ro,in,1 iii Sengin%T nieas. 'i'hesr

were also observed tiai,sporlcil eithcr oulo l h r bench or airoir t i le rccf lli~l down Lhc reef face into dceper r a t c i i'ingiricnts tiiiisLiy show, in t i rp la tc fmchir isg (PI . 3 Fie. 3. 41

Echi,,ornctra i r m l i i n c i

(10 4% or l o l d rragine,,t wr lghU)

O b s e r v e d d i s t r i b u t i o n nnd rno r l c o f l i fe: Liv- i r i g~pmimena wrre hidden beneath stories and coarse iiiatrria~ *tong roast "r IIIC O S ~ ~ ~ ~ ~ ~ ~ ~ ~ L rhasiier~ 'Thry ncrc also obrcrveil on Lhr reel flat of Tubya ,\l-l3ajcIa. ~ ~ ~ < : , , t i t i ~ ~ r i o ~ , : inistlilitiy i:iongn!,~ LCSL (PI. 4 . F L ~ 1, 2, 4) w i th poiygnious arribulncia mntain ing4 ]pore 1prirr to r a r h arc w r l l n5 iinp~rfoiaie s",i~crenulntr pcima,y r rnb i~ inc ra i t$ihercles (1'1 4. Fig. 2-6) in-

tciainhiilnria Ihve r vrrt ical series o f primary and secondary l \ iber i les l l o r i ron i r l s r i r s of 5 tirheri lrs to mill plate occur in the ainbilai iclgion. Ace l r ,

i la lm-ocet i l r r , l i is i $ ia i i i , n i i i a lus

(216% of loin1 ringrnesl wrights)

obrprvc.c~ d i ~ t ~ i t , ~ ~ t i ~ ~ ~ al i lr~ oc lire: very ionsliicuour ~ d , i ~ ~ i d r o d iypicaliy be- lwrrn rorn l lhrnilr on i h c r e d edge i n very rh r l l ow wrtrr Ollie, ii,,ing rpcc,,,,ess wrre ol,sewed wi th in c o r d rarpeir l< l~"~, i l ic~t i<, r , : ,,,&%i"', <:lo,,~~lc<l k S L (PI 5 , Fig. I ) w i th large polyporous n r s l ~ i ~ l a c m l plates

Rare pxampics o f lhe ed~ ino lhero id Aslhenosumr variirm were observed i n deeper w r l r r s o f the study area (> 30 m ) wi lh in thr reel M well as on muddy boltams. I L hns a largr, low, lherni~phcricd, "cry Rexiblr test will, inlbricnling plates. Fragments o i Lhc les l weir no1 discovrrrd it, the rnmple material lsoln.cd ernieples or I jchinuLl>rir calarnrris were ob- srrved at the base orcoral patches as well = on the reef RIL. Nei t lw i 1ragrornlr of 1.11~ pr i innry spines wi th a large cilviLy and m c x l ~ r n a l trnnsverse series d l huiss, nor larger cl irdemnloid ~plales which could be nr*igsed to i l l is genera were found wiLhin th* an- slyrcd snrnple material

Singleisolaled livingspeciincna as well ~ c o m p l e l r t-5~8 of the temnopleuroid Microcyphur rousscaui wcic obscrucd wi th in coral pa1dtes i n shnllmv w n ler as w d i as i n sand p r l i l m w i l i i i n reefs They arc e&-iIy rrcngsirablr the naked wehi of l h r 1nt~erwdmlacra (PI. :i, Fig 5, 6).

,;cg 22 U w d a L r u n d a l i ~ r i ~ r c l i i c a 1 U ~ m a d c UPGMA diir

beparated r curinc rneficient of 11.6 (N = 5 5 ) ~

3.1.2. Irregular echinoids

Tlir ,rregulrr ecii~noids oi the sttidy are b a i ~ rowing deposit rredrrs &,,d cnrnprise 53.17% thc tovri fragment weightr. NO s,a~erisi or i r r c g u ~ tar ec~,inoidi round in tile mnip~e iastriia~ dur l o tl,& m,all sire. liiegulai echinouds show a r i d c d is l r ibut ion ( F i g 4 ) i n all bottom arid scdimenlaiy racies. r\iiholigh oiort lrregliiai ~ ~ i ~ i ~ , ~ i d tax&

~'iPS""1 8 " ~i~"~ l i~* " l si.o,iiilr, only r"w ir;,greeu1s o r lhe irig~ ~ ~ ~ l m g ~ i d Mrtaiis sp wrrc dis~oueied ~;LSI.WAI~S 0 1 T u b y ~ ill Kchi, bin 25 nwl.crs d~pL11.

i ' l ~ p ~ ~ " . s l c i sp. (29 i:V. of W : % l fr~g,,w,,t w<.mghts)

Olia,.i.v<:d s t i i 11,d ,,rv,Ir o f l i f c : (:Iy".~'Lcr 1,iiiiiiIii (PI 6 , F ig I. 2) W h \ ~"BIII~OIII~ i,,,,,,,l tliioiiglioiil t1,c s i i i i iy arcs, ,mi rli~r1iiiw <oars<:

,,iwl*, iix s%,inIl sziid l~iltcl,rr r i L l i i i i sragius i i ~ r r d ~ .,ss, nnil r i l l > i n L l i i i1, i i i sand w:rsrrs on id IIrLr ~ 1 ~ ; ~ I l o ~ v l y l ~ v w d l > ~ ~ > ~ g s~wrtmtms wrc d > s m w m d P , .

0 x 2 ,I,,? C>LS,CT,, s1,c>rc or ,I,? "S<>L,~ ,~ ,W~S~ r1,m 1 'Ti,rm sprn,,,,wr or CI.Y,>CNLCT /?~W,,S WCiC

. i i ~ o ~ i ~ d I h r d $7, I io 3 r m sedio~eiit d q i h i n s2

.li:~~ili is $ 1 ~ s o i i i l ~ r ~ , imrici or the "Eni~rrs arc*'' <),,I? #lW,<l k S I S #Of <~ ' l~p~; ,~ lw ,?,,c"I~*I~~s WW" i",,,,<I

s l ~ ~ ~ l l c ~ ~ v & p I % s to $ 1 ~ w ~ s l , oi'TuI>ya AI-13ayt l~~ m d I , , cl"ep~r ,,wddy sm<1s l<l<!,,til<,:ati,,,,: rl.qXtb,<cr l,,,,,,,l,s ,s ~1,a.acLmzd I,, CIA, <>mI S~,IC~, ~I,,s~,I sm~t;,l ~,,,l>,,la~r"lh, n,xI sm;~Il pr taI$ (1,) 6 , 1Vig I , 2 ) CIypc%$t?r f c r w n $ is &ar- ,A<t<:.,d,,d \by a h r g c l>",al<d a w a ; a d 8, ~ I i ~ , , d l ~ <>,>,w i r m # d ~ ~ 1 h d ~ ~ r w ~ ( P 6 , Fig. 31 Only d c d L&s

( O f clypc&s,,T r<~t,?,,Id,,s WL!, a L h l ~ k and r",,,,<Id ..iig'. n c r v found (1'1 6 . l'ig I ) b spcc~cs c11sLiscliim

l i n g ~ , ~ w t r d imatr~rial WAS rial remb le i t n g n m l r tor L1W g,.,,,ir sir i rcogei iablc ILmiigI, l l ie Lest sir,. ,17id i.liiiklirrx, ~ i i l ~ ~ r c l i . size ;md dmsity, the p i r s ~ ,.iiic r n r l ri,oipliology or is lernal slipports and Ill<.

xin,il'lioli>gy or si,,l>,ilniral pnii-, lii t1,e pc,aIs (P I 6, I'lg 5 8 ) T n p h o r m r r ~ y : Cwziplrte &ad (crLr and larger (rag ,,,?,?is .r c*,,...lw a,,m,i;s w,,,, o r t m disrorcrcil iv iLhi i hr wcll oil l l i c rcdin,cid rurince. 1 .s i~ r i;,rirtio.s oiciir i n ,he nai l i re or 11,. rraglnenla w i th ih lpmrrncr o i bn l l i i e l r i - and ihlrapiale Srncliimi& (1'1 G 13g. 5 8) k i n g m e n t c l i s l . r i hu t ion ( F i g 1 1 ) The wide i l i s l i i ~ l,,,i,~ot~, wil l , a p r c s e i i ~ r i h nitnost, a l l cxaminrd s c d i ~ ,,,I.,,, sa,,,l'lrr. ,,,a4 piobnhie iepieseels r i o l n h i i i a ~ t i o i i o f i l i s i i n i t rpet icr disl.ribiilions.

i:diiiiiicyatmis ci ispi ia

( I I i i ' X or ,"la1 rrngtriciil, w~gi, l .s)

01,rrrvc:rl i l i s l r i l r n l b m nml mrde of l i fe : b i m d Ihrieg 91, aninplcs assor in ld wlli deep sesgmass ,,,,~<I>+S ,,i lial<>,~t,,l~r~~,., s%>\,<I<I~ s 7 d s i,, LIX ,O<>.lI\. ,.r,5 siid l",,ll,l~ia h%i,,, or ti," " l irst, area? l,lc~,,tifica,.i<,,,: \+,y small (< I c,"), Ila,,,,c,,,d low 1 c ~ 1 w i l h indm! i i ig i i i l r r r ia l s impni ls aid iprlsls ~ ~ I > , C I , 40 ,,,It rc,ac11 ,,l,c a>,l>il,>s (PI. 7, r i g . 1 ~ 3 ) 'The ".",", o r l r i l is Iiigl,~' l l iai, liir si*igi,i: Lhc periprorl, '"iiiliiirily Iiclwel:,, ,I,,: iiisigi,i *,id inoii l l , r:~';tpl~olrooly: ti^^^ rroi,, roe,l~~rtr~y c I < ~ , ~. ra i , ,p i~ i Lo i l i o s l rigiis or pirilniini,.

i,iih'""""' <k,>r'.s.sliiri

(8 67% or tolnl i i n g n m ~ t i r ~ ~ i g l i ~ ~ ]

Obrarvecl d i a l r i b o r i o n and mode of lift: I'uusd l iv ing very s11i~Ilm.iy lhi i iad in stlly rands or ,lie b*sioai ''En>," rlid " l h L nrrirs". I d ~ n t i l i r n t i o n : TI , " niLlrn?d penL1giilln1 or e l i i j i ~ Lwrl test hrs a r l i n m r l c i i s l i ~ ~ i i a i l i r p o r i l e pore %it..

~ ~ i . " d ,,, a ~i,,~,",,~ groorr ' ~ 1 , ~ petalr dc- v & p d (I'I i, p i g i, g j ;rr asr the in re r i i d s s p p o r t , ~ 'The p c r i p r o i l is close Lo Llic nrargiii. T l w Local lengL11 of the pr tn lo i r l nwa is amre than 213

i ~ i w v u , . n s u c r i eunt dbsrrwd in l l i c r l i i i l y . ~ w s Sih;;:str.ridic ;%re i n n w e io burrow deeply i~, //I,. SI.L!,!~,,,,,, Iltontiiicilion: C lhmr t r r i red by i l w pwsencc of

.I l'riil"lrir,,rs arid late,"-anal ia?c,nir; dreply de- ]"'."61.11 r,oi,,,.,1 nr,,l,iilnrriiiii and rilnkr. ,'?tills i l i s i , o i l l j sl>or,ei ti,nn t h r ru.kcn nsteriu, llaii ' i ' l i ~ lpctala sbui a g m s l llxc prripel.alous i&- <,<>I<. (P I H, l.',~ 7, 8) 'SIX! gc,,"ri< i<J~,~,,ifi~?.li",~ "i ""'S, s'l, ir~sli: i i i l fingniciiir r s s not passii,lr ivrag~ ,,,mi., o r l ~ , , , l # , ~ ~ m i , , ""'6,4).' "C" ":'"""'"d will, ,d,ar~,c,,~,,~l,ic p0.c pmrs <, iLtW i r o r l l d ~,,$I>,ila<,a Z l g ~ *ngg,,,g iii irrcgaI*r < ~ ~ , , I > I P ser,rr ' , I < , , , : I'hr w r y ti,,,, L<,S.r( we.? ,nost i?,~

i i i i l i IY 1,rokrs diiiing sil,,,plr t l k i l i g and Lrs,l ipoil k g 1 i l ( F i g I ) . Frnglnellts Co-

i.+l ly doiii,nnlr ilie ifer;>iv bzril,sl arcas. T h r y ws, w r y rni,>iii"il is, rxtra to vriy i iooriy sorted. wry ~ i , ,~ . r,aliis tiic iiliii~ s,,,I~~ ,,,,ml;,r,, i x , , ~ s

3.3. Muli.ivariat,e stiii.isi.ical analysis

Fig. 27. Dendrogrsnt or a hacrarciiird ~ ~ m d e U1'1;hf~ chster a n a i p m ucxrrrpuisr ~ ~ i i i z i o ~ d WI'DOUL C ~ ~ P C * ~ ' C I S P l?wmenr wei&l!tb. C iu~r$ :z~ rcpaiated at a cosine cuefiirienc of 0 8 (N = 6 1 ) ~

5.4. Conlparisott of echinoid fragments 3.5. Correlation to grain size fractions,

d8lTcrenl meciss.

'l'lie distiihution olerliinnid lrrgmeiils w ~ t b i n sed- i l l ici lLIry raci-3 is shown in 'S*b I I'1ie 'Curxl- gal ' t..iaa co.Lniss I,igl, amounts of <;l?pr~ler rn l+iciiiaiis nieCulri~n, Pnoeuodrris baculasa, and La-

4. Discussion

4 . 1 . Taplroncrnic bias

& v r i r i axa and S ~ I F r e s l r i c l m (> 2 rnm) ~m~poard by th r n~ ia ly l i c ~iirLl,od. ,\ll.lioagh the i i i r~Ll~odol- ogv s d r m from LapI,"llilmi< riiccii*ill.mm (iiiiludmg ~ r w s p o r ~ ) CLS well n\ #,hc d i l i c r r r~ l in l prascrvnlion po- ii.illia1 o i d i i r ~ ~ r r n t ~ l i i n n i i l t a m , advrnlngcs aresem $ 8 , Llic L,,,," ar..:i.&eii siriiiiiiilalio,, 0s r,g,ricsts of t i m e rioioiioiiily pnliIly dislr,bslcd Sauna! i : I r w r ~ ~ t s (NICIiICLSICK 1992)

h i , rxai, ipl? lo r i,spliol,oil,w h,xs is glue,, hy 11,- niliirrn rcl<IS,,rn, wliicl, li<.loags LO ilr mOSL cur,,. ,.ion a,,<$ < i i s t in r l ,,,e,hrl,"rs o f ,lie rp,sm,,:ai cci ,I,, oid fauna i n :he r i u d y area 'S lw rperirs c a i i t r i h u l r

insigiiificaiilly to f i ag~nen fs weighis w i l h ~ i t e saiiipicr reatc~iisl Th is is dur to the S u l t l i n t D i ~ n1em.n is i r p l i o n o n ~ ~ c n l l y very o,~slalrlr with i i a flrr.. >!>I<: tcli niit l very fragile q > i # ~ c s (ro in~mie GRFEN- STl,;lN 1990) 'TI>? d~slr i lml iorx d f r a g r n m l ~ acm%- i l tc S L U , ~ ~ sn:n W . i,onrlliclcss, rlili he foliuwrd wi11~11 l h ~ : study m a as reen $1, i i g . 7 and corw l a l ~ i i Lo the obseiu~:d proence, oSliving spsrirricrsr l u l ia id s,,lslirles Fvrgnrcnt rerngnil ion and qiiari l i l i d i o ~ c w m i n i e r t l ~ taplloson,ic inslabil i ty o r c r l i i ~

4.2. 'Taxon distributions

s , " r n n ol --N.- " '804 3 ? $ 8 C5;8 5 rpiouiynirjna.nl c. i ra 6 2 ra ti a a a a ia a -- a , o n n - - - . -a ,nrP $ c o g a O " 8 a n a a , r o n o s o n *

-PW~-,W yen -?egq - 4 - 9 ,a : : VS$$ 2 5 5 5 2 3 .$gas n - o r n n r ""7 - -

m m

W W ~ - - - W 8 8 8 8 8 8 8 8 6 8 8 5 8 8 8 8 cc8Fj 5s8:: 8 1 0 - 0 0 DOOD " 0 0 0 1 0 0 - n r o g - - o n d 5

%

~ ~ n ~ ~ ~ ~ d ~ ~ ~ ~ ~ a ; ~ 7 X S ? ? C S 8 6 a 1 8 3 5 8 8 8 7 4 8 7 8 8 8 8 $ " - 0 " S Z O Y = = o $ 0 0 0 0 -.,0= 0 0 0 - O E - n * 6 .- " " 8 -

W"," ,tow ","q,, y d z Z Z Z F zsa; El$$$ :sss S S X C , 5 2 3 8 - a X O Z $ - -

-S m ~ s ! n m i m n , o w a ~ % ? X F 3 ? 8 S ? S 8 ? 8 8 8 8 ?;8C 8 8 8 8 8 5

0-r". - n o * * - O n n " o o o C o o - 0 0 " " O B .-

.di,l,mzd*iJ z n s z ,,*, D N " s G 2 g - ra w 2 2 2 : : % i 5 5 5 5 : XC88 Z 2 * n n a n - r n a n a n n n a 22-; R 0-

2 - ~",""!qm'v,x, 8 >,,sq," 2 5 5 2 255; S $ Z Z $ ? $ ? 2 5 5 5 $ - *

ta .- %

'"," l , !uumm~,~-~,q, G g % W $ Z $ $ 5 5 ? $ $ $ 'v!ZZ ? $ $ $ 5 - B i w

f ~ D y , ~ m ~ ~ ~ u m ~ , i q ~ ~ V35: 2 $ $ 5 8 8 8 8 a,-o N m o * ? 3 8 ? 6 3 8 5 m a - e 9 2 m N - n a 2

0

qli,n2asmi..~r,L ir "% 8 3 Z 8 8 R 7 8 G 5 %S 8 '; a 8 gSS$ " " - " - a " - a - - - o n " - 0 - :;d; N n o r X 5 -

Mnm,as,Y+ !, R38C ?:8? 1 3 8 8 8 8 8 8 358 ; !+CS: S $ o O o n " " 0 0 0""- O O O " 0 " O n N I O O - * -

a2

a - 8 - E

O r o j n ~ o g r ~ , ~ p r ~ ~ . o ~ , ~ 1 4 . q 8 BC8c 4 3 ? ? $3873 8 8 8 8 $ " 2 0 3 n n o r * z a g r - - o - n o - a o o o - 2

'I - " * 8 "

r ! , , , d ~ r n r ~ r n ~ , , n v ~ 3 ;; ti 2 $ 5 8 5 8 8 8 8 8 8 8 8 CS86 8 8 8 8 5 2 " 0 0 0 0 0 0 0 " 0 0 0 0 " " N O D " " 0 .- - - 8 8 8 8 3 3 8 g EZj8G 4 , " 2 ; ,,,,

shurr redao~rislr 186, also been iepor icd hy POIL & ILEHNEI ISEGGCV (1966) and FIS l lELSON (1971) in the Rcd sea

" v ~ i i ~ n silldies on the dirtrib,rtioli sod =dimcnt de- l,endance o f n" irregular echinmd i r o m the Red Sea. The i r ru l l t i ienclhed i n them rludies. apcrirlly tho* ioncer,iisg ihr th is species. i1ose1y c",iesp"nd to ,he obvrvcd di~1,ih"ti"" 01 I.ovenia elongala i n Llrc area Therc are no i l rscr~ptmns o f rrhirir*Lerid cl ir lr ibulionr in the Red Sel . She adaptation atid rrf i lr ict ion of schirrrrLerids wi th n deep rooken anterior aiebalncrurn 1- s e n i n

4.3. Distribution o f t a m with respect to environmental parameters

'I'lie d i v e m l y ofregii lnr e d i u n i d r on l i d s u l ~ r l r ~ l c s is greater than ti la1 o i ir iegit lnr e i l i inoid walhin s p c ~ die redirnent type^. This is refleiled i n l h r fact l h r l . 1,iglily posi l ivr c o r i e l a l i ~ n r belwecn rcgltlnr echi- noids are common, whrreby thnsr between irreg+ la, eihinoids are not. Wbea two iireglilnr erhino,dr '10 s l r m g l y correlate posiliuely In one anolhcr, they

r h u ~ v d d s r l l y vsryii ig niorpldog,es, rcecling slrmt,:~ @cr, and burrowing deiilhs. Th is i s exemplified by lbe Clypeas~eroid Urlmodiscus s ir idus (a very sllal- low burrower) and Lhr s~~a taogo id Lovenin elongela (dffpc, t>,lirowers) in ncsr~shore terrigeilous sands. A f,,rther exampic 1s g 'wn by the co-occurrence or the very shallow burrowing Clypeasterord Legansni depressurn and the deeply b u m w i n g schir*~l r r ids in ",,,A*" <nx?<ls

T l w are a niiri ibci or fa i l u i s which ronLro1 the di r t r ih i i l iou or echinoid t sxa including Lernperuture, salii8ily. s i i l n lm ie i h a i a c l e i d c s . water movernenl. predaliaii. nulrienls, behnvior. d e p l l ~ and 'rbanrc" ( K I E R and G R A N T , 1965; EIlEIl'S, 1971; S M l T I l 19841 The hnntiiie of the substiate h;rr been nr-

geed to be the most impur lnnt contioi l ing factor o f 1ocd echinoid disLiibi i l ion ( S M I T I I 1984). Th is

I I O G G I L I ~ R ILOSE, 11984; ROSE 1484; CARTER, 1987, N E I l A U D E A l i 1991) rioe, s variety o f l i n b i ~ I . a s The corrcspoiidei~cr 1hci.ween rc l i inoid frasmen~ distr ih i i l ion arid the ruhr1,inlc is well dorament,ed for the Northern Bay 01 S&g* through their distribu- l i on w i th iesprc l l o bo t lo in b c i e s ( r igs. 19, 29 and

Lo poo,ly roi led, line sands. TI," rcs,,1ir coscemisg

. . d r i i s mels lar iar l iow a 1,ighly poailive corielation to

i o n i s grnin size frnrl ions and coiresponding negn- ~ i v c coirelation t o roean grain rice. 'Slwy nic prei- erenlial ly i m n d i n negatively skewed, madeialely sorted. coarse sands and gravels. The sedimentary pararnetcrs a f the substrate are, howeuer, no t con- sidered to control the presence of these n i e i o b ~ r s o f the epi iaana They rather i i l hab i l environnhents ( m a t red, C O , ~ carpel) w i th orgnnisras shedding large conslitucnt rrdimeutary pni l i r icr (for exnrnple co rd fmgmesk or marsive cidaioid sp lna) i i t lo the s"i,"a,,ding sediinenlr.

'The close c ~ i r c i r t i n n o f cchnoid distrnbil~iori to

Lrs)". 'This is s"ggert.d by the coiepaiison o f rbs* lute fragments weights o i i i h i z ~ i f e r i d s t o rn-an grsin

l n ~ c i p r e l i i i g lib s l r c i i g ~ l i o f hydrodynamic i,,flu~ enre ironl aedirnenloiog,cal parametcis 8s di f l ica l l i n the lh iogmx rcdimesta of the s L d y nrrn as l h ~ y air

diitntcd t o large e x ~ r n ~ by LI,C na~ l i rc tile psi^

i icuInte ron l i i bs l ing roniponrots. Nevcr~heie i r , diC fi:rrncc~ ir, lhydrodynaii8ii energy rnny he in.poiisiL lble for !he vsr ia~ ion o ie ih i i i o id i i ag i ren t piermir iii the c x a i n i n d shnllow r a t e r rediincnls. U~ l i i r i od i r rus auril8ir is found d u n g wit11 L o w n i n elongala i n thr. hn-1 and mod poorly suited nearrhore redimenis i n lhr "protcrled- "Southwed c1iaiincY. Crhinorrwiia rrialhaci is most comn>o,, ,,I shallow well sorled v d , ~ rilenls i n ,nore "exposed s iLn L h m Fib inr ia ovli l i io, which a i m shows n Iughly ~ i g ~ ~ i l i c r n t negative coi rcl,tion ro depth ( ~ d . 5) TI,= reslrici ion o f the iiagnicnLs or there sppcics Lo shaliuw wrler is again sem to re l lc i t lnccors related l o d r p l h and no1 l o d r l > ~ h as n i a r lo r itsell.

i r ~ g r n ~ ~ ~ ~ o f ttlis specm are most rommol l i n silty rands, r h i i l , d o n k a l e in seaginsr meadows. This =gain is seen as being n, apprreni .lrpcndency to sedimentary parailrelers.

The iircglrlnr ecbiuuidr o f the aLudy area arc de- p w i i feederr and highly adapted 1.0 l l ~ ~ d i n i e n t r i n which lhey live. 'They k e d on organic material in- c lud ing~rna l l in f~ i ,nnlo igasisms w i th in the sediment. as deduced by i,,"eslignlio,ls of gu t conlent D E IIIDIIER k LAWR,ENCE, 1382). The reslricLion o f mn.1 irregiilar ec1,inoid tarn lo redimenls w i t h w e - d i c p i n size cl,arnrlcr,rlicr iiich,di,ig rnenn grnin S ~ C , sortmg and skewness (zee Fig*. 31-33, 36 m d a71 ,..? tlnis the dirtiibution o r their r-d

supply. There may be a correlalian between lhe deeper occurrences (espciially i n the norl l iern bnsis d he "East ares") o f CIypensLcr rp. and Ed inocya- rrius crispus wil l , the sengmss I f d o p h i i n rt ipulacen w h ~ c h was observed l o d e p h 01 50 ni (I'II.LEli h I 'E I~VESI . l~ I1 198191.

4.4. Keiationship to facies distribu- t ions 01 i l i e grsrrriizril ldiier dirtrt l ,ul iom as piihlialied in

IILLEII & ?,IANSOilR (1990) 'T l x correlsl ioo of t o t a l e c h i s ~ i d ciuairrs LO lacier r l iows well the d i r unct ion or rshstrairr, dorninaled by irregular erhinoiifr, and I h r d riil~strsicr. dominated by regu- Inr r r h l n o i a ~ ' T h corielrlios, liowever. s i i l k rs irom t i le !ar,, that C/yi,,?arlcr s t > rmgn,entr, r l i i c l l are the ,nos1 rooirnos i n t1,c d u d y area, diow a r e l a t ivc ly undilleren,,atcd distrihl2Li"n. ,rho corrcistion o? regular ochir ro idr dusters is convincing ?or the Esr idans ",efi,iaria duster Lo sediment* associated ",ill, l id aubrlialrs, l?rhi,,o,nelra ,natl,nci Lo near ~ I m e randr I'nonoodaiis haruiusa to deeper soft ~ ~ ~ h s i r n t e r TIic hi$, i l i n t ?iipneusles g r i r ~ i i l n f i a g ~ e,entr relaiireiy raw and lhgl,ter l hnn &lie mnssivc

rnr ler ia l or I ' rmoc idn i in bacuiornmqv explain wl iy srngiass rrens are domisnled by Lhe bernains o! the large crdaroid r i l c best comparison; to i n c i n d i s t r i l i i i l i o n ~ is given hy llir clurLers of ilnrrgulnr ct:I~in&ls w i t l t o u t C I . ~ , w ~ l c r ap. w i t l i &he broad d i s ~ d m L i o n O! ~ t b u l a r i n ovuliirn r lusler LO coarse i l ln l inw water sedin~cnlr; 1he res t i i i l rd presenct of the Ed,inodrscrt.s aurrlur r l i a te r Lo thc "Southwrsl d m m d " ; 111~ d m r h i t ! o n o! s d i z ~ ~ l ~ r i d ~ in h - i n s ,:<,,,Lain,,,& dreper ,,,ads and 'iiiiddy sands; LIW prcc eiice o r i l x r ingliierr& depiessiim c lm le r iii roar*, s ,~dmest r ill inrigiiinl ~ ~ ~ n p l e 8 of tliese b e a i m and the n ~ ~ r l i t , o i , or LI,C ~ ~ ~ ~ i ~ ~ ~ ~ ~ ~ m u s crizgus to the

inoilhero ba~ii i OF l l w "ENI. area". s i t e p ~ s r i l i i l i l y !or l i i recl mmpari*os o r d i r l r i b " ~

eon r,lc~ rnrirs is i,, 'PAIIS. 3 and 4. shire is a more distinct predominance oi siiiglc echinoid tax* wi l l l i n 1111, ind iv idual bo l lo rn incies t i inn wi th in l h c ~ ~ d i r n ~ n t ~ r y inc~rs. T lh rsrees i n l l ie expected lo in1

~ I m ~ w t ~ w c r o i rcg,dac c c l ~ ~ t ~ ~ ~ i d ~ (rs1~cr~;311y F:txxJarx liirliilaiinJ ,n the c o d i r r p r i arid c o r d reel l h o ~ ~ o n i racics '7W c"rirsc>onding roin1gn1 srd,,,,ei,lary Pa i,cs ioiilriiia in, iower nvcrrgr pcicenteg<: or regii~ai c i l i~ i io ,d liaginrnir wi th s r r la l ivc ly large pciceotagc or r,y$""Crr sp and i,ngnn,,m i irpmr, ,n, . Schirn- d r r i d rragmcnis to la l ly dns,innlc t l i r m i d bot tom fa& i,, 11,e in l id fa& l,owever, they a,, car 1,s comnm. The ionl r ib i r l ion or Clypenslci

sp. iiagmenls d i fe r r highly in ~ h ? divcrsc Irotlorn racier, lo fhc scdimenlary fnc~cs, howevei. thcy rhow

firid olmrrvslioss. A n ir i ipoil i lnL role iwty also be pl6yrd by l h r fact tha t i n m y or the varisbles (15 iii a l l ) ured in l h c rlsssif icalios o issrnplrs i l i lo redh~ ,,,re,n,y iar i rs maj 1,nve l i l t k n l r r l l o n ccl,ino,d pres~

inride tile r ~ i s t r i ~ ~ u t l o n uiui~ler ihiogtoiii

n iggrs l t1,at cc l~ lno id pieseiiie r n d dis l r ibul ioss cri, lhc: xired i r nil iaddilionnl Loo1 l o sedimcstary facira iii lhl.!l>iwg l o deIiiirsLe Llic snalsirr and ~nL.:sl o i s u b

sedin,enirlion (1211.1.ER k MANSOl iR , 1990) I'L,,r, alll,oii61i inpl,o,ioi,iii proirsres may further aliecL thc ,,a,ure o r ftrginpiiL rrprcsenlaiion, the

higlily d i l l ~ r e n t d c d dislr ibulion orechinoid remairis o f this shnlimu water l r o p i i r l envi ionmrnl hhr agood c1,ance o i being prrserved and iecogi,ized.

The r w or echinoid i iagmcnls i n psleontological rwcsrrl, is, 01 coarse, ""lliing n c w Ti t is ir e r ~ l,eoally lrar for iegulni er17inoids having porn preservation potential ( K l E R . 1977A). have oG LC,, bren described and identified on tile basis or i r a g m m k d o n < (SCC TAUI3E lL 1951; K IER, l877B; ZARD1N1, 1988) R,,giilar eihinold iragn8entr have

L*,i,qm,,u ".spur rbriiow pni bvmvin

~~,b"in,,no""ium rhiikaw modrnc. h u m -

.nb le 6 Covnpiiution of mode of lice. grain size and lacm restrictions o f most iiripoilaiit k x a w i l l h the area.

ters such as coloration patterns, w e d t o help d i s t~nguish fragments, w i l l generally iuot bc present in roi-iis. ~ i f f i ~ ~ ~ t i ~ ~ in dis~inguisi,inp, fragments sympa~r ic ~pcc ies w i l l diminish the resolution of the method. The lack o f compaiablc compiele fossil sp*ci,ncns may hamper fmgmenl ldenlifiraLions as

wcll N diminish the pos ib i l i t i es for h n c l i o n r l o m r - plioIogiini nnnlyris o f t a t characters. N o n e l l ~ ~ l e ~ s . ule qilanti~fation edlinuid rragmei,ts in sieved non~roi,solidnted scdiments as well as on lnrcer ex^

Echinoidr can he q u a n l d i ~ d "ring fingineel ~ ~ i g l h t ~ fromi s lmdnidixed sediment sample= The data can hc subiequrnlly analyzed using ir,ultivsrinle s ln l i s i i cd t c c h n i g u ~ . T l i c advan- tages u i this rriellmd lie i s the k t that tapli* ,,o,nira!ly ,,ssLable elciiienls, s,,ch as ,nust reg- "tar ccliinoids, can b e recognized m d included i n the nnalysie. al l l ioi igl i they are ra r r l y p r n e r t l as complete well preserved skeletons.

,pd ~ u r f m s o f consolidated rocks rhould l e d 2. 'rile dlsiribii~ion 01 riagiilents is simiiar t o the to useful resuits i n thc reconstructioc~ of e ih inoid distribution nr observed i n the studs d i~ l i ,b " t i " "~ . area and as dncr ibed iii puhl8shed rccountr for

Ited Ser and Indo~U'csL I'sciEc echinoids. 'L'h*

dirliihation of i iv i l ig rc l~ inoids is the most i m ~ lportani s i t o r i s , deteiieini isgiragreenl d i s l r i l x - l ion, allhoizgli the dii ieienlinl tnphosomir pa l l i -

- ~- ~~ ~~

b'"iii size:, sorting and skrriirsr or thr rcd,,ilrliir ,,, u,~i,i~, tliey nirsr, .~. l ,~~~ mv,io ,,,,, co,,. (XlU'l~X 1% 13 . lY87 PaIc~ogcn~~ edwmds dastriL

tiids iii rcliiiioid dirfi ibii i ioii air nisi? revenid lhiilioi~s in Lhr Ailnritii arid c i i l i C o ~ s l a i Plains.

lw tlie close mirr l i~cioi i LO boitoiii aid srdirxier,, IPaInios, 2 3904Ul. 'rblra

6. Arknowledgcruents

'slim bL,,<l~ ",a> s,>pp<,r,"<l 1,y Lhe #,.<>J<><,b 1, 5877 a i d 1'7!iU'lGC;cu of the "l..osiir riir Fuidr rwg i l f r W Z ~ S ~ ~ P ~ S C ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ C ~ ~ ~ I :O.SCI~,~~~~ OS~.,.~~C~CI," ( , , , O J V ~ ~

leader IFiilr I' Slei,,ingur) I would Ilk,, to

i lwiik F i i l r F. Slci~iisgpr, wc~ernci P~ l l r r . 1 t e i r h d i:olchi"wrki. Peler Pervcsicr, Karl Kl.:ro,a,in, I'c. LC, Ilworscbnk (all U,,ivrrs,ly or Viennn), Cl,iial,r,, Ilupp (Geologirrlie BiindesaiisLall. Virnnr) and Ab- bar hlanroiii (l lnivrrsily or Qri,a/Arrii i~. IEgypl) iiii their licld suppoi,.; Eiii l, Krilscbei ror l,rip a1

1 1 9 - " N a l u i l m ~ o ~ ~ ~ ~ h r s hluseiim" in ~iesvtn. t'rtvr f Steiwngcr, LWrner I'iller and Joiiaon llol,eeeg~ 6,:' and Momkr Nrbrlsick for riilic&lly r~ndilib ih,. iiiriiiisciipt; woirgang iteiciiiiiariii fur tlir l,~io L"gia~'hs

IX IBEElt. 11, 1990 LIisLribiilion pnltcrns o f i e g ~ dar 8"a ".dins (Echinodciioaln Echiaoidca) across the Speiinoodc Shelf, SW Suiawe\i (In- donesia). - [I,?.] <:. Ilk: II1OI)ER. X. DlJBOIS, X.. LAliAYl: & M JANCOCX (&It) . Eclrin~ uderzii I t n rn i i l i , 185 189 (Balkrmn), Ro t i c r~ dam

. . 32357-67, Allieri

1XI.l.ON. \V I t . k 0Ol.I>STEIN. hl . 1981 Mlul~i-

l l i ! l l : N \ N , I I , 9 8 'she id,,gy uaiiale siinlyrir, r~ielhods avid njipliialiorir. 587 of F:cl8#8~<>car<Ji#>t~, rorLatt~,r# (EcI~t~t<><lerr~~ala. 1, (John W l q and Smm). New York.

Si~alnngoidn) iio~,i di!icre,i~ lhal,italr~ ~~ J I S I , ~

biol Ass. \ I . I<.. 48 !I7 I14 lIOl>L>, .I I 1 k SI'AN'ION. I t J . l!l90. Paleortol~ OCY: cosiri,tr mid nviiliiniions. 502 no.. (2nd

NEIIF;I,SICK, J . I1 .The N d h m Bay o f Saraga

~ ( j~1 , t ' uS . K. k 1ti)M.AN. I t . I981 Lei i:iliieldes ,Ie la Mei Rouge, hloengispliir ;ioologiiiuc: rl paliontoiogique. hlin i!nircrr. C:oio. 'I>;ii II~SL. scieiit hlerii sri s ~ , F ~ ~ ~ ~ ~ , %I ~115. Plr ir .

IIONOVAN, S. ti., 199 I . 'Shc Lnplioiiuisy deihinu- denns: calcsreoiis m~iltielerinen~ rkderuns i n the marine enuironinenl. -- [I.] IlONOVAN, S K (cd t ) The Processes d lomil,zalmn 241-269 (lJelhnves Press). London

DOTAN, A . 19901J. I'opiiIrLios rtriict,in. .of

the echinoid llelrro~cn,ml"s mmni i i ia i i r i (L.) dong ihc littoral zone or soii11,Mart Sinn, Coral rreir. 9-75-80. Hcr i is - I I~ ide lh~rg

DOWN. A k I'ISIICLSON. L , 1985 M o r p l d ogy or spines of Ileleruceiilrutlrs ma,nn,dlnl,rs (EchinodrrmnLn, Erh~noidae) and iLs ciulogi~ cai rign,(icnncr. [I,, ] 14 B Kregns k 0 . 1%. U. S. (Edl.), Ecliisudcm~ata, 253 260 (A A. Bnlkeriia), ltoltcrdani

DURllAhl, J. W., ?ELL, I 1 H., FISCIIER, A. G . KIER, F. M . MELVILLE, R V., PAWSON. 1). L. & WAGNER. C. D.. 1968. Echinoids. [I":] R. C. MOOltE (IZdl.). 'lienLire or liivrrle- brate Pdeonlology, part S 211-840 (University "1 tiansar I'rrsr and the Grologiral society or Ka"s&s), Lnwrmce, Kansar.

DWOHSCIiAK, P. C. k PER\,ESLER, P., 1988. Burrows of Caiiisna?sa bouvieri NOBl1.l 1904 from Snhga (Egypt. Red Sea) with romc Re- marks on lhe Biology of the Specis. - Sen& mbergiana maiil.. 'Ilk-17, Ftankluit/Mnin~

EBERS: 'I.. A . 1971. A prelimisar) qiianlil ir live survey of 1he echinuid ram,"& of Kcnlakekun and Ilonaunsu Bays, llawaii. P a c i f i c Science. 25:112-131, Honolulu.

EHNSL', G., 19 i0 I~'aasergchundcnl~el1 usd o ~ ~ ~ ~ ~ ~ I , ~ I ~ ~ ~ ~ DFI lrregiiiiren ~~h i , , i d~ , , dl:i iioidwwtdeutrchco Oberkreide. - - Pn lMn~ . Zeitsclir., 44:41 62. SllrllgaiL.

I ' t l l l IE l t , 1 , l97G R i e r ~ ~ o n n i inoriil#ulugy u f l h v r nia eioagnta (Gray) (Erliiiioiden: Spslnngoida)

Thalas. J~igoslav . lZ:l2l-128.

FEKBEII. I . k 1.AWHICNCI:. J. M., 1916. 1)~s- tribulion, rirbstrntlim prererence arid hurrow- i,,g hehavinr or i.ovm,a eio.gaia (Gray) (1Zdii". oderrnala: Spilangoida) i s the Gulf or Clr l (Aqdn) . Red Sea J exp mar. Biol Ecol , 22 207-225. :\,ztrteidaio

GlllOl.l>, J., 1982n. Fui,cl,~r>al ,norphol "~ of the Clypeasceroid ecl~inoid Ed?inocyamur puril- i ,us. -- [I.:] J . hl. ! . ~ W c v c e (Ed,..). Echin* dcrmr: Proceedings 01 ll ie lnternnlionai Conler- m c e . Tarnpa Bay. (Balkemn). KoLlWdnm.

GlIIOLIj , J., 1982h. OhscivnLions on the Clypeas~ 1eroid ljrh,nocyamus pus,iius (0 . F. Miiller). -- 1. Exp. Mar. Bio. Eiol.. 8157-74, Amsterdam.

GIBES, P. E.. CLAItK, A. M & CLARK, C . M.. 1976. Ec1,inodermr lrom narll~ern region of the Great Barrier Itref. --- Riill. B i . Mur. nalur. l l is l . (Zool.), 30:101 114. London.