MAY ONLY

ASPECTSOf THE BIOLOG Y AND LOCAl. DISTRIBUTION OF SEA STARSIN H.A81T1NG A SLOPED,IlOCKYBQTI ON.

IN LOGYBAY,NEWFOUNDLAND

CENTRE FOR NEWFOUNDLAND STUDIES

TOTAL OF 10 PAGES ONLY MAY BE XEROXED

(Without Author'.Permission)

IAN PAUL EMER50N

C,I

'A'SP~cTS

_.

O

,

F

THE-BIOLOGY ANDLOCALDiSTRIBUTION OFSEASTAAs INHABI TING-A"SLOPED~ROCKYBOTTOM

"IN'LOGY BAY; NEWFOUNDLA ND

Ajhest s

submitt.ed

in partial fulfil ment :,'

..St.~ohn's

,MarinesC1 enC~5'''R~s ear~hLaboratory'

.::.;-:" Oepartmen~Of.:Biology'

' ~~r1 ~1

^{Univer s ityof:}

Newt'ou~dtand "

Newfoundlanc!"

...

: '

^....

.','

.-:-·r , .~.-.

'.Frontispiece,."Atema1 e'.'leptasterias"pol aris'(Mull er'amiIrcschel,

·1842)in'a,broodingpo~ition.

.',

(

," "

. .

;'~c: '.,:::" ;. "

,o. Abstract

.?- .

'Th~^spath-land temporal,di st ri but i on~f^five species'of astero id swereexaminedoverIIoneyea r sampl ingpr ogr allrnefbymeans,of a

s~ries

^of

Ph~togra Ph~

^taken

~~eklY alo'~g

^a

~et

^{tra ns ectline}

extendtnc perpendt cu'lerlydcenesloped's .ublittoralrockybott~m:in

.

^.

Logy...Bay .Newfoundla nd.,Estimate sof ' the abundance and di strlbiJtion of three spectes ctasteroidsinhabitingIImor e'gent ly sloped subli ito,:al

p1atelw'~~u"ti~_9 ~.hf~~ ~nderwater

cli ff wei-e

al~o 'dete-r~1'ned

u.singavari ety ofquant~~ati v~, 'a~d qUalit~~J ve Samplln9'·techn fq~es·. :, The~es~1ts^ofsyste~ti.~.^{st udtes}

(If

se l ect ed phystcaj,

. . .

^,

chemical .anCl' biologi cal chej -ect erf st .tcsof these habitatsarepresentee. __Such

diaract~ri stics ,

^{includedepth. stope,relative}ir.radiance;'waterl, .

temperature.dtssclvedoxygenconcentrati on,cond~ctiv.1ty, and predato r end.prey abundanceand distribution.

Spect f tc studi es relatingtothefnte.ract f6nofsea stars wtth theirenvironment(t.e•.feeding,light prefl!rence,·pul'lcepectty, and

'fiit,:rs~ecific ~nd ,i ntra,spe~iffc-

behaviour) ,

un.de~taken·

^{to'-elucidate}

rectors affect.ing sea star djstrlbution.are also,presented.

A presentettcnof the effects of otherspecl f ic.biological acti ,vitles

o~\ l'~cal

di'stribution pat t erns..sucf

a~ migrat~~nand

mOvement.,rep~oductionand.growt h is al~ogi ven.

In the generaldiscussion, a hypothetitalmodel'forconsider i ng

.interactio~s

^{between a}

sP~cies

^and''its

e~vironmerit ~at a~fect

^population

dynam~Cs^and}ocaldi.st~ibutionis presented., Tre model ts·then. uti1i~ed·.

ina~ analysi~oflocatl ycbser vedasteroid distributionpatternsinthe' studyar:ea. From thisanalysisit,iSco'ncl udedthatthe-distribution

.'-,,

.'

" - , ) " '.

\

.. '

.

or:occtti~nce

^of

wat~l'

d1sturbance.prey.predato".and competitors

~rt ·

^,

'. ih~ ~or

^factor's

a~f.ect1ng 'th~"

abundance-di stri button'

'~tterns

^'otthe

."seast ars-tn"thestudyarea;,Ast derranyeof-interactions.beween"a

given-

sp~c~~s an~

^its

'e~v1ro"nme'rt

^may

~

^of

s~COndarY:JlI'IpOrtanc~,·tn·

^..

",.this-n!1at~onSh1P . ^{- '. "}

,

.

"

- ".'

:

..

,",

i;.'

"J'

I;" 0,

W _{" '::}

.

:, "

- :

"

j

.; "

:."

.'..

, '

·.

.

".•••oneis Impressedil0t."wi t hthe knowl edgegained.but 'rllt h;r w1th thenumerousquestions'stii','to bea.nswe~e~."

. . .

"Feder-and.chet stensen-

'Aspect sof AS,ter o i d Biology' . in,"PhysiologyO(EChlnoderma~ '~_•

.edited

?y

R.A."Sool oot 1a n.1966.

. ' I.

TABLE OFCONTENTS

lrtstoffigure s-;.- - ..;.---.:.--~-,--~.:.-~-^..---~---^{..---- -.}

L1~t

^{of Tabl es}

---~.---.:.-~----:_--~~-~.~--

^••

---,--- -~.-~----,

A~k~?Wledgernents---:;.----.-:.••-- ...-- ....-.,-•• ----.-- ..:-- --- Gener alIntrod uc t ion::.~--.~---- ~-- --~--:_. -~-~--- _:~.--,.:.--~~- -­

A. StudyAreasees ters• SystematiC::s,DesCl"i ptio,~,dnd GeographicDistr ibut i o n--- -- -.:.~-.--:.-:---...-.- - -_. -

~. 11 vi

64 21

•D.

. :; ::~::o~:·:,:;:;:~;;:~-::::::::::; ::::::::::::~:: :

~jgra t1o nand HovementStudies---,.__~__,":---.- - - - ~ --- - ..;:..- 119 E. reedin g St udies-- ----..--- ---.--..:---.~--- --'7-.-

•.----_:

¹³⁰

F. Reprcduct fve_{~i olcigy.}----:- -.---,.-""~...

-i"---.

.:..-~----. 171 G. Growth'of'J uvenileAs t erias vul gar is h _ 190 H. Pull-capect ty-Experiments ";..-- --- ---- -- ""--.----~---- .:.- 197 .

1. In'trasp eC1f1C'andfl'ite'rspe cf f icBehaviou r al Reac t i ons'--- 21B J. labora to ry.Ligh~.p~ferenceExper iment s---:'.--- - - ..:---. 235 K. Substr ate ;. ..._~... ...._.__._.. •• _.~.___{.~_} 2444-

! . ,

GeneralOiSCUS Si Or--.- - -- - --- -":"-:--- - ..~---.-:.---. ^248.

.Refe r encesCi t ed:-- ---- -,.- - - - -- -- - - ---.:--.--.--:--- - -- ...--- 285

- I .

Appendi ce s'. -- - - -- - - •••_--.':"--~"-~~~--~-~---~---:---: ²⁹¹

'·.1-

."t...•

Figur e1;

·

~

^Figure

~.

LIST OFFIGU~ES.

Phot ograph of the,abor alsu~(.~c e

,

of Ast er fas vulgarisuu--~-:uu'--~---:~ir··"'----·~.~_':-·-,··

Photo graph"oftheabora l·surfa~eof Leptas't erias pol aris - ...--- -----.;.>-- ~---;..- .;.:..- -.:.~:_.~ - -•••-.-7'---...'10, Figure3.

Figure 4.,

,

~~~~~~r~~~_~:_~~:_~~_~~~~~~ ~~:~~:.~:;:~~~~::~--.---

Photogr aph of the abora l sur fac eof Cross as t er

~

-_ .. _ ,-_._

...,..- -- ~_._~ .-...~~ -~~ . -_.

13"

·" r ^{5: . :~~~~1~~~~} ~~.~:~:'"~~~~~~.~~~:~::.~:.;.~:~:~:~~ -

15

I ^{~, gure}

^{6. Photograph0; theebcrel}surf ace ofHenr 1c1a sp,,- . Figure7.

~~~~t~~~~~~_~:_~~:~~~~~~~~ ~~~~~::_~:.::~:~~:::~~~u '

· Figur e 8. Drawi ng o.flogy 8ayshowingthe locat i on ofthe study area

r: : ' :....

-~.-

...

--.-~.- - -- .; -

..

-~.--?

·.Figure~ . Diagra m ofclinomet er-:..-.-~-•.---.~--- ~ --- -.~­

Figure10•.Diagranmatfc representati on of the stepwts epro- cedurefordetermining 'theslo peofan irregular bott om(TLQ3)--- --- -- - --- ~- ----~- .Fi gure 11.

i~~:t~.4~W~~gT~~' ;.ht:~~:~_~:~~:-~~_~~~~~~:~:~----

^.

Figure12. Annual\emper a t ure regime of the st udy areaat sixdepths-------- --- --~ --- --- -,----..-- ..;~.""••--.- ~.

Fi gure13.Annual d1sso1vedoxygen cqncentratt cn regimeof

.the studyareaat sixdepths .----:_--·-:··--··----..•

•Figure 14;Graph depfctingthemaximum vertical s trattetce- ttonof temperature .relativetrredtence; dissolved

~;~~e~·:~~::~:~~:~~~:-~~~·:~~~~::~~~:~·~~-1~:~~y

vi Ffgure 15. Map of logy Bay showing depth contours (inmeter s) Fi gure16. Underwater'phot ograph of: the photographictower.

.and divers)taken duringaphotographicser -ies-- .- -H-

•.,..{I'

--, -'- '

17 19, 20

24 30

31·

33 40 43

.45.

41 68·

Figure,17. Uliden'l'at er photographof the phctcqr-epht ctower showi ng theposHi onsof~amera'and h,ousi ng -.---- 69

79.

82

83

83

, -

85 Figure.lB.

~~~:~~t~heP~~~~~~;~~~~~gP~~jO~l~~~lio~~~s

^{:. .71'}

Representa'U~e

^pho'tograp h of a typi cal TlQ taken atTlQ-8.duri ng 11photograp hi cseries .Oct.,5, . 19JO. ~_.;:o_- --. - :_-:"- -~ ---- --;,.--~ ---.~--- -~-- -

- .

72 Graph ofthe relat iveetendence(ba;~'edon yearly ,

~~r~a~~~~d~~~~~~i~fe~~h~~~~~i~

^a:nao

~~~~~~s~:'~erf

^as'

~.for.each~--- ---======---

F1gure 21;- Graph'of absol utea'bundances(y~a!'lYl11ea~'Of ^.

e~~~1~~~ ~~~~~~~ht~~lc) ~e~~ds~~~~::si~~apa~p~~~~

~respecttortc's-- --- --- -- -- - Figur e

2~A.

_\^{'Gr aph}_{ast eria s}

~f the

_polaris

abundanc~'

_withrespect^(yea;ly_{-to dept h}^{mean) of}_••

~-

W-_":'_"-h Ffgure 22B:'Graphof the

ab~~dance

^{(yearlymean) ofAst e r ias}

vul gari s withre spect todepth

---:.--.:::::::::=--

F-i9.\lre22C.Graphof'theabundance(yearlyme~n)ofHen~ic1a

, eschrichti wf th respect to angle of inclination;

1.E!.slope(e )-~---. -- ~ --••--- --- ---.;.-

~fgur.e20.

Fi gure19.

~igure

^23.

~~a~~u~~TCQ~~~~~sa~~~~~~::_~~_~~~~~:~:~~~:_~~~~~i

^591.:

Figure24.

~~a~~u~~TCQ~~~~~Sa~~~~~~::_~:_~~~:~~~~_~~~~~~~~.: 9 2

Fi gure25'. Graphof temporalabundance ,ofIlenriciaesch~ichti in fourTlQblocks -- - ---.:. - - - --- - - - - --'-, 93 E1gure 26. Mo~elpr esent ing theinterrelationshipsof pro-

cessesregulatingpopulationdensities --- --- 109 F1gure27. Und!r;Wat'erp~otagraph tak~ninthe MALzone

showing pOsitionalrelatfonshipsof Aster1as

.. ~a~n~~ ::.~:~~~~~:~~~~-~~~~~~~:-~~~-~~~~~ ~- -:..-.

¹¹⁵

Figure 28. Plo_tof movementsexhi bi ted hya single spec1men

- .~ ,(11)of,Solasterendeca---.~--~~.---."': 123

-11~-

"-".

i ,

. ,

'~.

Figure 29. Plotof movementsexh i bitedbya'singlespecfnen ('2)ofSOla~tereodeca--- ---:- - - --~ --- 124 Figure30. Three dimensionaldrawin~of the Cliff area

defining_. predat i on areas.\ ^. --. --- --

.

- - - -- -- -- -- 135

"Fi gure 31. Relat ive numbers of feedingand non-feeding

individuals. andpe r c ent age.ofthe populetton , .

::~~~~ga~:a;eii~d;:~~~~ L~~~r~:yi~.::~:.~~~_~~::__

¹³⁸

, ."~

^{Fiture 32•.}Rela t fve nurnbersof'f eeding andnon- f eed i ng

- individuals-_~ndpercentage of thepopUlation _ .

:~:~~n~t~~l:~tf~1t~g;u~i;r~.~-~~-:~:.:~~:-~~:~~:~~~

¹⁴¹

Figure33. Rela t ivenumber s 'of feedingari~non-feeding

, ~~~~~~:U~}Sc~~~s~~~;~n~:~~o~~/~~ ~~~Ui~~~O~r'e_ .~

dationarea~1nlogy Bay---"--- ~--7:---- 145 Figure34. Colo urphotogr:ap"1lofHenricheschrichtiposit"1oned

On"the encrus ti ng tunicateDldemnumalbidum--- - - - 158

184. 174 Out1fne drawings"of thebrooding positi on"s

t~~~i:~ ~~~lQP4~f~~~o~a~~pnsi~rj~lyP21~r1971

. . I ·'

Drawingof thebrooding position of,Henrich eschr1cllt i ---~--- ----"---~~--.-"--.--.-~- 178 F,igure35.

Graphof temporalabundanceof Asterias vulgaris withrespect tofou rTlQb1ocks~pawn1ng pertodindicated---,-- ---:-- ~- --.:.- -~ 186

""Fi gure 39-. Grap hof temporal ~bufldanceof Henri ch eschetcht.t'

w1th res pectto fourTlQblock$.-:-wrtJil)r~

and,s pawni n9 perio ds,indi ca t ed-----.---.~-- 187 Figure40. Drawingof harnessusedforPull Capaci ty exper i-

ments-.---;.---;.--:--- ---.--- - -~ 199 Figure 41. Dra~90f·theexperimenta l,apparatusused in

studi esofPullCapacityof~eseveral speci es of asteroid s---;.---~ --- --- - 20 1

'I ;;

-iv-:

Figure 42.

. j .'.

of iepta sterJaspO~ar1 S.Asterias

icT~ :~~h~i*ITT~ral0~~~~~~~)papposus.

ngtePOS1tions ofthetubefeet . ' utili zedinsubst rateattadvnent:--- - - •.;---- ---, 215 ,fi gur e,43. I!ypot hetical model present i ng" the inte rre l ation-

shi psoffactorsandprocesses regul at i ng 'the...

.distr 1qutio nan~ ~bundanceOfseastars-:--- - --:- 250 -•

,p :

q .

:- ..' . " / 0 "

f > . .

',., ', ,

.;'I,'

.>.

",

.

~...-

34

.LISTOFTABLES

"/1 _., "..:i">/';J

~.:Tab/~ 1. -"~~-SJ.oP<d~ta

^fo'r" " TlQ

---~.,-.,--~-.;--.-~-~~

",:Tabl e· 2.

~~~~~s~~ci~~m~:~~rn9~8fe~~~~.,:~~_~~~_~~~_:~~~~_~.: .

⁵¹

..

~ ~'-- '":'. ', ^{" '- "}

. '

'<;..,·-fab1~3....fl.b.unda~ce_s_per TLQofselec t ed organismsfQund

"'\.:.,-'~.-~~~ai~~i~l~:~_~~~~_~~_~~~~~~~~~_~~_~~~:~~~~~~~:~:.^__~

~' .~

63 Table 4.

Jable, 5,

Numbersof-individuals -of the_ten~~ostabuplilant fauna(excludingasteroids )per 0.25square meter ';

as collectedbythehandsampli ng technique,-~~ ~---~. 58

t , . .

ok'!rl~tions.relative abundencqs, andhabitats of"the- ~'uscsobser vedin the.Cl if f Area-- ~¥.;';.---

90

95

"\'I .

...': ;_ .Table"'6; Analyses ofvarianceof the'yearly di s trlbutions'

.-.-r--

~:n~in:s:~~~~lc~~la~~~ 'J~Qtbl~~k;Vl~g~r}~i2~n:fl~

.~--- -- - ---_._._--- ---:- --,--_

. . ^-

.

~ .

^..

' -l~ble

^7.

;~~em~~~~~t,\g~~~i~~~~an~~s o~i~~J~~f~~~l~a~i~al ~

^"

. " "

·;~~:~rs~~~~_rT~~ ~~~~~~u~~~ ~A)t~~~ ~:l~e{~;s::~~~~

^__

~.--.~ ,

·Tab] e B. tnterpr etetfcnofosci ll at ion characteristics as' ,defi nedfor specif ic'per i ods of'the yearbythe

abundancesofLeptasterfas polarisin TlQblbck

,~- 1:-6--- ---.,_- ---~,--.-'--:'- ---- ---.¥---

..

-v-v- 97.-

)able~;..preta"ti on

^of'osci fl a t i on,char acteri'st1cs as 'def i ned forspecificpertcds of the yearbyt,he abundances ofLeptast er iaspolari s inTLQblock 7¥1~---:--- --- ---- ----. ---- -- --:-- -:---- - - ---.- - - - 98 Table"10.Timeperiodsof oscillationslist i ngthemaxi mal

andmini mal abundances(no.of ind1\lidualsper . "

.

~~~~~~i~e;~~ ~[Qu~i~~~U~:~2b~:~~~-~~~~~::~~~~~~~~:¥

⁹⁹

Table'11.Interp_rebtion

~f.

osci ll aii on'character-tsttcs

a~ ' ',..

defined for specificper iodsof theyearby the abundapcesof.~vulgar is in,TLQ blocok-7-12:- 101

.'.~~

l ,

Table 12.Timeperiods,of.cscfTlatt ons'li sti ngih~ ma~tmal^.

" andominimalabundances(no.ofindi vi duah per

. .squar e.net er cfuncbscuredbottom)of Henrich .

GZ

.eschri c~t1:for TL~block7.,12,.- - .~-;••--~--•.--.,•"" 102"

':'vi-

Tabl~22.

r

Table14. Sunrnary of analy se sof mean abundance s ofHem-tete eschricht f for thr ee period s ofthe year (consl der ed by palrs L usi nga t-testfor unpaired means(Wang .progr anme)for TlQblocks7-12~A)and19-24(B)-- - ·10S T4ble15. Time per i odsofosci llati ons lis ti ngthemaxi mal

and min imal abundances (no.oftndt vtdua'lsper squar -emet erof unobsc:uredbottom)ofHem-tete es chri cht1forTlQ block19- 24 --- ----::::::':":"- - ---- 106 Tabl e16. Interp r eta ion_{defin ed}for specific periof cs ct'lfattcns charac te ris t i cs asods of the yeer.bythe ..

i

..

l~~~~a~ :~~_~:_~:~~~:~~ _~~:~~~:~~ ~_~~_~:~_~~~:~

_ _ : '

¹⁰⁷

Table 17. Out li ne. ofthesampl in gme t hods utilized to deter-, mine the distributionandabundanceof asteroidS:

found1ntheMAL'zone--- - - -- ---- -..-~---~-- ---- .;- 111 Taq1e'18::

-~~~1:~~~sw~~hL~~~~~~r~~sMe~lj~~Se~~1f~sf~r,i~;

.~ --~---~.--

^{..--- --- . 113}

,

Table19. Positionsof Leptasteria spolaris'and Aster1u. vUlgar1~with respect to exposure1,ntJii"lilJr'Zone -- 114 Table20. Relativeandabsoluteabundanceestimates of•

Ast er 1asvulgal"fs andteptastel"1a spola r is'1n the

~..---- - --- -,,:"- - -- --- -- :-- - -- ---- --- -- --- - 118 Table2-1.

Sp~culatfon 0;

^the

,ma;~r

^activltfes

di~play~d

^bytwo

spect nens of~endecaduringmovement studies' .126 Sunrnary of thed;~iledfeeding st udyof teptastel"ihs polarislist i ng preyspecies and the,percentage eec preyspeci esrepresentedofthetotal preyfor each predat ionarea--- - -- - - --'-- ----:-- - --- --- .,.-;'- __ 139 Tab1e,'23•.SUlll1lary of the detailedfeedin g"study ofAst er 1as

~~~~a~:;yl~~~~nsP~:~r;~;~t~ ~fdt~~e~i~ie~;:;~lfor -_

eachpredat i onarea --- ----,;..-- .,.- -.-- -',;,• .:._- --.---..- ·,1 43

. ,

'

,.Table 24. .5U1111lary'of,t he det ai l edfeedt nqst udy,of Crossas t er "

~:C~o~~~:f

¹

~~;~f~sP~:~r~~:~i:~ ~f<lJ~et~~~~:e~;:~e

^for

ea.chpredet tcnarea--;---·- --·~~---..

- - 7;:----

^{146 \r}

-v1i- • .

Table26.

Table25.

~

~~~tl~~o~~~~r;fo~:;;~:tj~~~S_~~~~~~~_:~~_:~:~~._

^{••' 153}

List ofpr ey ofAste~ii:lsvul gari sfromfieldan'd' laborato ry cbservat t cns --.. ., -: :.______ 154 Table27. Listof prey ofCrossa st er~fromf 1eld

and laboratoryobser vat ions __n n _ n n n n _ _ _ 155

180 176 156

181· Table 28. List of preylaboratory obser vationsofSolas te r endeca

_====-

fromfi^n.eld and^{_ n • n}

.

'

.

Table2.~9..:'~N~u~m~b:eesand percenta gesof brooding Lept aster ias

~perpredat i on area--- - --- --- --. --- -~

Table 30. Surrma ry of the reproduct iv ebi ologyof thestudy area seestersv- c- ..-·..·---- -- -- --·- -- - ---·---- - ---

. ~

Table31. Ass essment,of possi..blefactors affect in gt~e brooding'lo.cation~"·o.fleptasterfa spolaris__-r--rr-»

Table32. Size -f requ'encydis t r i but io n of juvenileAste ri as vul garis coll ect edby spat collectorsin PJacerrt;1a Bay ---- ---- --- ---- - --- ---- - ---••-- ••••-,;.- 192 Table3i. Wei9ht-frequency dis t r i but io n of juvenileAsteri as

gUl gari s collectedbyspat collector s in Placent ia ay --- - - --.--- -- - - -- - - --.--::.:-- - --- -- -- ---,---;.:. 193.

Table 34. SUlTll1aryofprevtccsreports of·t he growth rat eof Ast er ias~.and~.vul gar is __• .,_ ~..__~-.-n-

..

Pull'Capacity (g)ot"Leptasteri as

pO lf~s,

^{Ast eria s}

f~~g~h~~~ ~~¥~~~s::~_~~~~~~~~~_~~~_:~_~~~:~_:~~::~ .:

195..

'03

'06 '05 Table 36.

\

:~~~r~:~a~~fy~~~~~Ui~~~~~~ie~fp~~p~~~;:r~~d ~~l:~i~;

_{endecafort~reetrials} n. ._ _n _ _ • . n _ _ n . ..

R~su1·ts^{of a·}two~way^{analysis of vat-iance to test -} thesi gnifi canceof intraspecif icdifferences on the same trial(A) and differencesbetween three .ur f als(B) ,;,-- -'--::-- - - --- --- -,--- --- --- -- - -:-- . - --- Table 38. Resultsof a one-way'analysis'ofvarianceto test'""

'th e significance of interspecificdifferences in valuesof PC/Rforthe first trial--- -- --.---;--.-. 207 Table37.

Table 39. Pull Capacity/ Radiusvalues of thesecond andthird trials expressed as:a percentage of. the first trial

(R·lt

tach'bil1tyL=----,:-~--- --- -,--._--- -viii-

'09

.\

·-ix -

,twould liketoexpress my sincere epprecfet.tcn toIl\Y supervisor. Dean LA,Aldrich.who providedhelpfuladvice on the

initial conceptualizatio nofth,isprefect;'gave comments on varfous . . '_

stag~s'

of research.

a~d

^providedthe necessaryfi'

carryout thi.s,rese~rchprograrrrnefromgrant-in-aidof research No. A-1368fromthe Nat1on4l Research Council of Canada. Special thanks

mu~t

^{be extendedto}Dean Aldrich"for

re,vl'~w1ng a~'d

^provtdtnq

corrrnents onthe.finalrough draft wolle he was~inastate~of 111 health.

I .em indebtedtothema~ypeople who'helped'with,the~1el d.

work. In thisregard.Iam especfal 'lygratefu l to Bob Sm,ith.Neil BanntsterandChris.Emer son,who partici patedaslily,diving partners,

ea~h'

^for

.extend~/p~r~ods

^{oftime. 'The'}

e~pertise demonsJ:\~ted

^by

't hese three divers under trying condfttcnshelpe dmake'm~~^{d..i ff i cU,h} dives successful .

, DUring,the 'COUr)ecf this work, I had the good,.

fort~ne

^of

having had'a~essto the fine fac111tl~Sof th;,M.S-,a.t.. For this privilegeI owe gratitude to the.found1n~,t1irector.Dean~. A '-;Al d;'i Ch.' and his successor.Dr.D. R,Idl er. I atknowledgethe cooperatton of the UniversityDiving Officer.-Mr. W.

L ukeman.

Mr.M.J•.Merce'randhis

. ' .

,staff, andthe fonnerAdm~nistratlve·Officer.·Mr.J. W.Cole. I'woul d.!

~lso

.

like i'o

.

especially thank^, Mr.Martin,Hickey of.theM.5.R.L^. who captained avar ie tY',ofsurface vessels oftenun~er a~ve~seccndfttons,

• _ .~ , : . :...,...7---'--~---- _ and Mr .TO~Stevensonfor~~n--helPi.ng-:to~S i gnand .

. constru~&a~ent.· Iam alsograteful for theadvice

.~rr~~tructi ve

cri t :Cismgiv en

at-vadOUs ,st.1~es.·Of

^this

r~searCh

.:.--- ~Ymember-s~fthesc~nt1ffcstaffoftheM. S, R.L..and notablyto

Dr.J.R.St ri c kl er , nowoftheJohnsHopkins,"Uni ve rsi ty, for:his .h~ l pfu l suggesti onsforfmpli ment a t i onof thephotogr aphicsampli ng

sert esandana lys i ng data. I amalso tndebtedto"fir.R.Scaplen forproVid i ng

~nformation ^an"d ~terla l

^{regardingmusselandsea star}

settlement.

I wouldalsoli ke toaCknowledg~with thanksthe help given b,¥Dr?John AllenandMr.BrendanHolden of theFe c uft y of Engineer",:

log for theirhelp 1n.analys ingthe distrlbut1.onof,waterdisturb\ce

· in L~ay. ,

^{• .}

Ai d was al so'give by Mr.R.Fickenofthe BiologyDepartment, andM~.G. Hf1lier andMrs.P.-Yor ke of~he M.S.R~L .in photo'graphing' figures used inthisthesis.

Thismanusc ri pt was typed by M

,

s.Harian',Atkinson

,

andfor this hel p and h.erass i stancein other ma t t ers I am grateful.

I wouldalso 11ke to expres s mygratitUde to Memorial University of,Newf oundl andfor.... providing mewi t huniversityfellows hipsfor 1969- 1970, and1970-1971,.andf~rfi~anCing^{bytri p toW}ashi ngt on. D: C;for the 972 Ech1noderms Conference.

Ihave reserved by finalacknowledgement'f or~.!"{felCa rolyn,

"w1i hou whoseassistance.patience,(a nd

en.couragemen't~ t~is.iroje(£t

^and

the th

.

;is'wouldnot

~ave

^{' )been}cOOlpl'e t ed. Forthe activerole

~he

played nthe typing of theroug.~·dra~t.typ!ngof final tabl es , photo- graphic assista nce.editi ng.. and'pr oofreading.Iam.n,tOstgrat~~ul.

-xi-'

~--- ^"', . "

.

^:

.

,:.~ .

^{, . : :' ':"} GENERALlNTROOUCTlON.•

> , -. ~ .

' ... .. '

.. .

~

.~ . ...

~

, I

Factors affectf.ngthe localdistribution ofseetstarshaslong

. ' . i

beenatopic ofinterest to asteroi d biol ogists .andyetitremai ns one I

ofth~ mo~t'POorlY,under~t~odasp~ctsofseastarb101

r

Ogy• Man~autho rs have cemented onthisarea but havefailed to cometo grips withthe multi plicityoffactors involved1nthesecompl ex rela tio nships.

Mead (1900)was probably theff rs ~author to attell1lt an explanatton of thelack of uniformity 1nthedistributi onofsea stars ata specific locat io n. He~stabl~shedthattwo closely related species.Aster1~~vUls,ar 1s(Verr il l)"and.A.forb~Si (~sor)haddtsstnf Ier d1s tri but1 dn ~'resul ti ngpossiblyfrom"natural ba-rters " wit hin the ,environmentsuch as de'pth , density ofwat er ,.andbarrenbottom (t .e.

.lack of food).

Huntsman (1918) discoveredt~ at-,A.vu19al"'iS~asabsent from sloped areas..Thi s obse";vat ionhe.equa:t,~'dto,fl uct uating teeperatures onme slope.

.

,

'

fn

a major researc hp','Ogr~to:ontro}'i:~heoyste rp~~da~or.~

''''~.·A.te re es t , Galtsoffand'loos anof~ ~1939~under t ookanexte~s 1 vest udy

ofthe distri but io n.of-thisspecies'in' thewat er s.off New Eng1<lnd. They 'ruledou~-~mp~rature ,salintty ,:and substrate . as being importa nt :facto rs

c:~ntrol11ng

^its

d1str1~ut1on ~nd

related large

conc'entration.~,

ofseasta rs tothepresenceof molluscs.

Smft~(1940) .,conversely.tne simll a,r..-st udy onA.v~lqaris~ n.

~hewatersar ound PrinceEdw.ardIsland"es tabl1shed,thatl.ow sal.1ni ty.

" .i-> .

~ ^",

-2-

and hightemper at ure coul dbe11 m1ting f~ctors. produ~;ngnatura l nv:irtal1~Y.buta'1sorel at ed~hi gh density ofsea stersto ahi gh d!,!nsity of'prey.

Recently . other authors,havealsorel at ed -the- distributionof asteroidspecies toprey. H.~ncoc k(1958) give'sevidencethatthe '

.dis t r i but ion of A.rubens(L.)1'1sprimaril y.aff~ctedby the di stribu tio n

""\ Of,creOidUlafo~nfcata

( L.y.

a~.d'Larsson(1968 ).reit~rat~sthisconclusion but re f er s toMytil us~(L)as'the.natnpreyofthi s asteroid.

The effec tsof temperature andsali nitywere examined with' .respect to'distribut ionby Ursin (l960).studying,theechinodermsof

theNorth sea. He.fo.und.that temperature was lm1mportanttn"11m1tj~g thefrdistribution, witht~eexcept-tenof one species ,Astropecten ir r egula r is (Pin'nant). Of interestinthis stu<l,Y', howe ver ,'washi s conclu s i on re.garding the genus,Henrich (Gr ay) . Notfindi ng that tempe r a t ur e!ndsalini typer se accounte dfor its'd,1st r ibuti on, he relatedtheirdi st r i buti on tothepr esence of Sagittaele gans(Verrill) wat er .!;f,e. water'rich'in

nutrie.~ts.

necess ar yto,s atis fythei r

'mod~

or- sus pens io nfeedi ng (Rasmussen,1~'65 ).

AnunDer of researchershave conside re d the~i stributi onof est erct dsincondi tionsobvi ouslylimited by.s ef tnttys~chas 1."the Bal,ticSea (Meyer, 1? 35; Bratts trom{1941;,Kowalski~195'5;~.chli eper.

'f

19.56); inbrackis hwat er,(Bt'attstrom.1941;Urs in .1960);andinsudderi 'fr eshwat errunof f sHu ations_. , ,(Crozie r , 1920 ; Mac. Gi'nt tie;'".1939;Loosan off' ,

andEngle,1940;LoosarlOff ,1945;Smith , 1940) .

Verri ll (l914)was one of'the firs t to addre ss theproble m.of seasonal fluctu at i on,inasteroid populati ons. His conclus ionsthat

,.\ " ,

J •

,·1-

-3~

sea stars migrated intoshalle:wwater-forfeeding in the fall and returnedto deeper water'duringthe winte rper iodto escapecold surfacetemperatures.were.h~ever.ref uted by Galtsoff andloosanoff (1939).

Similar annual'mi grat i ons/by other sea starspeci es appear to be well.documented. O'Oonoghue(1924)studied thesUlIIlIer'migratio nof sea starsof thelit t oral'zone at DepartureBay. Brittsh.Col wrbi a •.He

. .

note dthat exposureto air during periods.of hi ghtemperature and di rect sunlightwas'the~stli~elyfactor cont rollin g thei"migrat.ionto deep water..'Hewatt,(1~ 37 )found'that leptas t eriasaeQual is (Stimpson) migratedt~.,~eeperwater for reasonsessccf ated.wit h.reprod~ct1.on.

The problemof exposure

. .

tq air has been discussed by^- avari ety of euthcrs (SlJlJllloirized byFeder andChristensen; 1966). Recent ly •.

landeriberger'(1969)noted

th~t the. d1ffe~erices · in

^abl1ity-of

thre~

,speci es of thege~us'Pisa~ter(Br.ndt)to cope with exposureto air

hasre~ultedindifferenc~sin the'distri but i onqfthese·speCies.

. The questionof'subst rate .asafact or'affecti ng'di s t r i but i on',has been'dealt with'~stadequatelyby Gra.inger (1966) in,his dtscusstcn

~f

^dist r'ib utio nofsea stars.inthe Canadia nArctic.'He showed that species canbedivided'tntcthree,groupsdependin,~u'ponthe't ype of substrate on whichthey arefound.'Hisfi r st and thi rdgroups,compr~ se sP:cfes thatare found'only'onsoft andh~rdbottOms.respecttve 'ly , a'ndht s.secondgroup incl udes those'specf~s'whi ch'are found"ana wi,de range~fbot tom

.-

~pes.

Hence,'from prevtcusstudies . weseet;hat seastar~'istrib~tiori

.-maybe contr olled by a'wide,r ange of pa-ramet ers. 'severa1crfticisms'~

hoWever•.can be dt rected at the

s~ope

^.of

.~ny ~t

^these

" stUd~~~

^(but

I

• . .

-4-

notab'lynot /111); Usually. distributionhas been~ons1deredⁱⁿ terms ofone or two teeter s whic habsol utelyexct udee par ticular speciesfr~m'aspec ific habitat~ As yetwe do not have acomplete appr ec;.fati onof'why abundanceso~a parti cular,speciesvary withina parti cularlocale.

or

~hYthe,abunda,nce~di stri but1onpatternso~

d1ffere nt£tpecles vary; Most of thesestudies have deal t with the .probl emof

~

^{a,s peci}es is presentor absent

~-n

^acct tecte d sample .

and~ot.

0."

cbservettcns madewithin the ani mal s' envtrcnment•.M~ny types ofhabitats. partic ularlyrOCK bottoms. have received l1t ~leattention. and many specie s haveyet-tobe di s cussed. 'The majo rity ofauthors have fail edtost udyhow a species int era ct swith '.its envi ro nmentto produceitsin~iVi d ua labundance:d is tribution,

peptems .

Durin g'the sunrnerof 1968,wh~leasunmerstudent working at the

Marin~ '

^{SciencesResearchlab}or atory for Dr.

.i,

Ber ger of

~h~

~n1verSi~y

^{ofrcr cntc, I}

bec~

int ri gued wi th thi s probl em.. Followi ng conversat j,ons withDean F. A.Aldrich. thenDi rectorof theM~ S . R.L•

.and'with diverswho were'empl oyed to collect specbre ns for theM.S.R:l. ,

1-became aware that thevari ous seasta r~peci esfound inLogyBay haddiff e rent localdist ri but i onpatte rns-.o,.,ing·tolIlYint eres t tn' this topic.twes encourag ed byDean Aldri ch"to undertakea Mast er 's progranme toattempt'to eata bl.lsh what factor s'were responsiblefor

cau~ing thes~

^apparent

s~~~1es d1ffer~nc~s

^{in.dis t}ri but ion•.

Afte r'co~plet1nga~exte ns 1ve,.l1te rat ur e survey,lwas'a~ie.d to'discover that no,in fo~t1onexistedon.t hebiology'ofseast ars in

Hewf~uk~~and.

^waters.

'~hat l1ttl~ info~tion'~~s a~able

^{o,n;t he}

phys ical,cbent cat,and bi ologicalcharacteristics,of-thet r habitat.

.. . \ .. " .: .' __0 -'· --

\ .1

,\,\..

.

^,

-s-

and that lfttle.H-anY1,of-the.general.as"teroid1iter~turecouldbe \}

directlyapplied,t othis topic. Hence, Isetoutto studythe"spatia l andte~poraldlstributio~^ofthevarfou~^s-eastar~^{species,the..}

biological characteristicswhich may affectthelrdistribution,and the

en~1ronmental p~rameters affe~tlng

·t his relationship.

,':.1

j

,t I

'.

A. Study Areasee Stars: :~S-~~stemati~.~.Descrip~.i6n.^{and Geogra}phic,Distribution

J;: 3 "

~- I Asterias vulgaris---:---., - ---...-- :-- -- --- -- - - -.:.- --.-- - -- A-I. 1 Description --.:.-.- -. -- - ••••-r--r-r-...~.•,-..-. ---••---. -- 7 .A- I.Z Geographicand BathymetricRange---'~- .- --- .- -- . -- A-II Leptasterias.polaris

.--- ---~---.:---;---:.- ---

^.9'

A- I1 . 1 Description. ~ "';"';-.-- --- --- -- -..:- 9~

A.;II;2 Geographica~dBathymetricRang'e-.-- -'."':'- ••• ----.--. . 11 A~ nI ~endeca .:--- ---- :---••---,. - .--- ---:.---- 11

.~.:.I I,I.•

r

Desc~lption--~---;-•..-.-.--...--.---...--- -- - --.12'"

A-Il I.2 Geographic.andBathyrretricRange--- ..:-- -.----,. 12 A. IV Crossa~ter p~ppos~s"":""? "••-.'7 ••

-- ---.---.u----·:.-.:.

.12

~-IV. 1 Descriptfon -:'_. ••• • •__•• :. _::--..14 A...IV.2. Geog raphi c~andBat hyme"ti' i C Range.- - -,- - - - --- - ---:-- -- - 14 A-VHenrfc1a

~ .---.,--~----·..-7---~_:·-:.--·~--_:·:.,.----;-

¹⁶

A-V.l nescr-tpttc n ·---':::---:- --- ,.- - --.,- --7--7-- - - -- 16 A-V.2

Geo~ra.Phic a~d'

^{BathymetricRange}----::- - -.---:.--.;.-. -- 16 A,.VI Unide ntffiedSpecies--- - ----.--- .-- ••--~--- -- 18..

"

~".

-7-

~:

^:St u;y Aroa

S ,.

^Stars,

.~:"'ti", . .

uescr-tpr tcn;endG,ographic ' / Distr ibu t io n

Sevenspeciesof sea starstnhebfI tthe-study area,fi ve. of .whi.chcouldbeeasilyIden't'if ted to species,wher~asthe re mai,ning

two spectes presented texcncntcdifficulties'. The'followi ng section lists thedescr-fptt on;systen atics;andgeographic and

bath¥me~r~1

dist ribu ti ons ofthe fiveposi t i vel y'ldent~ f1edspeci es. Brief notesontheun~dentifi edspeciesareincluded.

.A- I Order Fcr c t pulate FamilyAster1 1dae Species As t e r iasvul gal"isVer ri ll1 Synonoll\Y:

"As t er 1as vul garis Packard.'~ 863(nomennUdum).

Aster i as stimpson;Verrill .1866.

Aste r i asvufguis Ver rHl .1866.

As te ri aspall ida-Per~i er.1875.' Asteriasfabricii,Perrier .1875.

A-I.1 Descr~pti on: (fig.1)

Di sc ofmoder atesize , armsfive,IIIOre'flattlm~dand pointed'th an

in·A,forbesii .'Abor alsurfa ce ; network of

na.rrOw· b~;~l k~ Plates.~ith

^.

.--JT'ort onese:0 963.cited by Swan.1966) states tha't'MA.vul~ar1s^{is "}

conspectftcwith'A;r ubens andnetevenseparable on a su sp:ecific basis.~

J~~a~~,e~~is~~~ 'r~bih~

^I

~~1~~~;~~~~~d I~O~

^s

s:~;ga}O~~d,li~r~~~ _ _ :~~~O~~hern

^..

the:BarentsSea, almost'as far as SvyatotNos 1n~laBay:.andin theWhite . -s ea.Recent ly.'.ithas.been'Tound off the coast of.Greenland. It sbathymetr1c~ r,"g." 0to 400meters•.ID'Y.kOOOV',19SOl. . '-'._~~:

\. ..:l

"

-8-

..

Fi gure"1.· Ph~tographof'the aboralsur face-of-Asterias vulgaris."

.'Magnificat ion3x. Refe rtoth~:extfordescr-tpefon.

I .

. )

-: J

"

-9-

~rgemeshes, making.afairlyW~~k'Sk~l~to~.·^Clear Ty percept tnIe rowof nedjen plates. All plates with one ormoreblunt splnes,with

rOU~~h

^__

t1 P(.··~~CirCled

^{by a}wr eat h of blUl'!t'pedi cell ar i ae: lar g:

acute pedtce tterteealong.stde s of-eras,risingdirectlyfromdermis.

Inferomargi nal swithccnsptc ucusseriesof long,truncat e spi nes.

Adarrbulac ralswith one or two,'long, flattened spines; Rr=3: 1..Colo r reQd ~ Shbrown.madreporlteusuallyyellow. " fromGraye;a1 (1968).

A..r.2 Geographicand Bat h'y!!!etr i c Range

'!he 9f;!ographic·range.ofA.vulgaris is frollt labr adortopape Hatteras,"occupyingabat hymetr ic rangeof0to650meters(GI1l y ....'-:

etel,.1968).

A~lI OrderFor ci pulat a Family Asteriidae

. . . .

Species .leptas te r i as polaris (Mulle r.andTro~ che1).

SynonolllY: /

"A~ieracanthion

pola r i s Muller,andrr oschel,1842 . 'r

r ?'

A.polareDuncanI andSfeden ,166l.

/

lepta sterias'

p01ari~

^Fisher.

1930~ ~

. Lpol ari s 'acervataFis her .1930.

__._.... A -rL

¹ pescr-tpttcn(fid. 2) . (

U'sually6 rays. R:r atleast

~.5: 1

^{to 6.3;'-' Aboral surface}

J ' .

^{astronq,fai r ly'}lose -meshed networ k.with spi nes whi ch ;ary greatly

Z -:,- ' , . ...

^!..

^;:::in

^{size.shape,and}

~rrangeine.nt .

^{in the}

d~f.fe~ent_

^{reporte.d.sUbs·peC.i f i c..'}

forms. Abor alspines.gener.allyofst ouu.c onstr uct .tcn, frequentlywith

"

.

^\

. '

^{. . "}

. ' .

-. bulbousends {althoughin sometheymay be-reteetvel ylongerand .

/ ' • . 7 '

l /

I. ..

^.;

-10-

. .

FigureZ. Photograph of theabo~alsurfaceof.Leptast er tas polaris . Magn,1ftcatton 3 X. Refert~.thetext"fordescr-tptt on;

.\

c .

I

-11' .

.

cY1indr~~al

^tothe tip). and

~f~n

^tn

gro~~s

^of

-S!'1eral ·s·ma~ ~.

^5pJ!les·

surrounding^II'hr gercentralone,or less'frequently as so1itarv spine\Of"ary~ng^{de.nsUy. Not}1~freqUently^abOr:ll1~P1ne_ 'g~uPS

.withsome.s 6ll'blanc:eof regularity along the carinal rowsatleast•. : and sceetteesadditionally in

,

one dorso la te ral^. rowlongi t udin all y alongeither sideof.thecarinals; elsewher eusu,allyscat tere d.

Hargi nil:"and

~~-n~ro1ateral

^spinescyli ndrica land

lo~ger ~nd:r'elllthel~

moresl ende r'thantheabor a1s: in 3 or 4consp 1cuou~lyregular long1tudinal'r~s^{along muchof} th~^{le ngthof}

the

rays•.Adarrbulatral. spines,cyf fndr -tcal,relativelymoresle nder an'{ shor t e r thanthe la t erals~1ne;.·freqUentlY^as2~^{less'oftenas 1}per plate ..Somet imes long consec ut iv e seriesof l: pe r-pla te pairsof.spi nes alterna te regula r lyinpcsf rtcngivi ngsuperf i c hllythe.impress ionOf'~rows' of spi nesal ong each side of the groove...·frOlllGrainger(·1966).

A-IL l Geogr aphic andBa thymetri cRange

The geographic'range.ofth 1s.s pec i es 15 frtllll the Be~1f!9Sea·. eastwa r d toEast Gr e:!nl and and sout hwardtnthe westAtlanticto NewEngland. occupyi ngill. depth,range of 0to110 meters (Gra in ge; . 1966).

A-I) I"Order Spi nul osa Fam11y.Sola steridae

. ,S pecfe·s Sola ster endeca

(L:J.

,Syn?noll'(Y:

~Solasterendeca.Duncan,andSl~den. 1881;

·S,en~ecaD,an1e1sse n and KOren. 1884.

·S;ende~~ Dod~r!e1n ;.1900.

':S.endecill. Fisher . 1911.

- - -

,.

.S.endecaHorte nsen ,1927.

~, " ' '

..

^:

;'.'

'f··,

-12- A- Il L l Oescr i pt ion.( fig;3)

Rays'7-'13, Inost frequently9'or

~ o. R: r

^{about2.3:1to3.3: 1,.}

Aboral paxill ae small ,.hi gherthen broad,moreorlessscat~ered'.

w,i th fromatleast4 toabqut 15spines. Inframargin~lplates lar.ger than supr amargina1s,transvers el ye1EllJgate; Supramar ginal

.paxt 'llee onlyalittle larger thanadjac ent~orsol ateralpax1,ll'ae9

butclea rlydistin gu' s hable fromthem;.Inner adanbulac ral spines set inthegroove,fr o'!' 1.or

2

t,o4in nunber.lIluchshort~randless conspicuousthen thelarge ~out eradanbula cralspi nes. These,fn transver sel yplaced rows , vary from about 3to 7or 8in number.the

,

..

^{, '}

.

inner most sct nesthelongestandst out es t , the out ennost thesmallest . Tube- feetin2rows. Oral int er radi alspaces with pax.illae ....from

~ra f.nger(1 966).

A-.JII.2 Geographic,andBathymetr~ cRange'

Thi s'sped es rangesfrom the""eastemarcti cto thexereSee , south

t~

^New

E:ng~and

^{.t n}

the··wester~'At1antiC. a~d

^southto'the'Br i th h Islesvt n·t he-J.st emAtlantic'.

I~:lhe eas~r~'

^PaCif1 c!,1t rangesfrom' Point'Bar row",Alaska. south to

va~couv~r

^lsland,'.

~e~ ·range,. in

nor t her nCanada iSo'"8.to 118 meter s(Gra~nger,~19?6). ' A-IV-Order Sp1nulosa

Fllm'lly Solasteridae

Sp~des

^Crossaster

papp~sus

^1,(L.-)

.

lM~ny ~~thors .

^{'do nOt}differentiate '

b~!'twee~ t·he .g~nera ; ·Crossa~\er··

^and

Solaster;however ,convincing a,rgumentsby D'yakonov~edon

~cesout1f.nedbyFisherjustify the.usage of.the,genu$"t rcsseste r , Recently.many authors haveused'Crossas:tet'ptlpposusas ,the naiii!OTtJiTS· specie s (eg.Castilla,andCris p, 1970;Mauzey ,etal,1968 ;'and Feder .

and Chtiste nsen,1,966 ). _ ' .- -' , '

.;13-

.Ftgure 3; Photographof the-abor al·surf ace ofSol i st er endeca.

t1agn tftcatt~n :3

^{X. Refe;tothe text-for}descriptton.

..

'-,

, .

..~-: : '

....

^;

I' -14- SynonomY:

'"cros~asteroa?posus DuncanandSiaden',.~'BBl

.~.Ql!.~!~!"~ucder tet n, 1900.

~·Grieg.190i .' S.pabp~·.·Ftsher-.·~·91 1. 's"papPOSUS-Mortensen; 1927.

~Hed1ng . 1935.·

... .

A-IV.1 Description (fig .4)

E'ightto16reys , lOO,stfreque~tlYlO- 12l:r about 1.7:1 _~o, 2~7:1. Aboralskeletonan open reticulatednetwork. with.up~o50 spines perpaxf lfa. Inframargina l plates with a,si ngl ec~nsptcuous row ofpaxil 1~e;supramargina ls a11OO5tindistinguishable'f rom abora l pa~nlae. Inner ad'arrbulacralspinesarranged parallel tothegroove;

nunbering

us~.allY

3-4,

occ~siOna;lY

2 to atleast 7;outer'adam- bujecralspines tnrows at rightanglesto 'thegroove-numbering usually about'S-7.rarely

.

3-9. Orafplates

^.

with 6- 11

^.

marginal spines.

TUbe-f~tin 2r~s.N fromGra1~gey{1966).,

A-IV.2 Geo,graphicandBilthymet~1c Rance

c,.~app·osus^{is anarctic}bor~al^species'exte~dingsout~ard ~o~

40° N'(appr oxi mat el y Ne,wJersey) in the western,Atlanti c . andtoGreat Britain,in,the easternAtlan~ic. Inthe Pecfrtc,it exte nds'to'Vancouver Isl andandJa pa n ontheeasternandWeste rnsides. respe ctively.' It

' " ~ , . , , .

h~as,an extrema.lyWi de,~ath~etr1c_ra,n9

r

^of,0 to-1200 aete rs(0':yakanov..

1; 50) . ' - \

7

,d y~'

. .

,

F19ur~4.,Photog raph of. the aboralsurf aceOf~.. ,'~.' Magni fica t io n 2X. Refer t~ th~-t ext'for.descr-tpr tcn .

-16- A-V Or~ei''Spfnulosa

Famil yEchinasterfdae

Species Henricheschr i cht1' (Mul l e randTrosch~l). Synoncmy:

"~~Mull er andtrosche'l, 1842.' Cribr e lla~Dunc a nandSl aden.,188l.

Henrie fa~,eschrichtiiHed i ng~:1935.

Henrkia'eschr i chtii ~schricht11 Gra i n g~r.1955.

A-V.l Description (fi g.5)

Fiverays.R:rabout.3:,f to4:1."Ai?oralske.1et on'1 rregul arly· and closel y ret i cula t ed.-dcrs c'lete r e'l,marginaland oral surface ske-le t onfa1rl~regularly'1~br1cated .withpapularareasmuch \ narrower . than'plates. Ca~jna1plates'not,r~ad llYdfstfnguish'a~le._~

Aboral pextl'leewithabout4-40 fairlysmall,spineswithusually3or

' . 1

4lateral'"ridges ext endi ngbey ond thetip~t"acrownof30r4

. t - .

diverg,ingpO.fnts" Mar gi nalplates la rge.c~nspiCuous .in eVen'imbriC<f)d rowsand incl os e contactwithadjacentl'pws. Adarrbulacral plates.wit h usually 2,(somet i mes3) transverse rowsof'about 8-18spines.the ,. la rgest near the groove" Tube -feet"in

i

rows:,' fr omGra1ng~r^'(1966),

. .' >

A':'Y.2.Geograph ic andB~thy~.tr.fcRarige

Si nce ~~Ohconf us fonhas existedwithrespect~othe Ident ification of this spectes , ttsdfstr1butie:n is'dif fi c ult to determine. Grai nger (1-966)defined"its North

Amer1~n di'str1b~tton

^'as

'be~-"9

^{'f r om}

NeWfou'n~l~nd to Baffi ~

lsland:'and havIng.a·dept h

ra~ge

of'14(sic)

t~

148 meters;'.

I . - - .- .

l.,stnced~fflculttestn1dentificati oIJof this-spec'les'eX:1 st"'spe~imens weresent toGr~in~e rwho.~confi rmedmy identificati on. . ^$-

·. 1:-., .

,i

...> " ,

^-17-'

..

Fig ure 5. Phot ographofthea·bora1. surf aceo~'Henr1'di1f~hr1ChtL Hagni f 1cation5X.:'Refe r,

t!l

thetextforde$.~ri ptton.

.:'

":.'.

.,,;. >-.' '

, .

-,'

/ .

-lB - .

.A-VI. Unident ifi ed Species

Of"th:'two

:Un~.d~n~i.~~ed S';eCfe<p~e

^ts

, ~~t

defi nite lya

speciesof the gem1sHenrfci a (fig.6'). Since~rtDersof this genus ,have proven.tobeI'chro~ic^{sources of}tr~uble^{tostudentsofthe}

.qr cup'1n

_.

nor-thernwaters"~Grainger._;. 1966) .. aninte nsiveeffort

to

ident"f fythis species was under take n:. 'Itwas-felt'thattht sspe~ies

. mo'~t.'clo5elY

tesembled H. scabrior

_(~1khail ovsk; i) .

^but

demonstra~~d

^/' sev~ral.characterist1cs^{whic h did}~~t,ag'reewith the descr ip t ionby Gra i nger'(1966l4"O~th~S^{sbectes, Afterexaminl1\9}~evera_l^{ofthes e,} '~pecim6is"^Dr.Grainger(pe;'.~on..alco~o.nicati.on )^comnented:

"You,:,~Imus'tque~tion.although'! can~ot give itenctbe rname ".Eitheri tis somet hing not ye't na~doroH.Scabrio r.haswid~r:'~tructural l~imits than I,once thought.'"

~

Sinc e identificat ion

o~ thi~

spect es. fsunclear',

It'is' ·refel.':ed. ~ o

^:.:

'to as"'Henricfa sp,

th',~ughout

this thesis.' ' .

,":'A si'mil ar

sit~~tion e~isted

^for

th~

^other uni dentifi edIs peCi es

·(fig..J). Based,on a number'of taxgno,"ic charac teris t i cs. it should '

°be,assig!!ed"~othegenu's Le'pta sterias; however .positive's p'ecfes ideritifi cat ioncouldnot bemade.l. Since

~pecimens

^{of thi s}

specl~/"

closel y~sembledj~veni1e-~vulga r1s2.' ,bot h observatfon s"of. thi s

spec;e~ 'a~d o~JUVen~le

^A.vulgaris

ar~

excl uded from

th"~S 't.hesis' ·

.t oaVoi dco~fusi on ."

,

lspecimen~

_{too~asullable}^of

thi

_t

~

o,positi vely ident ify^{spec ies}

~ere

^also

e.x~

.them.

' by.Or~

0" ^{Gra 1ng\r}" • ^a.•^1)d

h~

2und~'rwat~r""~l:tserva;ion's

^{of this species}brood;in"geggsin aeenner .

"sim11ar to Lhexact 1s (Sti mpson)."(Chia.1966).demonstrated,that. .

..these,spec1.~not.~arfant~~~.a"non~brooding SP~~1es. .

I .• - ' .. ' .. ..

I . . .

,f :;'"-'

^:Pi

7 !

F1gure6•.'Pho tograp~of :,heabo~surfaceoLHenr1da·sp.

Magntf~cJt~on5X.

..~

Figure7. Ph~tOgr~Ph^..Ofthe aboralsur! ace ofleptaster1as~p, Magnification5X.

··1

-/

B. Stu dy Area

~ B·I

Introd~ction - ---.-~---

^...

----.-.---- --.-~ -~---- .

^23.'

.B·II Topography--.:-:--- :...---.-- -- ••----_•• -:.__,._-:._._.__ 26 B- II . 1 Materials.,andMethods.-- - ---:.-.--- •••••• 26

J .

B-II. 1(i) TransectLine-. -.---:.-- -.- - - --- -:- _•• • 26 0-11. 1(f f).Depth.----~---~--••--~-~--:--.-:---. 28 .B.I I.l(~f·i)' Slope --.-- •• -•••-- -- --. -.---•••• •---__ 28 0'-11. 2 R~SUltsand Discussfon-- - -- --•••-.:~.---.---•• ---- --__ '32'

B-II:2 {O .~-Ala r1a-lamin aria(HAL ) Zone.--- - - ••---- 32 B-I I.Z(fn C11f f Area.-- --- ---....----~:.--•..:.--.---.-... . .32. B-III Physi cal'andChemi calDescript ion--- --.-•••••--- -- -.:-.-,..35

B-II I. l ttater1alsa~d^{Met hods----}: ---.•-':'~-~^••--~---__ ^35. B.I ~ 1. l(n·^.Temperatu~i^Depth,Condu7^{t !v!'ty}(Sa1fnft~) ^{.; 35} B.IJI. 1(ft )·Dissolve dOxygenOete.:m1natfon--.---- --.,;-.--- 37 O-li1.1(fti ) Relative lr radfance(R.I:).~•••--.--- ----~--- 39 B·II 1.2-Results andOfs~uSS1 00-.:---~--':'-^•••---.--:-~--- ³⁹ B- Il J. 2(i) Temperatu re--.::••~---••---:- - -••__':'. _•••__ 39 B-I lt.2( H) Salfnfty'(c~nd~Ct1V1ty) ^• .L~_________ ⁴²

B~tt1.2{1f i ;, ~i.ssohed·

^Oxygen

Co ncent~~t1on

^{." ••••}

-.----:--~

⁴²

B-III.2(1v) Relat iveIrr:adi ance.--.-- :"'- -- • .44

·B-II ~.2 ( v):Wat er 01sturbarice ----•••---__• ••••_.. 44 B~IV Bfota:.--- ---:.:. -:---- - -.:---.••••----.- - ---:-- -.__.• 52 B-IV.1 MethodsforDetemlinatf onof Bi ot a--- -,..---:- 52'

·'B- IV.l(i) Gener alObservat1o~sof Flor aandreuna -.•-- - -.-- 52·

.B·IV.1(tt)

PhotOg~aPh1 C ~alys1s .--~~_

^••

~

^{•• ._.,;}

~

^{.;,. 52}

'...

B-~v

^{•.i( f11) ,Han.d,SamPl eTechn1Cjue}

-.---~.-.-~-_----.-:-.:-._

^52..

~

-22-

"

~:.

8.I, .2...;,fts endDiscussion

---:"---r----~·-- ' ~3

B-IV.2 {f) Biota of HALZone,----__:.':'_':' .;._~.... .._:-.53 B-IV.2{H) Bio taofCli ffArea ------ - ..- ..--- -':' -- ----- -- - 55 B-IV.2(11.a) GeneralConsiderat i ons --- -- - -.,.-....-- --....- 55

~..IY.Z(11.b) PhotpgraphfcAnalysis .....-..----..----.--..--- 56

B-IY.Z{ft.c}'HandSample Technique,---::--~:_--- .56

B-IY.Z( 1t,'d) G:eneral

Observ~tt Ons

^and

sl.lllllia.ry .~Of

^{Fauna-- 59,.}

' .

.~

' .

.'

.... ...

^:

^..

-.

-23- .8•.StudyArea

8-1. t"ntroducUon

.

Selectf~m

^{of a stu4y}

'~tte

^was

goYe~ed

by three eajor ..

crfte~ia

^{which we'renecessary}

f~ o~e;'

^tocarry

~ut

this-study.

1. A habitat'was'requtredwh1t~possesseda largedepthgradient ,. so that the distributionof'ast eroids couldbeconsideredover

arangeo~phjsi c:al;chemical.and b10log1Cal·paramet.ers.

2•.A highd1versi.~Yofasteroid spectes was-re qut red tor inter-:

specificecnaerf scn.

3. Ino~erto elimi nate proBlemsoftranspor tatI onof equipment andpersonnel ,'a c'loseprcxtn ttytothe faci liti es oftheM. S.R. L wasrequired•

...k! ideal locati on according tothes~cr:ite r ia wasfoundatthe., .head of LogY Bay,a bay facing nor th east er l y1nto the)l.orthAt lan t i c.

Th~ geOgr~ph1C

centre of this study area··hadcoordinatesof 520

.sh,

^54-W

:.l0ngi\ude,and 47037' 37-N.lat1t~de. .

.Boundar1~ .of

the study area

(~tg.

8) were·dete nDined.such that

l

a unifonPsubst rate of reelcongl omerat e bedrock characterized all. .~ortion~of the stucfy.area.:Bas edon topographicfeatures thest~dy

areawas div1ded into b!0 Alajorsections:

1. A"shallow

wat~r

'Plat eau',roughly

reCtillng~lar

^{in.shape, whi ch}

exten ded·seawardsfromMa~ ROck·1,:.~·20^{"meter s having}a widt h of 20:.30meter s ·wasnamedthe~afte r its three~or benthtcccepcnents ,

!:tl!!l.!!!.

edulis ~Ala r h es cul ent a (LJ an?. Lamina~1a^d191tata(Huds.)~

:...'.

.

,~

,:."

-24-

Figure 8. Drawing of logy'Bay showing thelocat i on'

or

^the.st~dyarea.

A-·MAl·zone.

B-Cliffarea, C -San'dbottom'

r»

.p

~fJ

I.

·f

LOGY BAY

....."

...

l

' N

o

...

"/

"

~ --25-

2. Asl oped areaof

b~ttOm

^{extendt,ngfrom theedge of}

~h~

^plateau

atapproximately,7

~ters

^{toan almostlevel}

s~nd bottom

^{at ..}

31met ers. Si nce no benthiccomponent swere dominantover the enti r e

~ange

^{ofth,:'slope.this}

~rea

^was

na~

^the

cfflf

^{Ar e a}

~fte~1ts dlXl1nanttOP~9raph~ C ~~atu're^•.

-.. : .j

,/ '-

,.J

.:

.•..

.

^{'; ,".}-:' .

. " .:: .'

'..~:. '..»i .

B~U Topography B~Ih1 Materi als andMet hods, a-I L l(i). Trarisect line(TL)

Inorderthat-apoi ntofreferenceexisted forthe var ious, divi ngsurveysunder t akenthroughout the,course of

~he

^{fiel d}

study~ . '

a

trans~ct

Hnew as posi ti oned in the cl iff.area2; .'

I ' ' " , - , , - -

NotetheimportanceoftheTlisnot restrictedtoitsusage,for merel y surveyi ng the topographyof the cliff areabut as.a key refere ncepoi nt forot her surveys·,admi ni ste red inthecli ff area includi~ gsurveysof biot a .and asteroid dfstrlbut fon, .

":"2 Thenwasnot extended'int o'theMAL zcnebeceuse of thehigh

wat er disturbance,'peri odically.characteri zingthis zone, which, would havedi Sl Od9 ~dit. . -

;,

-27-

The'TLwas a\',~braidedPOlypropylenl~rope~hich w~sheld

secu~to

^the

san~

hottomby mea:sofa large"burlap 'bag,fl1led with sand. Toensurethat,the rope did ncteove on the 'rocky slope , the Tl wasaffixed to~hro~-steel P1ton~'l^(Hia:tPN7) whichweredriven into cracksonth~r~ckyslope.,In total:

lengt h, the Tlmeasured 36i11~tersandext ended from ad~Pthof' approximatelY9met er to a deptho~.31meters(the horizontallyflat sandbottom):.The Tl was alignedsothat.t trandown the gradie~t.

,pa;allel to

th~

^{incl i ne.} At fnterWlls'of1.5.seter s,nUmberef' . lam1nated pl as tic car ds (2" X 2") wereattached.to then tiymeans.

of short,lengthsof nylon cord. In orderito maintain the nylon cordat

a

fix~d_posH~ion,a small quantttyof "Sea Gain" pcxy

'r: '-

putty was..-J}1aced at,it s-in" t er sect.i.on,w1t hthe Tl. Thus when

~7~~Pl~ted , theT~~~~1l'siste~of twenty-four1.5 metersegments

'.(call edt_r~,ns~et~1~equadr,ats,'!L~~S),numbe,~i ng'f rom 1 to

2 J'

I'

i ncreasi n9, ~r1 CallY

^fromshallowto,deep water. "

'I

'--'---'---=~---'----:...;

..

I

. ..

-28-

·8-1!.1(tI). Depth

S1~e

^anannual

.

^{'Udalheightrangeof}. ^{5.2 feetlexisted.}~ .

in.logy Bay,"the topof'HadRock2 was'selected"asthesuriace'leve.t•. Measureme~~s0(,depthwere markedon.then'at,de pth s~f^10.15:. ,_

20. 25-aOO30 eeters.onJuly6,'1970o,ad,!y when~e.surfaceofthe

·waterwasfJat call1l. The tidal heightat St. John' satthe ti me

· theeeesurere ntswere'

~aken

^was1.2feet

^{i .}

^A

mea~Ur1ng

^linewas

attached'toiII..surfll,'ce floata~dmarkedat'on~met:er.inter~al s .withtape; andat5meter-dept h...1n te rva l~withIItiedloop.

To det ermineth'rposi t 1i;nforea~h^5eeterdepthmar king...a

·~ook:attachedtoa di'verlslead we i ghtwa~hung"f rom.t he required loop. Apieceof stringwas.ti ed around!the TLat the pos iti onwhe~^. theleadwe1gh.tjus ttouched the,bot tomwhent~eropewas taut end..

perpendicula rto the surface •.Each.5mete r depthmarkpositi onwas

.

^{. .}

recordedwith respect to'~enQ n1Jlltleredtags-,.Thedepthof'the"

sand bottoinwaseees ured from thetapedone..ete~intervalswith the,weightat-

th~

^30meterloop. This val ue

w~s

^.1aterveri,fied by useof-asoundingttne froma sur.f acevessel.

".a-I L l'(Hi) . Slope

To accuratelydetermi ne theangle ofin,cl1nati ~nof the i~regularly~l opedstUdyarea.eeas ure nents of both~helengt h.an(,

"tile~lope of-~veridifferentl yslcpedbortoesegmenthad to be

.

.. .

lCanadian Tideandcurient'Tables..Vol.1. The Atlantic Coas t

~nd

BayofFundy: Canadian Hydrogr aphic Ser vice,1970.

2Fig~re'

^8showsthe

l~ca~-ion

^{of HadRock.}

~ear ~~.

^uppe';end'

of.

the.."

tt, ~neral ly,'it15'exposed at low tideand coveredath1gll tide~ .'(under calmwate rconditions ) . .

'3AS^{in 1.}

' . ,&. " :-••... .

"

...

,'

.

\~. . ... ."

"

\-/::.

-29-

taken. Witht~e^i<;1,i110mebir ''( fji g',

9)

hetd

paraiiel':and

adjacentto then.,

the-~ifferent'lY

^sloped

r~gjons

^weremeasur"edan"drecorded

".w.1t h respect to,t he TlQn~mbered tag~. F1~.10 shows'the stepwise procedu:efo~:m~asU'rffl9;heh~e9Ul~r.Sl op.e'at.TlQ~~ C~i nom:ter readingsIx};because of;theposit~_onof-the pro,tra.ctor,hadto',

.~e

^{.converteltoangle: .of-inclinati on(8)by}

th~' ~ormula : e

'~~Oo,'.:

.x.

. i-' r

"

/ '

..,~".

,

.

- i ·

"~

: .

a -protractor

b-Pl.umb1i.~e

.-J'." . ". ' . ' . .

c-sc~lemarke{every 10~ondextonframe

\"~:"

.'

~

..

^"""

..

'

."

.4.

".-. :

-31-

Figure 10. ,pi agral11Tlat 1c' rep,resent at1onctf·the·stepwis.e.procedurefor

.

.

^{: '}

det ermin i ngthe slopeof anirr~gul arbottom.(TLq3).' The clinometer was positionedsuchthatoneendwasflush withthenunberedtag.at. TlQ3(Al,'an~thedi~tance(t o

<thenear es tlO'on) ofu~1fo~slope wasmeasured(here• .0.8met ers ). Angl eas measured by the'C] i nome t er (86° ) wasre~orded. Todete rminethe:angl.eofinclinatio n(9), angle (x) wassUbt~a.ctedfrom 900 •Le~B,.90-~ . B s'hqws'the'measurement,ofthe

n~xt

slo pe"(1,.e..thedis tance .

frci~^{the-last m}eesurenent tothee~d^'01'~he,ne~tunl~otm segment) andc'ltncnete rreadf ng. Val ues were'0.7mete r sand

O~

(x) orij"'"

9~~;'

^.

Sfml1~rlY~

^-C'shows,

me~s~re~nt'

^of,the

next slope,from~helas.tmeasurement to,theTlQ.4marke.r

~ag:. Val ueswe.r~0.3meters,and 77°(~ror-.13°.(9).

, I

.

~ ,

^{, , . .}

. ~ .'

.

.

^'.

· A. " .

" . .'. . " . .

"

<.

-32- B:-l1.2 Results~andDiscussion B~II.2(0. MAL Zone

Except for a few trenchesdi~playingthe northeasts~rike_

0 :

thered conglomeratesubstrate.the~Llonewas:'almost-unifonnly flat. incTfned ata

Sl ight-ang"le~~hat

^{depth of}the pla tea u .

-c- .

ranged---frbm 3eeeersat the base of Mad Rocktc.emaximumof7

~ ~e~s at,~ts pe.r~meter.

^Ithadenapproximate.area of600 square

~, meters.

B-l1.2 {ttl-.CliffAr~a

The slopingl>ottom of thecli f f area wastncttned at'amean angle-ofapproximately_.40°.to the horizontal;'however .fourmajor areas of'differingslopes couldbe'clea rlyrecognized. These areas were d'esign.itedB. C,O.E. from JL7'met er s depth to 12.7meters.

(area Bl.thered conglomeratesubstrate was gently'slopi nghaving~an

an~le

^of

incli~ation

^{of 310tothe}

horiz~.nt·~l .

^{Asimilarlyslope'd}

area having anangle of incli nat ionof 28⁰was'also presentinthe 20.'5.to 24.0meter depth range,(area D). Between thes eqent'ly, slop i ngareas (,Le-.12. 7to20.5met~r.r;a~ge ) .the bottom was more .steeplyinclined(54°) (area C) and asimilarlysteeplysloped

(51°')area was present between20.5meters

~nd

^the

s-~nd

^{bottom at}

'i

31

nw:te.r

^{s (ereeE). ,}

. '1'

This patter nof alternatfng.steeplyslopingandgentlysloping

.t-'.v', ., , _.~ _ _.J.. • .' . • .'

areas is showninFg.11; a crcss-secttcn ofth.ecliffas mea.surett, a~ong_the TL. The slope and depthof'eachTl Qisshowni~ Ta~l~1.

The widthof the study-areawas detenn i nedby horizonta l

.boundari es·Of.each equal lyslcped region. 'Thebo~ndary

of

zones 1ncrea'-sed

~

· j

-,'- ,' ...": : ..' . " "."

.·..F1·gur~11: Ltned~awtngofthec1'1ff-praf tl e show.tngtheloca~1on~Of:

T~Q·S~.

^{Verticalseale}

t~

ⁱⁿ

· me"ters.: ~.

...

.

...:

...

•"l""

.• L·

:.'.'