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(1)

AN ECOlOGICAL STUDY OF EPIBIONTS ASSOCIATED WITH THE

SHEll Of THE SEA SCALLOP, PLACOPECTEN MAGELLANlCUS

(GMELJN, 1 791) CMOLLUSCAPELECYPODA)

PERMISSION HAS BEEN GRANTED FOR THIS THESIS TO BE XEROXED WITHOUT RESTRICTION

DAVID HUGH BARNES

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AN ECOLOGICAL STUDY OF EPIBIONTS'ASSOCIATEn W!TH THE SHELL

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OF_-THE SEA SCALLOP • PLACOPECTEN

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f:J!!~ELLANICUS

,.. (GMELIN, .1791) (MOLLUSCA: PE]jEqYPODA) · ' - .7

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Submit~ed to

The Departm~nt of Biology

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B'-emori~l Uni~ersity

1 ·of

N~wfoundland

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In Partial Fulfillment .. of the Req~iremetits fo;~ the pegr'e~

Doctor .of Philosophy

·' . ...

. @. David ~ugh Barn.e~, B.Sc.;

M.sc:

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"' Marc}1, 19.74

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ABSTRACT

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epibiotic organisms associated ·with the shell

~f t~e ~~" · ~callo~,.··

The obJect of the study was· to. describe t.h~ ~ssemblage of

n

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·P~aopeaten magelta~icus

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' '(Gmelin, 1791). Qualit<it'ive-·informat~on from appro.xima tel'y

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3 do.zen 'shells was combine_d ~ith ,qtiantitative data

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obtained from

12 . a'dd{t;Lo~'llhetls.

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,-to· statistida:.t;·-:-inalyses

in

order to

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~e· n~edca? ·· ~ata were . ~~~~ected~

examine dqminance, diver.sit;Y and

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...

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affinity withi-n ~Jle a~sociation.

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Live sc;~ll.ops •were ·~ollected

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by divers ·from an area in &t.

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Mary's Bay,_ ... on·the .sa'utheast coast of the·-~·sland of Newfoundland • . T~

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external .surfaces of the shells were examined using a dissecting , .

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· microsco~~ · ~nd_: ~h~ area' o'?d.1pied by each· ~p'ibiotic speci,es was

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mea~ured.

y

X~r~diography

was used to investigate the effec'ts of 'boring

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spedies within the shells. A total.of 74. epibiotic species of animals · and plants was recorded. '

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The., affinity ·studies revealed. tlult the scallop shell supports .

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an epibiotic community consis~ing of separate and. distinct component . .

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co~unities -on its ·upper and-lower ,-81,ves, each having a predictab.le ''.

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specias. composition. No seasonal._or depth..:related effects wl'!re

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detect~d

.:regarding the .

·c~mmunit'y

:'d'n adu!t .

scallop~

over-10 years .of .

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_' ~ge,: bti_t: the

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structures 0~ /1 the associations on young scallops l

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and

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were diffe~ent· itf. several reSJ;IeCt's from the adult shell ,.

· .·co~unity.

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Th~ dominant spe9:i:!'!~ are sessile or sedentary_· in hab~t ·'and are

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either filter ... or deposit- feeders.· They .are not obligate members of' th~ commu~ity,, but nevertheless'reproduce and ~n some c~ses ~ complete

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the~r entire ~ife cycle within the community. Diversi~y -indices

'obtained for _the

~ommdni~y · ~er~ ~e~erally

low (ranging

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for a ~}n~le valve) and were lower for ~pper valves than f

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or lower valves.

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ijo~ing.species play an important role,in the development of the

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co~~nity. Their actiyities modify the surface.

of

the snell, making it

more habitable fot certain later-arrivi~g species. Damage to the· slie'll

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is rarely extensive and the scallop'do~s not generally appear to-suffer

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any harm from the pre~ence.of these or any other of . the-epibionts •.

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. 0 ACKNOWLEDGEMENTS

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Financial ~upport

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for this study was provided from within a

from the National Research Council of Canada •. I

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am grateful to D'r •

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~vans also· for $Uidance and e courage'ment throughout -the program, and ~ "!

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for crit'ical reading 0~ "the IWilOU '

Dr. G. I-. MeT • .' Cowan and ~ C. C. Davis also re.ad the manus- cript and· ·offered valuable suggestions for its improvement.

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following

per~~ns peic;o~m~d d{~i~g

se:Vices· at various titpes and were g~n~r.ally helpful ~ith the ~el~ ~~~: Barbara Fowler, Robert Smith, James Osborne~ Jonathan Moir and Neil Bannister. ·

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c.--s.

Andrews and J. Bartlett gave1much. help. with the writ~ng

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of the computer program .for the· analysis ·of the numerical data.

..

,. Techni-cal·.assistance in the photographic proces~es involved in ·

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the preparation of the f.P,gures 'lias given by Roy Ficken and Graham Hillier.

Mrs •. P. BetU!.ett of Technical TyPing2:-typed the figa~~anuscrip't

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~nq is therefore largely responsible for.the~appe~rance .of this ~hesis.

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Taxonomic Specialists Q

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The au~hor is grateful for the assistance given by the following (

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persons in the identification

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certain 'of the associated, ap~cies • .. : T~e"' numb.ers correspond to· those in' squar~ bracket9 following the particular

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.. spe~ies, -listed in Table l .

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Protozoa: Ciliata: . ·,; '/

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/ •Jl. Dr. ·J. B. Lackey, Melrose, Florida •.

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Proto~oa: · Foraminifera:

2. Dr. G. Vilks, Bedford Institut-e of Oc;:eanography, ·~

Dartmouth, Nova Scotia.

Cnidaria: I . . . . ' < J

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3. Dr. D-. R. Calder, Virginia Institute ·of Marine Sd:ence', ,.

.

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Gloucester Point, Virgi?ia.

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4. Dr. K. W. ;Petersen, Copenhagen Vniversitx Zoological Museu'!ll•

Aschelmfnthe~:

Nttoda · . , ~

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MJ:'.

R.~ H. Mulvey, Agriculture Canada • Ottawa.

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6 • . Mr. B. E. Hopper, Agriculture C~nada, Ottawa.

An~elida ~{,olychaeta:

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, 7. Dr,. Mary Petersen,, 'Copenhagen University Zoological Museum.

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Mr.- J.

Q~b~rne, Can~da

'fi·epa.rtment

~f.,

the Environment,.

·St. John's, Ne~oundland.

Mollu'sc,a:

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I. Smith, ~atiomil f:1useum .. ot' Natural Hi'st~ry~

9. Mrs. M. F.

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~ Ottawa.

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10. Dr. 'K. 'B •. Clatk,' Florfda Ip~titute of T'echnolog,y, t-;elbourne, Florida.

Arthr~poda:, Crustacea:

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' 1~. Dr_. E. L. Bousfi.eld,. National Museum of Natural Sciences,

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Ottawa:

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12. Dr.

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T. Shih, National Musetim •of:Natural Scien'Cea,

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Ottawa.

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13~ Dr.

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Lindquist·, Agri~ulture Cana:da, · Ottawa.

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14. Dr. T. J. M. 'Schopf; Department

'of

Geophysica~ Sciences,

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University ~f Chicago.

Chordata: Ascidiacea:

15. Dr .. N. 'J .. Berri.ll (retired).

R~odophyta, ·Cyano_phyta'-:

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16.· Mr. R. Hooper,- ·.Department of Bto~ogy,

Memorial' Un~versi ty $Of Newf~undland.

17. 'nr. A; Whittick, pepar~ent of Biology,

Memorial University o.f Newfoundland:

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ABSTRACT

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ACKNOWLEDGEMENTS ~ t

. ·, LIST OF TA:BLES •

·:,

LIST OF'FIGURES

~TRODlJCTION

. . . .. . . .. . . . . . . . . . .

. . . .

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. . . . . . . . . . . . . . . . . . . . . . . . .. . .

. . . . . . . . . . .

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MATr.;RIALS AND METHODS.

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Field Work i .•

. . .

Determination of the Age

of

the Scallop

. . . · . . . .

Qualitative Analys'is ..

· . . . . . . . . . .

~ Quantitative.Analysis

. . . . .

~

. . . . . . . . . . .

Analysis of·Data

. . . . . . . . . . . . . .

DOm.inance

. . .

.

. . . . . . . . . .

Divep~ity,

.. .. . . . . .

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Affinitie' between Assemblages

. . .. . "' ' .

RESULTS AND'OBSERVATIONS

. . .

.-)

. . . . . .

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. . . .

.

. .

. N~tural History •.

.

Spatial Dominance

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Diversity

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A~finities between A~~blage~ • • • • • • • . .

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~,i?elity,rVitali~y an~

Periodil. it . • • • ••

DISCUSStON · • • ... •. · • • . • • • • • , . • • . • •· • •

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Tqe Scallop Sh~ll_Surface: Na ural History

• • • •

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Species.Present

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Domiqance

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Diversity ·-

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Evaluation of the Present Study The PZacopedten Community. ~

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Future Directions for Research SUMMARY

. . . . . .

.

REFERENCES CITED • • •

APPENDIX 1 • • • •

.

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I, 'APPENDIX 2' • • • • •

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_Page 122 125

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• • 138

141

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TABLE -1

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LIST OF TABLES

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EPIBIOTIC ORGANISMS RECORDED' FROM SHELLS'OF PLACOPECTEN . MAGELLANICUSJ FROM SALMONIER • ·• • • .• ; ... • • • • ~ • • •

X

Page·

40 2 DATA

FO~

.12

scALLOP~

usEJ; iN

·Q~ANTITATIVE ANAt~srs

oF\, -· .

ASSOCIATED EP1BIOT-IC OI{GANISMS • • • • .• • . • • • • • • • 51

~.1 ."'tOTAL AREA, AREA OCCUPIED BY ·EPIBiO~TS, PEl\ CENT

AREA

OCCUPIED, AND NUMBER . OF EPIBIOTIC SPECIES PRESENT. . DATA FROM uPPER .VALVES,. SUMMARIZED. FROM APPENDIX .1,

~ABLE A • • • • ~ • • • • . • • • • • . • • • • .• . • • • • • · . ·53

·

..

3:2 TOTAL

AREA,

AREA OCCUPIED BY EPIBIONTS, PER CENT

AREA

· OCCUPIED, AND NUMBER OF. EPIBIOTIC SPECIES PRESENT.

· .. DA1A FROM LOWER VAtV,ES, SUMMARIZED FROM APPENDIX 1, TABLE· A • • • • • • • • • • • ·, • • • • • • •· ~. • • • , .

.· ~ ··

~

ACTiVITY OF BORERS J;N THE SHELLS. OF TilE 12 SCALLdPS , ,, • :

LISTED IN TABLE 2 • • • • . • • • • ·• • • ·• · • . • • • •. ~ • '56

' ,J - r "' ' ~ .

5.1

SPAT~

DOMINANCE. TEN DOMINANT EPIBIOTIC SPECIES RECORDED FROM THE ENTIRE. SHELLS (BOTH VALVES CONSlDERED

TOGETHER) OF THE 12 SCALLOPS LISTED 'IN, TABLE 2 ·• • • • • 63 5.2

~

5.3

5.4

. . . )..1

5.5

. '• . . . ' ~ ~

. .

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SPATIAL DOMINANCE.· TEN DOMINANt EPIBIOTIC SPECIES RECORDED FROM .T~ UPPER ·vALVES OF THE 12 SCALl.OPS ..

LISTED IN· TABLE 2 ~ · • • ., • .;' • • ~ .~ • • ~ • • .•. •.

. . .

. . ··f"'t . . . . ' .

SPATIAL DOMINANCE. TEN DPMINANT EPIBIOTIC SP~ClES

RECORDED FROM THE LOWER VALVES OF THE 12 SCALLOPS·.

LISTED IN ·TAB~E 2 ".. •. •. • • • . ~ • ~ -.. • ~ • • • • • •

~t . . . ·~ , (} I

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SPATIAL DOMINANCE. TEN .DOMINANT· EPIBIOTIC -SPECIES RECORDED FROM THE UPPER VALVES. ·oF

nm·

5' SCALLOPS. · INCLUDED Ilr~HE "BAsE" GR9UP ·

e'OLD"

/SUMMER/ 6', ~T.ERS) , AND NUMBERED ~ THROUGH 5 ./ ~ .•" ; • ~ • • • • • : • • • •

. .

SPATIAL DOMINANCE. ,TEN DOMINANT EPIBibTIC SPECIES.

·R~ORl>ED FR.OM.

THE

LOWER VALVES' OF

rim

5 ·sCALLO~S ..

INCLUDEl:>. IN THE "~SE" GROUP . ("OLD*' /SUMMER/6 MET~S), , AND NUMBERED ·1 ~OUGH 5 • • • • , • ~ ·. • • . • . • .. ~ ~. ., •

.

. 5.6 SPATJ:AL

~OMI~CB.

TEN :OOMrnANT EPIBIOTIC

S~EciEs

' .. RECORDED FRO THE UPPER VALVES ·oF THE 2·SCALLOPS · TAKEN At A WER DEPTH (20'METERS)·, NUMBERED 6. AND ·j •• .

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. -

" ' ::..·

. '' ~

I

~

...

.··.·,

..

.

.

. It:

(

.. .

I : .

(15)

I

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.. ~ ...

I

..

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.

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:1""

't

. '

.

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.,

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TABLE

.. a 5.8 .

I

- ·

I ,

.§ :' .

, ' I

' r •• lJ. " ' '

- . . . ...Q '

SPATIAL DOMINANCE~ TEN DOMINANT EPIBIOTIC. SPECIES . RECORDED FROM THE LOWER·.VALVES OF THE 2 SCALLOPS · TAKEN" ~T A.LOWER J?EP'lil (20 METER'S),' NUMBERED 6

AND

7 .

'

.

SPATrAL DOMINANCE. TEN. DOMINANT. EPIBIOTIC SPECIES 'RE(CORD~D FROM THE ~P_PER V~I.VES OF THE. 2. S~AttOPS .. .

TAKEN. "LN ·THE. WINTER Sf:4~0N (6 METERS DEPTH) 1 . -

~-g~ 11 ; ~ 12 • ~ . • • • • • : ' • . • • • • • • • . •.. •

5.9

SPATIAL DOMINANCE • . TEN DOMINANT EPIBIOTIC ·sPECIES RECORDED 'FRoM.il'HE LOWER VALVES OF THE 2 SCALLOPS

. . I · r 1

T~N IN THE WINTER SEASON (6 METERS DEPTH),

I .

. • .

. ~~ED . 11 AND/12 • •. ,; ,. • . • ' • • . • ~ • • . . • . • • . • • • I

,. . ' . n,.

5 .10 . SPATIAL DOMINANCE. TEN DOM~NANT EPIBI.OTIC SPECIES

Page

I

73

~-

·' 75

, ~ECORDED

J'ROM-

THE UPPER \"AI,VES OF

TH:E

3 "YO~G" ,, :. ·-·

I ' ~CAi.LdPS (SUMMER/6 ·METERS)' ~ERED. 8, 9 AND 10 • ,. • • ·"' 78.

· .

...

Q 1

. .

•\

. i

·~

. ·~

<Iii ', .

"

'

.. '

• ' \ I·.

>' I

, ,..

.

. . ' "

.:·- 5.1,1 SPAT~ DOMI~CE. TEN DO~N~ ,EPIBIOTIC SPECI§S . . . . . . RESOIU>ED FROti piE LOWER V~VES OF THE 3 "YOUNG" . ..

·• · ·· . :'SC4J.LOPS (SUMMER/6·METER~), lilJMBE~. ij, 9 AND 10, • • : • •. · . 79. ·

..

.

,"

. ..

.,

.

.

' '

,, .\

·. r

-:

D-I~ERSI'fy . IND~CES

FOR THE EPiBIOTiC

·~ASSOCIATIONS

ON

~-

.

·.. THE SHELLS OF THE . l2 SCALLOPS LISTED IN. TABLE 2 : . · .

6

. ·.

B~ 'VALVES • . ~ • • • • . • . • . • • • ~ • • ~if"l:

· • • • • • • •

. . . . ·. . . · , c:;.J . 'y •• 87

7 ·· :DIVERSITY INOicEs FOR

THE EPIBIOt~c A~CLATIONS, ~N .

~ THE SHELLS OF THE ·12 SCALLOPS LISTED tif·TABLE 2: '

. ·_,'.BY R~GIQNS • • • • ·. • • • . • • • • • . • . •. • •• •• :. ~ . • • • • .•.

r9. • '. • . . 91 . '. -~ . . : . .

_, .

a · ·

.·.COMPARISON OF EP,IBIOTIC· ASSOCIATIONS:.

HITRA-SCALLOP· ~

. ; BY V~VES (UPPER . .VERSUS

LOWER). ·SCALLOPS

NUMBEREo \\

· l

THROUGH 1;2, AS LISTED IN TABLE 2 ·• • • • • • • • • .. ~ , 95

' ; L o \

9 .1· ·. ,

&OMPA~USON ~p.

EPIBIOTIC

ASS~IATIONS

I~ER-scXl.LOP,

.. BY· VAL,VES. 'BA~J(~OUP OF 5 SCALLOP~ ("OL~?/SUMMER/ - . 6

:§Et~S)

, NUMBERED 1

~OUGH

5 • ;, .• • • • • • • • • •

.• '~ .

.

' . :

.

97 ·,

. .

9. 2 . COMPARISON 0]!. EPIBIOTIC ASSOCIATIONS:· .. UfiER-SCALLOP, • . BY VALVES. :. BASE GROUP (NUMBERS 1' TO •5) VERS,US 2 . ·

'ta f I•' .,' ',

-~

1 S,CALLOPS NUMBEJU) ·6

AND

7) FROM LOWER . D&FTii (20m) ;, ~ . . · • •

.

· . 98- . 9. 3 : I COMP~IS OF EPI~IOTIC ASSOC~TIONS: • INTER-SCA40P ~

~y VALVE • . BASE .GRoUP (!WMBERS. 1'To ·5) VERSUS' 2· .·: i'( -• .

{I L ·SCALLOPS (NUMBERS 1l: AND i2) TAl{EN IN WINTER-, . • • f)· . ·.• :·.99

t' ' ',I ', ' ' ' o ~\ •-~; ..- I

9. 4 . CQMPAR.ISON

OF . E~IBiotiC . ASSOciATIQN~· : }NiER-;SCA:LtO~

,.. . ; ·."' . 1

-·-~

· Bt VALVES." -BASE GROUP (NUMB:ERS 1 TO 5) ·VERSUS 3 · •·

.. . , ··.vo~G

·sCALLoPs (NUMBERs

.a, 9 ~

10). · • • . • •. ·

~

· ••. •

~,

· .. . - .-:

100 ·' -:....

.

.

~- .; .

'

.

.

~l ••

.. '· ' I

.

. :

..

. : ,,

. .

:

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• ! p

.

. .

.. . . ..

, ' -.

-~;

""~("! 1 .~

.

1- '

· ... ....

.. .

, .

.

.

'

.

·~ . :

. ; ·. ·~· . . . . · . : . ~ · ' . .. , : '; ~ .... . ... ,: : ; . . . . -~ ·i<,j~

(16)

(

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. .

••• f

"

0 :ldi

..

- J

-.. ' . I

TABL~

-~ 1'0·

t

.

~

..

J '•

AVERAGE INDICES OF AFFINITY, •c,AJ.CULA~ DATA

: IN TABLES 8 AND 9 •· • ·~ • • • • •

i ·. . . . .. .

. ·

COMPARISON

?F

EPIBIOTIC

.

A~SO~IA~IONS: .

..

"· INTRA-SCALLOP,

.

' Page 101 '

I. BY REGIONS • " • • • • • • •. • . - • . . • • • . • • :. • • • •.•

. .' "" 0. . ,.., i05

12 · , FIDELI-TY, VITAL:J;TY, ··-PERIODICITY AND FEEDING TYPE ..

·.' 1 (OF ANIMALS) FOR· DOMINANT SPECIES IN THE PLACOPECTEN

"- \::...co~ITY. SPECIES ARE .RANK.ED BY .lf.liEIR BIOLOGiCAL . INDEX VAL'1JES ," IN THE ORDER. IN WHIC1i THEY APPEAR :LN

TABLES 5 2 AND 5 3 • ..•: · · - 107

. . ·• • t.

', I / /

1

t h

EPIBIONTS RECORDED ~OM; THE SHELLS OF THE TwELVE A '

.. -

SCALLOPS LISTED. IN.TABLE _2~ (PER CENT ARE4.0CC~IED;

PER CENT OF TOTAL AREA .OCCUPIEr¥ BY EPIBIONTS: NUMBE)l . .. OF INDIVIDUALS I~ GIVEN ONt~.;fflEN ~ERS OF. SPE~IES

ARE COUNTABLE) . • • . • • • • \ •. • . • • • • • • • •. • .; • • 145

0 '

A.1 »:PI.BIONTS RECORDED.- FROM SCALLOP N~ 1, UPPER VALVE • 145

.

. .

A.2 EPIBIONTS RECORDED FROM SCALLOP NO. 1, LOWER VALVE. .•. • • 146 ...

.

'·. . () A.·3 '· EPIB,IONTS RECORhED ~E>M SCALLO~-~0 •. 2, uPPER VALVE •

1.• ~ '

. .

147

.

,

..

, A.4 'EP!BIONTS RECORDED FROM SCcUl;OP NO. 2,_ .~OWER VAL_VE'.

. ..,

148

X.5 · ·

EPIBIONTS· REcoRDED

FRoM·. ~GALLOP

No. 3, .'

trrPER

VALVE

. ~ . .. .

. .

·• 149

~.

' . .

I I .

A.6 . EPIBIONTS RECORDED FROM .SCALLOP NO. 3, LOWER VALVE 150.

.. .

''\ '.

,!\ '.

~

• .

'r

}) . .

.

. · 5 ~

:. r~~~ . , • ..

"

. . .

. .

'

•' I

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. . .

- ? ·' ~ .. c ~*:; p

A~7 ~ EPIBIONTS·RECORDED ~ROM SCALLOP·N0.'4, UPPER VALVE

. . ....

a .• A.8

Er~BIONT~

REC_ORDED

FR~M

scAL\OP NO.

4,

LOWER VALVE

.

• •

:>

A.?

EPIBIONTS RECORDED 'FROM SCAL~OP NO. 5, UPPER VALVE •

. .

I '• ...

A.10 EPIBIONTS RECORDED FROM· SCALLOP NO. 5) LOWER VALVE

':

.

.

·' A.l1 " EPIBl!ONTS RECORDED·' FROM· S~LOP .. NO·. I

6,

UPPER 'VALVE

. .

"

A.12 EPIBIONT~RECORDED FROM SCALLOP NO. 6,· LOWER·VALVE • • •

''\ ' ' .. ' . c...: . . •. ; .

A.l3. EPIBIONTS RECORDED FROM SCALLOP NO. 7, uPPER-V~VE ·• • •

il .1

' A:- 14

EPIBIONTS RECORDED FRoM .SCALLOP NO.

0 . . . '

7, LOWER· VALVE .I .; • 8,

UPPER

'vALVE • • •

. .

.

. -

EPI~IONTS RECORDED<~~ SCALLOP.NQ.

: . . '/ ';. t . '

8, LOWER. VALVE · •· • •

·- A.1CV

' '

EP:IB:ON!S ~COIWED ~OM ~~LOP NO.

151 152 .. ~

. J.53 (154;

~

155 '

156 15.7

'

_r::~5a

-t6o

161

~- ·

~ ..

. .

. ""' .

.

..

. ....

...

. ~r

...

7

0

'I

·, .r : ' .

•'

I·

.r

...

. .

i . .

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..

: . . ,. .. ... :.!-···

(17)

>

:~.

..

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; / ·

. <;!

...

-· . ~~

. 0

,.

.

(

TABLE

9, UPPER VALVE

.

. .

.

/·A~

11.( EPrBroins

~coRDED

FRoM sCALLoP. No.

~ ,I

A.18 EPIBIONTS RECORDED FROM SCALLOP NO.

.

9, LOWER VALVE~-.

" t' A.19

k.20 A.21

(1 .

·; .· A~23

I

ol

.

.

_A.24

.. .

....

EPIBIONTS RECORDto FR9M SCALLOP-NO. 10, UPPER VALVE EPIBIONTS RECORDED FROM SCALLOP ~0 •. 10,

EPIBIONTS RECORDED. FROM SCALLOP NO. . '

. .

11,

EPIBIONTS RECORDED FROM SCALLOP NO. ll:,

...,;,,

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0

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t; .

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.

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-- . -

'":.' '

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·-

0

·

.•

LOWER VALVE UPPER VALVE .

, ...

.

~OWER VALVE

.

.

'

. . ·1 ..

. ~ ~~.

. I

xiii

Page

.

~ ·162

,.

.

'

.

163

. .

, ~ 164

,, '

. '164

J.J .. 165

• • 166 .

. . .

167

. .

168 I

J/

I I . 'Ia • •

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·. ~

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·.

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·.. . ' { .~·

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