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 ·ofN~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::r t
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~~ .. . . j' .t. t'
~e· n~edca? ·· ~ata were . ~~~~ected~
examine dqminance, diver.sit;Y and
• ..
...~f' .
affinity withi-n ~Jle a~sociation.
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Live sc;~ll.ops •were ·~ollected
. ....
· .:'1', . • .
.. A' 'd· .
.
.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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~ r '.' • , : ... ·. . \'-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.
yX~r~diography
was used to investigate the effec'ts of 'boring' • , 0 ' i' ) A
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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 .'' ' • • I: '
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_' ~ge,: bti_t: the
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structures 0~ /1 the associations on young scallops • • • l
(2
and::.·_.):···~~'fa_ . ~ld)
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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-f ~~·,,Q.J ' ~o . J.~
for a ~}n~le valve) and were lower for ~pper valves than f
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or lower valves.I'
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 itmore 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 afrom the National Research Council of Canada •. I
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am grateful to D'r •.;'.••
~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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followingper~~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. ·' . .· . . 'y
c.--s.
Andrews and J. Bartlett gave1much. help. with the writ~ng•' "'
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\ . . . t • I) ~
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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 · . , • ~·· 5.
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R.~ H. Mulvey, Agriculture Canada • Ottawa.' .
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,I
Ottawa.
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13~ Dr.
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Lindquist·, Agri~ulture Cana:da, · Ottawa.Ectop-rocta:
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14. Dr. T. J. M. 'Schopf; Department
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Geophysica~ Sciences,~ a
University ~f Chicago.
Chordata: Ascidiacea:
15. Dr .. N. 'J .. Berri.ll (retired).
R~odophyta, ·Cyano_phyta'-:
' '
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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MATr.;RIALS AND METHODS.
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Field Work • • i .• • • • •
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Determination of the Age
of
the Scallop. . . · . . . .
Qualitative Analys'is ..
· . . . . . . . . . .
~ Quantitative.Analysis
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~. . . . . . . . . . .
•Analysis of·Data
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DOm.inance
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Divep~ity,
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Affinitie' between Assemblages
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• • • •RESULTS AND'OBSERVATIONS
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. N~tural History •.
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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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'viii
Page ii iv ix xiii
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Domiqance
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Diversity ·-
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Evaluation of the Present Study The PZacopedten Community. ~.
•Future Directions for Research SUMMARY •
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•.
REFERENCES CITED • • •
APPENDIX 1 • • • •
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I, 'APPENDIX 2' • • • • •
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_Page 122 125
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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~
.12scALLOP~
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 CENTAREA
· OCCUPIED, AND NUMBER OF. EPIBIOTIC SPECIES PRESENT.
· .. DA1A FROM LOWER VAtV,ES, SUMMARIZED FROM APPENDIX 1, TABLE· A • • • • • • • • • • • ·, • • • • • • •· ~. • • • , . 4·
.· ~ ··
~
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 CONSlDEREDTOGETHER) 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' OFrim
5 ·sCALLO~S ,· ..INCLUDEl:>. IN THE "~SE" GROUP . ("OLD*' /SUMMER/6 MET~S), , AND NUMBERED ·1 ~OUGH 5 • • • • , • ~ ·. • • . • . • .. ~ ~. ., •
•
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. 5.6 SPATJ:AL
~OMI~CB.
TEN :OOMrnANT EPIBIOTICS~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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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
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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 OFTH:E
3 "YO~G" ,, :. ·-·I ' ~CAi.LdPS (SUMMER/6 ·METERS)' ~ERED. 8, 9 AND 10 • ,. • • ·"' 78.
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. . ' ".:·- 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. ·
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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.
EPIBIOTICASS~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 ·' -:.....
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TABL~-~ 1'0·
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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.
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, A.4 'EP!BIONTS RECORDED FROM SCcUl;OP NO. 2,_ .~OWER VAL_VE'.. ..,
148X.5 · ·
EPIBIONTS· REcoRDEDFRoM·. ~GALLOP
No. 3, .'trrPER
VALVE. ~ . .. .
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·• 149r·~.•
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I I • .A.6 . EPIBIONTS RECORDED FROM .SCALLOP NO. 3, LOWER VALVE 150.
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A~7 ~ EPIBIONTS·RECORDED ~ROM SCALLOP·N0.'4, UPPER VALVE
. . ....
a .• A.8
Er~BIONT~
REC_ORDEDFR~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
•
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·' 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
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161
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9, UPPER VALVE
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11.( EPrBroins~coRDED
FRoM sCALLoP. No.• ~ • ,I
A.18 EPIBIONTS RECORDED FROM SCALLOP NO.
.
9, LOWER VALVE~-." t' A.19
k.20 A.21
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._A.24
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EPIBIONTS RECORDto FR9M SCALLOP-NO. 10, UPPER VALVE EPIBIONTS RECORDED FROM SCALLOP ~0 •. 10,
EPIBIONTS RECORDED. FROM SCALLOP NO. . '
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11,EPIBIONTS RECORDED FROM SCALLOP NO. ll:,
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xiii
Page
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• • 166 .
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168 IJ/
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