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F-lot K n t-ftnK r F scrssFie l)92 

INTRASPECIFIC SHEEI. ARIATION IX THREE SPECIES 

OF ROCKY SHORE GASTROPODS FROM HONG KONG: 

CORRELA [IONS AMONG HABI IA IS AND 

A (OMPARISON 1’ UI H ‘I EMPERA [F SPE( IFS 

A Richard Paltrier 

Department sit iooioe’ 1 ni\elsit\ of Aiheita, Edmonton. Alberta T6G 2EQ 

and 

Bamf Id Marine Station, Bamfield, British C thimbi i VOR IBO 

Canada 

4BS TRACT 

lo det rrninc if suf ropieal ga tropods exhibited magnitudes and patterns of intraspecific 

morphological sa iation comparable to those of m re thoroughly studted te riperate 

spec ie. I col Ie ted three specics of rock’ hore gasri opods born a total of e ie en in 

teitidal sites along the shores ol Hone Kong Tsso species ssere predatory muric ids 1/otis 

nit i ,o and T 1 0re \1s>11,o i and the third \ cis a common. herbis 01 OUS troch id 

lIt’iit c/SOt/U ‘ji’t I e\anlined Ii aits sensiti\ c to pollution Ics eE (penN sI/Ct ti aitc 

relate to desiccation iesistanccfsusceptibility to dislodgment (aperture area projected 

area, shell capacity I and traits related to predation intensity/resistance (number of re 

paired shell injuries apertural tooth height. lip thickness development of shell sculpture. 

shell weight and occupied \olume) 

For both Ps1it fast ‘ro a and T leuft ofuma masc uhinized teniales ss crc rno C 

common at C ape d’Aguilai than at any other site. pcrhap because ot its proximit\ to 

major shippine lanes. \lore samples ilong Hong Kong’s southern shores ssould he re 

gun-ed to suh’tantiate this pattern F-rn traits xnhcie data could be compared quantitati\ ely. 

both T lot tdera and Wi,nod nit la/no exhibited ma6nitudes of shell s ariation com 

parable to those of tei iperate species The trcquencs of repaired hell injuric also 

compared favourahls with published values for rocky shore spec es from the \orth 

Atlnti Hence H ng Kongca 

5trcpods did ao app a tss he unusual ‘a cithr rc pcct 

Of ‘ome intcest. he rclatis e variahll1t\ of morphological traits sas stgniticantls oi— 

re ated hetss CCI1 I lot i in to and If ia/an, traits more ‘ ai iahic ci 1 10,1tt it al ‘SI 

649

650 A. RICHARD PALMER 

tended to be more variable in 34. la/no, In addition, the species found oser a broader 

range of sites 1. /aciecra> v as most s ariable. shi1e the species found at the fewest 

site, T. leiIteasroma) s as least ariable. ss ith 11. labia in betseen. 

L sing regression techniques. I compared shell traits both among populations and 

among species. \perturc si/c exhibited the most striking s ariation among habitats: for 

both Thais lavii,’e,a and 1on’ dotzta labia, populations from more wase exposed shores 

had proportionally larger apertures. These taxonomicallx and ecologically distant spe 

cies thus exhibited similar morphological responses over a wave exposure gradient, 

Somes hat surprisingly. I found little cs idencc ot correlated responses to s ariation in 

predation intensity. In both species. the relanse development of defensise attributes \as 

not significantly correlated ss ith shell repair frequenc among sites. In addition, the onl 

significant correlation of comparable detensise traits between species as a negative 

one thicker lipped 7. clasi’era were geneially tound on shores sith thinner lipped 34. 

labia. Thus wave action and predation risk did not appear equally capable of inducing 

shell sariation in these species. The interspecitic comparisons resealed that both T. 

c/al ii.era and 31. labio displayed a remarkahl similar eighi of shell per unit body 

ss eight and per unit shell capacity. Hence. in spite of their different sizes and shapes. 

these tsso species ins esied in roughly equis alent amounts ot shell to protect the same 

body mass or living space. 

INTRODICTft)N 

Most e\tcnsise studies ot intraspecific variation in the shell morphology ol rocky shore 

eastropods have focused on species trom temperate latitudes, mainly muricids (Kincaid 

1951’ Luckens 1970’ Phillips aol 1973 Kitching and lockwood 1974: Crothers 1985, 

Appleton and Palmcr 1988: Palmer 1990) and littorinids (Nesskiik and Doyle 1975: 

Smith 1081: Johannessin 1986. Scele 1Q86: Janson. I 9%”7>• A few have examined 

tropical or subtiopical species Struhaker 1968: Palmer 1979: Hamilton 1980: Wel 

lingttin and Kurs 1 Q2: Fni 1989e Coupled ith the much higher ds ersity and 

ri latis ely locr ahundance t tropical pccies i \ erinej 1978: 1987) one is left oh 

the iriprc s on that ampcrttc species arc tntrinsi ally more vriable morphologically 

t n trapcal ‘Ic adire c a u t’m. howes . ‘mi>ar saolpling intensity and 

cluanttiitis tcchniq es has e to be applied as er a comparable range of habitats 

Among rocky shorc nastropods exhihmting pronounced mntiaspecitic ariation. shell 

form c’rrc!ates most conimon>y ss ith the Iescl of ss atcr movement and the intensity of 

pi’daiion i’. shell-breakmg predators Kmtching r at. 1966: Struhsaker 196%: C’othcrs 

l98: Joha nessori 1986: ttrr 1 988hi Ithough it may also orrelate sith growth rate 

nd fxd a ilabmliey (\ermeij 1980: Well ngton and Kuris 1983: Kmp and Bertnss 

1 83 Ap eton and Palirer 1988> Ia determine sshether suhtmpmral gastropods exhibit 

romparable magnitudes and dir ciions ot intraspecific morpholoomcal s ariation in re 

S(iisC to a s ari:ihle ens irinineni. I condu ted a quantitative ‘tud at such varmaton iO 

threc specIes at rock\ shore gasiropds from the shores of Hong Kong: the prcdator 

IilCrc its The/I !a\ ., ii K tister and 1 ?‘o 6 arc ‘anna Hoiten. and the herhs orius 

ti cl id Won Santa alto L

Collection dtes 

JHAIS AND WO’VODONT SHEILVRIATION 651 

N4 4TERI LS AND MEl HODS 

Snails were collected during low tides from a total of eleven sites within use different 

geographic regions along the shores of Hong Kond between 12 and 21 pril 1989. 

I Cape d’ \guilar, exposed (CDI) 114 l5’43 E. 22 1302 N (approximate). Thais 

clas igera and Monodorna labia were collected from the upper 75 cm of the 

S’Iegahalanas sokano zone in cres ices and on shelves on a steep bedrock shore along 

the southeast edge of the cape. Because of high swell and a poor low tide, only the 

upper shore could be collected. I. kuteo stoma were collected from a high pool where 

they appeared to havc been thrown during storms as no food was available, and the 

shells appeared bleached ompared to those from CD2. This sample of I l’4tentf ma 

may not hase bcen representative of those present intertidally. 

2. Cape d Aguilar intermediate (CD2) 114 1 5’43’ E, 22 1302 N (approximate). 

T daiigcra I kutro.stoma, and W. lah,. were collected from the upper 50cm of 

the il’ft ç’ahalanus solcuno zone in cres ices and on shelses on a steep bedrock shore 

at southernmost tip of the cape. 

3. Ping Chau, exposed bc ch (PCI) 114 2C07 E, 22 32’22” N. I’ clavigeia and I 

leuteosk ma were collected at mid shore from among Tetradita on bedrock ledges 

and crevices at the southernmost end of the island, 

4 Ping Chau, high pools (PC2) 114 607’ F, 22 3222 N. T davigcia and I. 

lcutost via vere collected from among a dense cover of what appeared to be 

(‘hthamalus intermingled with articulated corallme algae on a bedrock ledge just abose 

water line of a laige t depool into which wave continued to break even during low 

tide Most snails were rapidly growing juveniles and young adults, and hence this 

sample may n t have r Ileeted the adult form in these pools accurately M. labto were 

sparse here, and only a few individu ls were found amone a rather larger number of 

mid ard upper ch re tide pools examined Most of those found w re small 

5 Hoi Ha Wan exposed HEI 1) 114 20’2( E d 8 51 N I c/as ig i a and 4,4. lahio 

were collected from he middle a id upper portions of thc Sac ostrca / me from 

bedroLk e vice , hould rs and pool at the north rnmost tip of Ko n iso Kok at 

the easter cdge of the rrouth of the bar 

6 Ihi H \ar, inte mediate (11W) 114 WOO F )2 S 32’ N T lasigcia and Vt 

lahi s crc olleeted I cm the middl f the Sa stra ione on boulders and cob 

1 s ak ng the soutf tern edge of ti e ir t r aj pro nor tors alor g th east hore 

d the bay 

7 Hoi Ha Wan pr tee ed HI-li) I 20( 1’ F 2 2804 N. I dasiçeia and 114 

lahi v ‘a. e )Ileeted Ito n the niddle and uppe portions of tie Sa coSt? ra /one 

rnor e rix d h uldr arid hdr ek alc ng Ic west ed of the sec nd ira 

pr non a o W thL ast horL f th bay. 

8 Hot 1 a Wai cry pro ta. cd HH4) 1 4 1 ‘ F. ‘‘ ‘7’47 N I do ‘i a.ia nd 

‘14 oh v iv o l ted from am n Saivostiva ml )ulders surrounded by sand in 

fr at the s 14 1 ‘Ia 

) H Sing War, espc ted 1181) 114 1444 F. _2 6t)) N I lcr cc a er 

coIl et dIr Tither idle and 14 / rvn akn th ufpc.r d .of liv. Sty cri 0 

mt. on a bedrock headlard proje tmf ou into bIt Harbour.

H Sm \Saaprc t 

cli dt )I t radii 

11 itht rtf’ alp 

tvec HS1 rdtF 

Ia TilT i. 

IL Vt 

Identificaf n of liters species 

; ii r u 

111 1 1 51 

t. (cdtkpatc i 

h v act cshihiDd co 

I ny a ong mai a 

lcrti ab a tiats lull 

ti tO era inc pr 

dj c it t hc utur 

e 

Prorta x mea ur me I 

ti chss c aidail ssci 

a ire tipfed s aDrpr H pc 

Ts treaurmert lid 

51(1 ai ii r at t It 

t irca oii ss n w oFt. 

cuing bru h and tHu hsorh 

RI H RD \LMFR 

‘ITS ii 14iHL,2D 2 N 1 ,i 

c 

anT 1/ rcia ncheupprecL. t i Co to 

1 pp siraist thd tI ‘. all h i 

Ha 1 the ay 

C 222(2 NO 

ft cud ‘flat 

ails t e u edt. 

tth’ H is 

tI cit 

o H 

\t . c ) 1 st’t s sps 11 cedly Fog 

Ott ci rt on ret sI I 

11 tilt lfl mfic as S p1 

55= 

5 51 ci mdi ih eq a iii cc rdisd c I the 

a I n al i thi aioI st t ey V t i ciucar ace at dc 

cp tI / d xl ic I rph ned iat tc ict lOs F pie 

a C dp ri Iacd p icr t ii tec a Or a Ii f 10 pes 

a a n 1ItS1 t cetdb 1 cC nt V I I t it) F 

r I tIll I ed eretak t F Ia, 

j 

I 

(C VO’ 

r 

t 11 is 

gc iyFd 

n th id 

1 10 d 1 (1 c to t 

a cc kr I tth told 

11 1 V I I 

Sr FIt F aidr ore r i’ni ii nd 

tal I ra raT ate I 

1 ( apr ret 

rim tF 00 rhi as r etc iF ti prdol H 

e Fcaaiatiw ics a Vg 

Ily II IC raT uals II u d tha 

h d ti u h d crc t c t Fl y ti 

r ac hetsse r o’ I re I te 

Firf Ia or a then rib tiache 

rtrvs krb 0 al seen earalla tiP 0 

th Iti rF d xhtl ted tss sman stia Itt r kr ) rlw 

I Ira r sr r ss c ru h i r xi xc c cp ‘ted xxi a 

ag tb’kq c h ç cthi.ntf jui ‘n’irc ii 

the c r 2( C of k H He ci alCoty ol F c’c cIt eters 

preseit a p cs Fe a iii rg at ire iid vidual o Icc ideiti tahi 

based to hell scuipturt and colout 

Shell handling xeights, solumes, sexing 

uhf 

ci a 

los If 

ais) Od a 

ud a 

icc 

c Al 5a oait en 

tidatdu Is aYbifu(u 

ruml n dthu 

a rw U ki F 

he ci ida 

10 1.001 

TI s a sr 

hc a 11 tI 

tIE (J 

‘aT 1)hc 

na ht lusI 

shcH vcr clcarcdrf n u tirnog tin a hit fly 

rdralintif nmnu Ire r rtt one ihllwh 

and coated e itl at oacr lat In It paver I b a ion, 

hc nad whdc thc anin als sre st I al s cht o th 

and th un copied I a If tIc sFelI It 1 ire Pr It 

al crcclaudla kiit,thcrslclls tI ft iiod 

r t tissue a presced I rn I up cain t the If rculum tr

11141S4%DVflOD . 4SHHI 4R14110N 6’3 

amie mW tra rawtr s iti kttttto u1. ttht lhad 

drd.ialserw°adcaiual,iic.c. 

L c.up dv lumccfashefl sr.e ie by, i.. ic.te rc oft tvite 

i d tohl tievo r dtheshellb t en F cp c.uuir o th lrvm.zsn i idthc 

pai t pr c. fa c.oid ..tt 11 v htof r Ith lalncev tied 

wit sic •tsoi 1)1 heac.dwt wul Ic ,.tdlrc.shvtrwa 

ttnitrc ucdntcti pe thaP iflett shi hldi ii .ox w r 

unit uru(ed C ncwa hldaprxiuatd Fat w ii ate c hub 

be 1 cribiis,t ijpedar undthto unc uptc va hc.ipc. ftheo na ,uli 

thdaw e t it Li tel T ill th too ii x.don hini tnod haL 

iI’ wit ti panccftiejc.rur patH t tesrfae fthe ethrgoar Adi 

ho lhel s’tc.rva i trod ted to i ip rtuc.Litltic. satert iredaflatsurta 

(ieitte nwed ncr sacg’d) lust sith th. out r lip rd tic. ventril sari il tie 

Iuirc.lIt \ the e relfat urac.e tin thet.dwe F otwtradde 

tc hc.aprtuc.y recorded ‘Wihc.u th r di ‘table nelilc.s t mi 

Paia IW 1nnuFc ttht.ppr Iunc.s rercpc’rtcdi raoffi shwar Thc.c. 

weght ma ec.orver flat alvclun sa taHow% 000m. 1 ml c. 

An dx. weiiht weie rip! ed rulssei pact ibolir. atu r era! 

nnuc. dt I F etat di cmthe 11 asic curv di c Sradsw r ad 

tithe I)pnis r I wind ) tlf’, Istalati Ttc.si, cttl pm 

iiy s i e tsn a .nc. ngtt a dd ie r a afrac ci 4 te 

rgttteiale. acre iFirtI a £ th . md as tc. ramc.o uedra 

& p1 t orcjj l cs ii ira at i ci na eratec tre i .cnad c. ci ‘ 

ULJL.di C hitlic. t4 4. 4 urn dgc’, i na 

Six. cadri Ian ht t iO (‘ii idn )vtnandleahedtoflienc.arest 

nc Pc. h& weiji a rl nea ur diwasubsairpcotaltlircc.sc. ies.Altei 

reinn th bcd tronth sIlo aboc,hcflslwadrtctoac.cnstantwightat 

6( ( nac ingot nflc.ween ‘id48h) Thest ubs mple ieIdc. veryac. urate 

rcducc. r ajm a i RMA) rear 1 sw it Y Iosjb dy dry w”i’ht)as a tunc.tion Cf X 

—lq( t net dbxly w tw &ht) hg 14 slop. andadjutc.dmens±siw cc. 

S t t Ia ii c. mad c.usicmctnicdc.Illrc.grtsontc.hnique) 

B thIla sçe p hi RMiscpe= 04( 0(2J) mcanX 2.92’ll 

i tt”IY fim mx 2NXOC 00DM (F ,— 1949,1

654 A. RICH RD PALMER 

‘ii 1000 

100 

50 

4000 

3000 

• Thai clavigera 

0 Thai leuteostoma 

‘ Monodonta labio 

•4 , 

300 500 1000 2000 4000 

Estimated body wet weight (mg) 

2000 98$ 

1000 

0 1000 2000 3000 4000 

Body wet weight + bnoccupied volume (mg) 

Fig. 1. Subsamples from wheh est rrates of A, body dry wmght and B to ci nternai 

volume of the tell ( hell capac ty were deterri ned to threo specees of gas 

tropods from Hong Kong See Ma enals and Methods for regresson equahons. 

dislodg bubbl It was lien pla d o a lump cf las with th p r ural p me parallel 

to he vu hing pci Cf Iiitl b lance and erraind r I tie ii tr I s un I lie 

witt wat r a icr un e upied v umc, Care was tak n nt L alh v y ci t di place 

vale in it ape s h r ooiti(n no the shell on the balance IThial irte ral slur 

aleulat d Is suhi aeO i. thc hell dry weichi lr( ii tFe s Light (I tIe Fell ill I tI 

frc h a e [he sub r p1 ci id I d er aceur c RM \ r gr si )r t )r Y tc tel 

nt a I olume s a uncti n t X estimated body ci weith oeeupi d rlum 

F g lB slope am I dju t d rr a Si) 

hr h h I a s pe u RMA sip 0)46 ( () ) 26) ieai X l83[ 

eseete I at n iX l69 0 0’ 1 (26, P J0)( I) 

F r Wi to (a a RMA sk pe 0995( ( 0 0l4) me ir X 580 

exe uc \ 0 near X 562.11 (05 (1 I I P ()(001) 

9 

9 

9 

0 

A)

TH4lc AND MOsODOI 4 SHELl ARl VI ION 655 

Total Internal solume for all remaining snails was estimated via these regressions Oc 

cupied olume was computed simply b\ subtractmg the unoccupied olume, which was 

measured directly, from the estimate of total internal volume, 

Shell morphometrics 

Ses eral morphological attributes of shells were selected for measurement because of 

their utility for inferring functional relationships. In the list below calipers’ indicates 

measurements recorded to the nearest 0. 1 mm using Vernier calipers, or to the nearest 

0,02 mm with dial calipers (lip thickness only) ‘Digitized’ indicates measurements 

obtained by superimposing, via a camera lucida. the image of a shell under a dissecting 

microscope over that of a calibrated graphics tablet (Mac I ablet Summagraphics Corp., 

Fairfield, (1, l SAz 500 x 500 DPI resolution) 

Shell length (calipers)— from the apex to tip of the siphonal canal for ‘I hais, or from the 

apex to the distal most margin of the aperture parallel to the axis of coiling for 

Monodonta lahio (repeatable to ± 0 2 mm). 

Aperture length (digitized) from the posterior margin of aperture at the suture to the 

tip of the siphonal canal for Than, or from the line described by the adapical margin 

of the aperture adjacent to the suture, to a point perpendicular to this reference line 

on thc abapical margin of the aperture, for 1 lahio [mean ° errors — sr (N 10) 

were 0 7 + 0,1 7 and 2,4 ± 0 38 cc for T clavitera and M lahu respectively]. 

\pe”ture ama (digitiLed) arcd described by the outermost margin uf th aperturdl 

lip and the polished parietal callus when siewed perpendicular to the plane of the 

apertuie [mean (f errors ± srrvi (N 10) were 1,8 ± 0,50 and 3.S t 0,96- for I. 

claiigeia and 11 lahio. respectively] 

Apertural t(oth height (digitized) the elevation of thc tip of the tooth perpendicular 

to the inner margin of the aperture when viewed roughly perpendicular to the plane 

of the apcrture (Ihans only. the tooth heights of M lahi iwere not reasured becausc 

the were consp cuous in all samples) When present the heights of the tw most 

abaxial teeth were masured In specimen where teeth were quite pro mnent these 

teeth were the second and third teeth abapicall from the suture’ the fourth and fifth 

teeth w re usually lareer hut were more difficult to oriert for accurate measurement 

Lip thickness (ealipers)—dn Thaii, I p tfi kn s was measurcd f om r point irisid the 

lip at the lo’ator of apertural teeth, t a point outsid thc lip lying bets een tlic two 

pro ninent spiral rows of knobs ncares the suture, t was mea ur d trm the tip ot 

the apertural tooth at this position f onc was freseit. Before condu ting statis icai 

analyses howevcr the heieht of tf apertural tooth was subtracted trcm his mcas 

ure of to al lip thick iess In 11 ab , lip thickness v as me sur d lrm a p )Int 

hetw r the third and fourth apertural te I adapical v f om t[ olu lla t a pc tnt 

bet see i the piral c 1 d on the oute serf ace f the sh 11 as perrend ular a p hle 

to the maicin of the aperture lip hickne s s a i pat lie t 006 r m 

Projeete area (digitized Projected area wa rnea urd a the total a a of tI srell 

vi wed p rpendicu ar to th axi of ci hug from the rieft i e nf the hell 

Sh Ils were attached t an adf s. e v rtical urfaee (duet tape a ta F ccl to a r an 

gular up )rnng block) with the axis of coiIin perperdicular to the line of sight

66 A RKHRE P\IMER 

rrnean (( errors ‘ ‘a ‘a = It ) w 1 1 )7 - r d I I lot [. lint ii’tii 

and M labio respectis l\ 

Knob heictbt dieitiied) Knob heiht a n sured as h des ation of the ttp of the 

knob perpendicular to the outer surface (I h shell ‘a hen s iev ed rouahf\ perpen 

tiiilar to ss hat ss ould hase been th. plan t he aperturL at the tine the knob ss as 

hein’ piciduced if /iul onls I ss 0 l rosh o etc n ,isured on each indo dual the 

highest tss o of th first foLir procc ti3g t’ m the apertural mat gin alone the 

ross adascnt to the suture Knobs in tnt’- ross are eanerailL more pronounced than 

those tn the other. mere anterior ross . 

Statistica’ anak ses 

Stattttsal analr ses ss crc conducted xs fib the micnsompueci statisttsai packages Stats less 

ii Vei 11>3: descriptise regressions and Super-\NOV-\ i Vet. 101 anals ‘as of 

cox ariance both from Abacus Concepts i Berkeley. California. f’S eSlopes from least 

square- linear reeresston IIR ss crc cons cried to reduced mami asis slope’- ithe R\IA 

is one form of model ii regression b\ di tding them hr the correlatton coeflis tent: the 

standard errors ot the slopes remain the ‘-arne LaBarhera 1 9h9 i. The standard errors 

for adjusteu means ae not preciseir deftned for model Ii regression. so to compare 

differences among means n the figures I graphed the standard error for the expecte 

Y at mean X as determined I torn the LLR CflaiX -its. To consti net the I igures ihat corn 

pare traits among populations. I computed ‘. alucs at a standard 5iie u’ine onir the 

regressions haed on indis tduals from each sPe as to loss 5: 

logi standardiied s r1tte { [log(standard X) 1 arnpk mean of X R\I \ slope i 

loge pected Y a - mean X 

Ihesc standardi,ed s alues were dctransfoi med pr or to graph ng. 

Statistical inference of significant ditf rences am tg populat ons ss as complicated 

fo most traits because populations differed not c r lv in the as cia e development of 

particuIai traits, hut also n their allomctiic relations e p c. idices i—iii Hence 

although ascrages differ nces wer cleary apparcnt among p p lations, the precise 

valu of the ditferences dcpendcd upon thc rcfcrcncc tic uscd to compare populations 

Ii anid p’ t( ntial lv mi Ii ading p titer s rha, mi hi ri nit her st,m tine popul tion 

‘a ‘ans usirg a comm r I pc across all poula ions whcn I s di in I ct diff r I 

computed regressions of fun tionaIly r lated pairs r f traits for ach population scpa 

ratcly rather than ss ith analysis of covar ance (ANCOVA). 11 cse population-specific 

regressions were then used to computc expected means at three diffgrent si/es br cach 

population fhis allossed me to determine to what extent the am ne-population pattern 

reported I doss s ared in resp n to the choic d stan lard iz Qualitatis e patterns 

f variation among popula ions that dcp ndcd upor th chot )f reference size are 

r(ted in the te\t. All of the population specific rcgres ions f he traits e\amined 

arc presented in Appendices I III 

lo examine most patterns of s ariat n in tng populati )ns. the reference size used 

V as approxirnatelt the averace size icros all populati r tot each species sepaiatelv. 

F-or traits ss bern sluantititis e comparisons hctss i spccics ssere also telex ant. I used a 

single reference si/c for all three species shtch v as close to the aserage for them all.

in IS \\D tlO\ODO\l 4 SHELL \ \RL\ [1Os 

When examinina correlations between char icters ithm species or hetren speLies, 

I computed both a sta idard parametric correlation coelticient and non parametric 

correlation statistic (Spearmans coeftjcient of rank Lorrelation: Soksl and Rohlf 19811, 

Variation in adult sue 

RESI1JS 

estimated adult siz b computinc the average shell leneth of the I irgest 2O of the 

sample used for morphonietric analyses. Because a roughly uniform distribution of snails 

was collected across the size range asailable at each site this seemed like a reasonable 

measure of adult ize. 

Adult size aried sianificantl among sitcS for all three species. hut was most pro 

nounced fur 1. clai igere Fig. 2i. Of interest, the largest sized adults of both T. e1avirera 

and 11 nisioora /ahie ss crc found at the intermediate ‘ites of Lien Ha Wan I HH2 and 

11113) Also. for 1. lai i’e, a the smallest sized adults tended to be found at the most 

wave exposed sites (Ping (‘hau and Cape d’Aguilar) 

T. clavigera T1euteotoma M.io 

CDI CD2 PCI PC2 HH1 HH2 HH3 HH4 HS1 HS2 TI’ 

Collection site 

Ffg 2 Average shell length (mean SCM) of the largest 20% of each sample of three 

species of gastropods from 11 sites along the shores of Hong Kong Sites are 

ordered roughy in order of decreasing wave exposure (see Materials and 

Methods for site abbreviations), missing bars indicate no data. Data for P02 

were not included because the sample was unavoidably biased towards smaller 

Snails. See Appendices I—Ill for sample sizes. 

‘ ariation in penis size and sex ratio 

Penis size saried in an unexpected manner among the four iegions studied (Fig, 3i. 

Among ihai. from Hoi Sing Wan, Fbi Ha Wan and Ping (‘han, pcnes were either well 

des eloped or effeetis clv absent l’his pattern was the same whether or not smaller snails 

crc included compare Fig. 3A with 3B and 3C ss ith 3D i. At Cape d’Avuilar the di’

, RICHARD 

PALMER 

tribution of penis si/e ss as 5tili bimodal, but the penis ize in presumptis e females v as 

conslstentl\ larger there than at the remaining sites. Significantly, both I. clavigera and 

I. Icuti’ôtünia exhibited the same pattern. 

80 

60 

40 

20 

80 

60 

40 

— 20 

80 

60 

40 

20 

80 

60 

40 

20 

t 

iiThoi Sing Wan Hoi Ha \Van 

L Cape DAguilar Ping Chau 

A)T. davigera, All lengths 

— Tentacle 

BIT. clavigera. 25mm length 

C) T. leuteostoma. All lengths 

D) T. leuteostorna. 25rnm length 

none

TH4IS A\D 1IO\DO\i.t SHELl \ ARIATIO\ €)59 

Females were, on average, more common than males in the samples for all three 

species (Table I). Sex ratio varied among sites. hut departures from 50:50 sere onl\ 

significant at Ping Chau. where females formed a larger proportion of the sample. and 

at Cape d’Aguilar. where males were more common. 

Table 1 

Numbers of males and females of three species of rocky shore gastropods 

coNected from various sites around Hong Kong 

Site M 

CD1 21 

CD2 34 

PCI 10 

PC2 3 

HH1 16 

HH2 12 

HH3 27 

HH4 5 

HS1 14 

HS2 20 

TT 

All pooled 163 

C hi-square: * 

d.f’ 

Variation in shell repair frequency 

The incidence of repaired shell injuries varied b several-fold among sites (Fig. 4). For 

Thuis Lial’9’era. the average number of injuries per shell aried significantly from 0 to 

0.32 chi-cquare 17.6. d.f. = 9. P 0.040). Rather rernarkabl. the incidence of re 

pairs dropped consistently with decreasing wave exposure among sites in all four regions 

studied, Only at Ping Chau. ho’ever. did there appear to be an overall lower incidence 

of repair. 

Although the incidence ot repairs also varied among sites for Thais leuieo stoma and 

1onodonta labia, these differences were not significant statistically (chi-square 1.09, 

d.f. — 

3, 

P 0.78. and chi-square 9.1, d.f. = 8. P 0.33. respectivek.Somewhat 

surprisingl. there was no concordance of repair frequency between either of these 

species and L L’laviera. 

Thais c/a vigera Thais leuteostoma Monodonta Iabio 

F ? M F ? M F 

19 0 

21 0 

30 0 

8 0 

21 3 

26 2 

18 0 

14 1 

24 0 

20 0 

200 6 20 

25.5 

9 

0 002 

2 20 

8 14 0 0 0 40 

6 16 1 

6 9 0 3 3 3 

9 19 2 

11 11 8 

12 16 1 

14 9 7 

0 0 24 

0 0 24 

1 5 14 

41 22 50 63 123 

0.74 4.46 

2 3 

0.69 0.22 

Note: For Thais, individuals with a penis that was clearly larger than the right tentacle were con 

sidered males, those with no penis or a penis clearly smaller than the right tentacle were considered 

females. For M. Iabio. sex was determined by gonad colour (see Materials and methods). ?—sex 

either ambiguous or not sexed. See Materials and Methods for site abbreviations. 

*Contingency table analysis on M and F categories only for sites containing > 10 sexed snails.

fl4-i 

A. Rl(H.-\R[) P\L\IFR 

MIL1h1o 

CD1 CD2 PCi PC2 HHI HH2 HH3 HH4 HS1 HS2 IT 


t repairs per sample total snas 

Fig. 4. Average number of repaired shell njuries (Iota 

per sampl& for three species of gastropods from 11 sites along the shores of 

Hong Kong. Sites are ordered roughly in order of decreasing wave exposure 

(see Matenals and Methods for site abbreviationsi. misstng bars indicate no data 

except where ndicated 

flometric variation 

All populations 01 [hai. lu (cent and T. ! ureo\r(ona. and all but t\\ 0 pOpulatiOnS of 

Wolif)d,Jllht 1(11)10. exhibtied po’itis e allornetrr (or ltp thikness rciati’e to aperture length 

(Fig SAt. At all but one site T CiU 1:010 exhibited the most pronounced allometr\ and 

it v a also the most consistentlr allometric of the three specie’ statistically significant 

8 of I () populations). In addition, three of the ses en statistically sienidcant allorietries 

tm T. las igeIa, and two ot the three for W lablo oc urted at Hot Ha \Van sites 

In contrast. all populations of flints k zite stoma and Mm lonta lubn and all but two 

populations of T dat item, exhibited negative allometry of aperture izc relati e to shell 

capacitr (Fig. 5B>. M Ia/no exhibited the most consistent allornetrr (6 f 9 populations sig’ 

ntfieann. wherea’ that e’,hihtted by T. i/rn Ct ma is as somewhat ic” consistent ‘ of to 

populatIons signifieann. Among geographic regions. the extent of allometri in if. ithio tended 

to increase is tb increasing is ave e\pd)siir I Tai Tan — Hid Sine Wan —s Hoi Ha Vs an —‘s 

Ping C hum. ‘1 he sample Ironi (‘ape ci’ Aguilar. howeser, was not consistent with this trend, 

Very similar patterns were also exhibited by all three species tor aperture site relatise to proW 

jected area, both within and among gcoeraphic regions, although the average allomctry 

exhibited by all three species was somewhat less pronounced (see Appendices I III), 

All three species were approximatel\ isometric for shell dr weight relatis e to hod 

dr\ weight. when as eraged across all sites Fig .5C The magnitude of ailometr\ 

how cc er. varied rather markedly among sites Although Thai leueo 51(000 and 

ifonini’inta la/na did not exhibit an notable patterns, the allometrr of 1. let ieeni 

s armed in a curious w av. First, with one exception. allornetrr increased with decreasing 

exposure among sites within each of the four geographic regions examined: the exLep 

Lion was HH I Second, in contrast to the tirst pattern the acerage allometry for each 

region increased with increasing wave exposurc 0)85. 1 04, 109 and 1,19 for Hoi Sing 

Wan, Hoi Ha Vs an Ping Chau and (‘ape d’ guilar, respectis clr).

1.0 

TH4IS ‘\ND i’fO%Ol)Ov7A SHEI L VARIA1 ION 

Laviera T. leuteostoma 0 M. labzfJ 

A) Lip thickness (mm) vs Aperture length (mm) 

00 

:i[ll**i1U 

B) Aperture size (mm 2) vs. Shell capacity (mg) 

ft7 

l4j 

06 

CDI CD2 PCi PC2 HHI H112 11113 11114 HSI HS2 I I’ 


Fg 5. Coefttcmnts of aNometry (RMA slopes) for A, apertural Np thckness relahve to 

aperture length B aperture sze relahve to shell capaoty and C shell dry wmght 

relahve to body dry weght for three specjes of gastropods from 11 mtes along 

the shorec of Hong Kong Dashed nes ndmate mometry Asterisks nidicate 

samples exhthhng sgnifcant allometry (P

66 ‘\ RICh SRD PALMER 

ariation in antipredatory shell features: tooth height, lip thickness, and knob height 

Of the tv o Thai spcies. only I. clasigra consistently exhibited apertural teeth Only two 

of t4 I k iu toma vere touni with apertural te th anti in tese idisiduals tieth were 

not os ny well developed (0(5 and 0.2 mit from 1111’ and PCI respectively) Ihe great 

est di. elop sent of apertural teeth in 7 da 1 aa V as at Hill and CD2 Lie 6A), but the 

extent (I t oth di lopmer t did not vary in any notable way ther witi among 

geographic regions Ihe lack it apertural teeth at PC2 was probably an andact of has ing 

npIed pr dominanth uSi. siles and small adul s at this site see site dese iptions in 

Materials ai d methods). The least well developed teeth among the remaining san pies were 

at 11112 

\ I tFre species exhibited it ughiv equivalent variation in lip thii.kness smong sites 

(Fig 613, note that lip thickness did not include apertural tooth height) As I it apertural 

t.’oth he,ght bp t kaes i” iL’ C ‘ highe’t at HHI md rp , and lowe 

at 13112 1 m few sites yielded 1. knit stoma for any patte ns to merge In AIr iodon a 

lahi Vp th kness tended to ini.rease with decreasing w ye exp su not only among 

stc wt1i geocraph e regions (Hoi Ha Wan, lhi Si it. Wan but al o among ceographic 

ieei(n (114 1 48 1 14 1 50 and 1 56 mit for Cape d Agu lar Pirc Chau, Hu Ha 

Wan hloi S ng Wan ard i i Ian re pecti Iv 

F c t nu uallv elI d i.lopi.d kn bs the ii pool sample In m Pins Chau 

PC2 n he ne of I/a xhibi ed v’rv pronounc d var or r Fe d ebpm nt 

)l sic s ulptur o tls es at sined (Fin 6C) ‘\o i. hclc s both F a ,cua ard 

I tar t ‘s produe d t kirg y larger Inhs is the I rg (F ioOs gg st ng a 

troig, rsnnlv exp it. d effc of enviro ment 

‘I,’ ariatior n aperture SIL and shape 

pee s xhib ti.d i Ian patter is I r a i r ri dat ye ap U 

c ur c i at a v it sh I eapac t ) a r u e size i e e s d it 

u r ot onl m it g it s h s g igra F ic g n Fig \) b 

o c egi s Tail ‘ iese pat ri it r v r r p r d 

iii (Fig 713 a 

re hare i.xpr i.d ap t r I ar a a i g aper r I ,, F 

h c m i t nt (at en Ii 7 ) Ta las ia ho vi ape d’ \g 

i. n U na reia iveis viner perturc ‘s ii ir os ru vi er pe n am 

or Cy d Au la d rl tiselv v d n ap r r s r e I P g 

Frif ecs i F s e Vs d ie ptt r at 

Variati m in shell weight and o cnpied volume 

/ cx 

in r asrg 

C ar on 

n salle 

t x 

a and un 

C it rora 

(h ‘sc 

with r is R F i e v Ft ii d i Ii ire gi Ii n 

g 

p t n ti iuh it d d my r 3 t e i/c lh 

pr v t icn c udfi is ri 

( S 

at rs s r. po d 

it 

a ato i 

akf i 

(r I en e 

V r r n I n s. 14 p a Ii 

Beuew 

r ,aI L’SIL it i f d I e ci p i s, r. s Ii I 

s s r par rclati shill it ‘ight and o eupie 

sir’ 

e ure 

1 i 

Icr ‘t I as 

luantt ti 1y urn.

O4 

TIl tic AND WOIvODOIvT4 SHEL I vAR1ATION 6(3 

T. clavigera T. leureostoma 

0.6 A) verage aperture ienth 

02 hi1i± 

B) 

Fit iIii.JaiIiFiEi 

4 C) aserageapertm len )LhJ 

11iIJi 

CDI CD2 PCI PC2 HH1 HH2 HH3 [1113 FISI HS2 ‘11 


Fig. 6 A average apertura tooth height. B aersqe apertura lip ft ckness, and C 

aierage sculpture height for three species f qastropods fron 1 1 sites along 

o shores of Hong Kong Af bars are mean si for sna Is at a common ap 

erture length sp cif c to cad spe ie approx nately a e age a a a eng h 

19 3 21 3 and 12 2 mm fo Tha;< claviqe a T e tao, toma ar ilono a 

lab a respect iely tes are ordered r ug y n order of de r as çj ‘a’ a 

posure see Mater a s and methods for site abbreviation m s ba s indicate 

no data unless noted oy a .ero. Approx mate s’gnificance Ic e s r m ANCOVA 

for common slope - 

0 

equality of slopes among populations a d equaty of 

aciusted means assuming equal siooes were as foiiows T c!av;gera’ A — 

801 A) 

j . 

160 

90 

80 

A. Rl(HRD P \LMER 

T. clavigera T leuteo,sto,na M /uhioj 

rage shell capacitj 

.I1iIL1i 

B) E1IsheI1cacltJ 

/ ini11,i1fl 

n 

20() erageapertureIejJ 

180 

160 

140 

1.I41[1r1, n 

CDI CD2 PCI PC2 Hill 1*12 HH3 HH4 HSI HS2 11 

Collection Site 

Fig. 7. Vanation in apertural traits for three specms of gastropods from 1 1 sees along 

the shores of Hong Kong A, aperture area at average shel capacity (1215 1726 

and 897 mq for ma/s clavigera. T. leuteostorna and Monodonta labjo, respec 

bvely): B. aperture area at small shell capacity 491. 556 and 363 mg for T. 

clavigera T. leuteostorna and M. /abio. respectively: and C. aperture area at 

average aperture length (193. 21.3 and 12.2 mm for T. clavigera. T leuteostorna 

and M. Iabio. respectively. All bars are mean srv. Sites are ordered roughly 

n order of decreasing wave exposure see Materials and methods for sOc ab 

brevations. mssing bars indicate no data. Approx mate ngnOicance levels from 

ANCOVA for common slope — 0. equahty of slopes among populahons, and 

equalty of adjusted means assurrurig equal slopes) were as follows. T clavigera: 

A, B

THAIS AND WOODOPvT I SHELL \‘ARIA lION 665 

Table 2 

Average aperture area (mm> at a standardized shell capacity for two sizes of three 

species of rocky shore gastropods from five geographc regions around Hong Kong 

Thais clavigera Thais leuteastoma Monodonta labia 

Region Ave.* Juv * 

Ave. Juv. Ave. Juv. 

Cape d Agullar 151.5 86 0 173.5 84.9 120 4 67 4 

Ping Chau 149.3 834 1756 85 1 108 7 64 8 

Hoi Ha Wan 136,2 77 0 103 4 60 0 

Hoi Sing Wan 126 9 70,2 99.5 56.8 

Tai Tan 99.0 55 3 

Note Tabled values are averages of the adjusted means from Figure 8A and B. 

*Tbe shell capacities at which aperture sizes were compared were computed for each specec us 

ng the followng RMA regressions tabulated in the Appendices 1-144 1151. and III 112 for T. 

clavigera. T leuteostoma and M labto, respectively. The shell lengths for which average’ shell 

capacities were computed were, 27.5 30.0 and 18 0 mm while those for juvenile shell capacities 

were 200, 20.0 and 13 0 mm for T. clavigera T. leuteostoma and M fable respectively 

species as well as within species. For these comparisons, I chose a reference value close 

to the average of the lou-transformed values of body dry weight for all species com 

bined (150 mg. actual detransformed mean 148,2 mg) and shell capacity (1000 mg, 

actual detransformed mean = 992 8). 

In spite of their very different shapes, both Thais clavifera and Monodonta lahio 

exhibited remarkably similar overall shell weights at a given body weight (Fig. 8A). T 

clavigei a, howes er, exhibited much more dramatic variation among sites than did M 

lahio (1,77 vi 1.18, ratio ot maximum to minimum adjusted mean among sites). 1. 

dasigera also exhibited a dramatic decline in relative shell weight with decreasing 

exposure among sites within three of the four regions examined (Ping Chau, Hoi Ha 

Wan, Hoi Sing Wan), although the pattern was reversed at Cape d’Aguilar No trend in 

relative shell weight was apparent among regions, however. Of the three species ex 

amined, T lcutemtorna produced the least shell per unit body weight, but showed no 

significant variation among sites, 

When expressed as a function of shell capacity Thais (lauf,rra had the heaviest shells 

of the three species examined (Fig SB). Significantly, the pattern of sariation among sites 

was very similar to that exhibited at a standard dry body weight, hence this geouraphic 

ariation was not primarily a product of variation in relatie body size. As before, both T. 

leukoskma and Ilonodonta labia exhibited only slight sanation in shell weight when snails 

with a common shell capait were compared In contra’.t to Figure 8A, howeser, both 

species had comparable shell weights esen though their shapes were quite different 

Thais dasie,cia occupied a larger traction of the internal shell solumc. than either T 

kutu stoma or lluiodonta labia both of which vcr quite similar to each other (Fig. 

8(j This fraction of thr, internal solume actually oc upied by a snail also varied sig 

niticantly among sites for both T lavira and 11 lahi althoueh the v’ nation ‘. a 

much more pronourned tor T. cia iv a The three hithcst saIue for I a iç’rra oc 

coned in the two no t was exposed regions (Cape d’Agu ar and Pinc’ Chau) hut I’! 

labia exhif t d no noteworthy pattern cf sariat on Once aga n ditferences amor sites 

for I kut istoma were not si?nifiLant

A. RICHARD PALMER 

T. clavigera T leuteostorna M. labiyj 

gdrydyveight 

2800 B) mg shell capacity 

2400 

2000 

1600 

I NI 

I 

900 C) [t 1000mg shell capacity] 

800 

700 I 

I iIIII 

CD1 CD2 PCI PC2 HHI HH2 HH3 HH4 HSI 


HS2 TT 

i .i. . 

Fig. 8. Variation in relative sMell weight and the volume of occupied shell for three 

species of gastropods from 1 1 sites along the shores of Hong Kong. A. shell dry 

weight at a common body dry weight 1150 mg for all three species): B. shell dry 

weight at a common shell capacity 0000 mg for all three species): and C. oc 

cupied volume at a common shell capacit (1000 mg for all three species). All 

bars are mean r SEM. Sites are ordered roughly in order of decreasing wave 

exposure (see Materials and methods for site abbreviations), missing bars indi 

cate no data. Approximate significance levels from ANCOVA for common slope 

= 0. equality of slopes among populations and equality of adjusted means 

(assuming equal slopes) were as follows 

Selection of characters and analyses 

1H415 AND WO\cILh)\T-l SHFIi \ Rl\TIO\ 667 

DISCI’SSIO’% 

In an study of morphometric variation, the characters measured and the number of 

analyses conducted are in principle limited only by ones imagination and patience. In 

practice. the characters measured and the comparison’, conducted should he selected in 

ads ancc to address particular questions of functional or ontogenetic interest. In this 

manner. one may avoid drawing unjustified conclusions based upon small numbers of 

statistically significant associations after having conducted many possible analyses on 

arbitrary sets of characters. In this study, the characters I chose to measure and the 

analyses which I chose to conduct, were selected for the following reasons 

Penis sre was measured because increasing exidence suggests that ‘masculinization’ 

of female gastropods may he induced by the antifoulant tributy itin (Bryan et al, 1986: 

Gibbs ci al. 1988>. 

Shill /entIi ss as used primarily to pros ide a description of the ‘size’ dependence of 

attributes for all populations in units commonly used in other studies. e.g.. see RMA 

regression equations in ppendices I Ill. Although convenient, shell length may not 

always he a useful descriptor of size’ Palmer 1990). 

Sperture len tli was used to scale out size difterences for traits functionall or 

deselopmentall related to the aperture apertural tooth height, lip thickness, knob 

height. aperture area This avoided potentially confounding effects that might have 

arisen from u,ing shell length asa general size metric, due to differences commonly 

obsersed in the proportional or allometric relations between aperture length and total 

shell length among populations Crothers 1985: Palmer 1990>. 

4perrure area was selected because of its relation to tenacit (Branch and Marsh l978 

and to desiccation resu,tance (Lowell l984(. \perture area relative to projected area 

should approximate the relative susceptibilit to disioclgment by’ moving water: te 

nacity (proportional to foot area> relatise to the maximum drag force experienced 

(proportional to maximum area in the direction of flow). In addition. aperture area 

relative to shell capacity should be an index of desiccation resistance water loss rate 

(proportional to foot area) relatise to water reserse (proportional to the total internal 

volume of the shell) 

Lip tlzukness and apeituial tooth hezçht were selected for measurement because they 

are directls related to vulnerability to attack by shell peeling crabs (ermeij 1978, 

l982d>. 

Knob heii.lzt was measured because such knobs reduce sulnerability to shell crushing 

fish (Palmer 1979). 

Proiected area was measured because it is proportional to the maximum force likely to 

he experienced by shells in breaking was Cs (Denns ci ci. i985. 

Shell Lh’\ weis’hr ‘.5 as measured because of its general aloe as a deterrent against ‘.heli 

breaking predators of all t\pes (Vermei 1978: Palmer l 99 . and also because shell 

weight am a ins en body weight should he roughly proportional to the energetic in 

scstment in morphological defense. 

Bwlv des u eiht was measured to allow the relato e ins estment in morphological defense 

ss emht of the shell relatis c to weight of body ( to be compared among populations.

t6S A. RlCF1\RD PL\IER 

Size/I opacity vs as measured as a component ot desiccation resistance and as a means 

of comparing the relatis e amount of shell invested in protecting a gis en volume of 

living space ot a shell. 

Occupied z a/time was measured to compare the relative amount of living space actu 

al1 occupied by the snail because this may vary in response to ens ironmental stimuli 

(Palmer 1090). 

Masculinization of female Thai 

Ihe concordant geographic variation in the extent of penis development in presump 

tive females of [hats davigtra and T leuttostoma (Fig. 3) suggests very strongl a 

c.omnion cause. In tVu cl/a, masculinization of females is widespread along shores 

exposed to even very low lcvels of the antifoulant tributyltin (TBT) (Brxan et al 1986; 

Gibbs a al. 1988). These data thus suggest that Cape d Aguilar experiences higher 

levels of TBT than any of the other sites in Mirs Bay or Tolo Channel. Little more 

can be said, however, without a more thorough survey of masculinization on other 

Hong Kong shores. 

Shell variation: genetic or ecophenotypic? 

Patterns of interpopulation morphological variation by themselves are not very in 

formative about the mechanisms responsible br that variation, Consistent differences 

may arise among populations via at least four pathways: (1>. the cumulative effects ol’ 

selection over several generatiOfl.s coupled o, ith low gene flow among populations: t2 >. 

intense selection tor a subset of genot pes from a large pool of genotvpe arris ing in 

each generation: 3>. selective recruitment of particular genotypes to particular habitats: 

or (4>. ecophenotypic effects vs here even genetically hornogenous populations may dm 

verge from each other in response to ens ironmental stimuli. 

I think the first mechanism ma\ he rejected for the three species examined here. If 

‘i4izudoiita labia and the tvv o [holy species studied here are like their congeners. they’ 

‘.vill have planktonic larval stages Fretter and Graham 1967: Spight l97fo. vvhose 

planktonic period vs ill he on the order of nne Fretter and Graham l962z to two weeks 

or more (Webber 1977). respectivel\. Hence it seems unlikel that larvae released by 

parents from a particular site on the shore vvnuld return to that same site. 

The second mechanism would also seem unlikel to account for the variation ob 

served here This mechanism should result in decreased character variance vs ith 

increasing si/C. hut scatter plots on log-transformed ayes rev ealed that the variation in 

larger mdiv iduals vs as at least equis alent to. or sunless hat larger than, that of smaller 

ones. 

Although I cannot reject the thmrd mechanism re1erental settlement is alvs av’ 

possible vet ver difficult to detect ithour reliable genetic markers—- I feel that thc 

fourth mechanism, ccophenot pie plasticity is a more likely espianation br the hulk 

of the interpepulation ariatiori reported ab )s c Numerous studies has e now demon 

strated experimentally that gastrop d sh 11 shape (Kemp and Bertness 1984: Etter I 9%Sa. 

shell vs eight and apertural d fenses (Appk.ton at d Palm r I 988: Palmet loot>>, and life 

histoty attribute’, (Cross I ar d C 5 h 1990) car all ho piaslicil\ in response to 

ens ironmental stmmnul i

IHAJS \NDt4O 5ODOyI4 SHELl VAR1VIIO\ 669 

Magnitude of shell variation: differences among Hong Kong species 

The morphological variability exhibited by the three species examined appeared to 

depend upon the range of habitats over which the were found Thais da igera was 

found at all ten sites which were thoroughly sampled (Iai Tan was not included here 

because the tide was too poor to determine which species were presenu, and for all but 

one character (relatixe aperture length). it exhibited the broadest range ofsariation (Table 

). Ivlanodonta labia was found at eight of the ten thoroughly sampled sites and was 

morc variable than I. kutostorna for two thirds of the traits they shared in common, 

T kurtostoma was found at the fewest number of sites (four) and for the majority of 

characters examined, it exhibited the least sariation. Although the samplc size is not 

large here, these data are consistent with thc siew that ecological generalists are more 

morphologically variable than ecological specialists 

Relative character ariahility êkn appeared tn he cnrrelated hetween the twn species 

which occurred oscr the broadest range of habitats’ characters which were relatively 

more xariablc in Thais (lavigera also appeared to be relatisely more variable in 

Monodonra labia (Fig. 9). For both species. apertural traits (size, shape, and allometry) 

were relatisely less variable than those related to morphological defense (shell weight 

and lip thickness), Here again, too few species haxe been examined to draw any strong 

conclusions. Nonethelcss, the proportionally greater sariation in defensive attributes 

compared to those relating to desiccation resistance or tcnacity is intriguing. particu 

larly if thc bulk of this variation is ecophenotypic. Perhaps selection has been stronger 

for plasticity in defensive characters than for haracters related to physical stresses (but 

see below) 

1 6 

1 2 

08 

Aperture size 

allometry 

Aperture size 

Lip thickness allometry • 

Shell weight allometry 

• Lip thickness 

Adult leng th • Shell weight 

• Occupied volume 

Aperture shape 

1.2 14 16 18 2,0 22 

Thaza clavigera Max/Mm 

Fig 9 Assoc atian between the relat ye vanab hty of morphoiogca trade m 7 iais 

c’avp,a and Morodo a iabu 0 82 P = & 007 Speairnai P — 0 O6’) E ch 

pm repress ts the. ratio, for a angle trm of the max rrun populahon rean 

‘ r e m r mum populaho i mea obser ed among all t( e crIes sampled data 

om Table 3)

Tabe 3 

Ranges of morphologica) variation in three species of rocky shore gastropods from Hong Kong. 

Monodonta labio 

Thais c/a vigera Thais leuteostoma 

Mm. Max. Max. ‘Mm. 

Fig. Mm. Max. Max.. Mm. Mm. Max. Max/Mm. 

Trait 

Adult’ shell length 2 279 41.2 1.48 32.4 45.2 1 40 19.1 26.7 1 40 

Lip thickness allom. 5A 1 33 284 2,14 1.14 1.55 1.36 085 1 75 2.06 

Aperture area a))om. 5B 06 0.68 1.13 0.62 066 1 06 0.57 0.64 1,12 

Shell weiqht a))om. 50 0.77 1.29 1.68 1.11 1.21 1 09 0.8 1 15 1.44 

1.15 1.66 1.44 

11.3 132 1 17 

937 120.4 1.28 

103,9 1061 1 02 

1610 1908 1 19 

571 608 1.06 

Tooth height 6A 0.06 0.44 7.33 

Lip thickness 6B 0.79 1.45 1,84 0.63 1.01 1.60 

Sculpture height 60 0.94 2.74 2.91 1.53 3.25 2.12 

Aperture lengths —- 18.4 20.7 1.13 21.0 21.6 1.03 

Aperture size (area) 7A 122.1 158.8 1.30 173.5 176.8 1.02 

Aperture shape (area) 70 128.6 155.3 1 21 174.0 181.4 1.04 

She)) weight 8A 1271 2253 1 77 1025 1184 1.16 

Occupied volume 8C 601 839 1.40 556 607 1.09 

Note. Tabled values are the minimum and maximum adjusted means for each of the three species as depicted in the indicated figures (units as in figures). 

At a given aperture length. 

At a g:ven sheli length. computed tram regressions -112 to -121 for T. ciavigerai. 11-38 to lI41 )for T. leuteostorna). or 111-83 to 111-91 (for M. Iabio(, in the 

Appendices 

At a given shell capacity. 

At 150 mg dry body weight 

At 1000 ng shell capacity

IH4JS AND ‘1O\(1DO\T4 SHELL’s \RlA[tO\ 671 

Magnitude of shell ariation: comparison with temperate species 

As emphasized in the introduction. most of the extensive studies of morphological 

ariation in marine gastropods has e Lonceiitrated on temperate species. t’nfortunatel 

lack of a common methodolog rather limits the ability to compare quantitatis el the 

extent of intraspecific morphological s ariation. To conduct a preliminar examination. 

I computed comparable measures of shell variahiliir for three notoriously s ariable 

species of north-temperate .\ mel/a upon ss hich I has e ss orked. Rather surprisingly. Thab 

laiier, in the vicinity of Hong Kong exhibited a range of variation nearls equis alent 

to these species (Table 4). Ross eser, the rather impressive magnitude of variation ex 

hibited h T. (lavigcra may not he representative of tropical Thais. It is more a 

subtropical than a tropical species because it appears to be near the southern end of 

its range in Hong Kong (Abe 1985b). 

Allometric variation 

The patterns of allometric variation (Fig. 5A C) were somewhat difficult to interpret 

The consistent positive allometry of lip thickness exhibited by all three species (Fig. 

5A> occurs commonly in marine gastropods (Verrneij 1980: Palmer l990. Such 

allometr could arise if the maximal rate of both growth was limited h the rate of 

shell deposition (Palmer 19811. If this were the case, however, sites exhibiting the most 

pronounced allomet should also exhibit the highest occupied volume, because the 

bodies of rapidl grossing snails should have expanded to fill as much of the habitable 

volume as possible. Yet for both I hai v elaciç’era and fonadonra laimo sites with the 

highest lip allometry were associated with relatively loss occupied volumes te g.. in HHI 

and HH2). Perhaps the occupied volumes at the time of collection were not representa 

tive of differences in growth rates. among populations. In addition, none of the three 

species consistentl exhibited any allometry of shell weight relative to body weight Fig. 

5C). further suggesting that the rate of shell production does not limit the rate of both 

growth in these snails. 

Perhaps the most interesting allometrv was exhibited by aperture size. On the whole, 

all three species exhibited proportionally smaller apertures with increasing site (nega 

tive ailonietry, Fig 5B). this was true even for populations from wave exposed shores 

where aserage aperture site was larger (Fig. 7A, B), suggesting that the size-depend 

cnce of apcrture area was less relevant to tenacity than to some other factors, e.g., 

desiccation or predation resistance. 

Concordant variation of characters within species 

Assuming that most of the obsers ed variation reported here arose ecophenot\ pa ally (as 

argued ahoy ci. concord mt variation of characters among populations vs ithin a species 

ma reflect: (a>. des eiopmentaIl independent respon’.c ot different characters to the 

same ens ironmental stimuli: (hL des elopmentall independent responses of ditfercnt 

characters to separate but otherwise correlated ens ironmental stimuli: (ci, correlated 

change due to developmental interdependence, either via some form of tradeoff or via 

linked deseioprncntal pamby a s: or d. geometric non-independence. I has e avoided 

making any comparisons which might tall in this last categor . and has e tried to con

Table 4 

Ranges of morphological variation in three temperate species of rocky shore gastropods. 

Nuceila emarginata’ Nucella lap/I/us 

Nuce//a lamellosa’ 

Mm. Max. Max 1Mm. Mm Max, Max. Mm. Mm, Max. Max. Mm. 

Trait 

Lip thickness (mm) 

1.81 3.56 1.97 

Aperture length (mrn) 13.6 157 1 15 16.8 20.3 1,21* 

Aperture shape (area. mm) 91 4 104.4 114 

She! weiqht (mg)” 2477 5402 2.18 782 1463 1.87 2811 4578 163 

rn 

Note. Tab ad values are minimum and maximum adjusted means at roughly average size for each of the three species. 

Computed from data partially presented in Appleton and Palmer 1988. 

Palaer unpublished data from 10 sites over a wave exposure gradient in Barkley Sound, BC. 

Computed from regressions in Palmer 1990 

At 20 mm N emarginatat, or 25 mm (N lapiflus) shell length. 

At 145 mm aperture length. 

At aporoximately 150 mg dry body weight. 

Th s saiue compares quite closely with the range in a much larger data set presented by Crothers (1985: figure 29. excluding the Severn estuary).

IIItIS AND ‘i4OvODOTA SHELL VRRJlON 

0.3 -fCD2 

/ 

-IS2 pci 

02 

- I 

---+- ±/ 

0 1 tim HS1 +CD1 

PC2 

0.8 1.0 1,2 1.4 1.6 

Thais clavigera Lip thickness (mm) 

Fig. 10. Correlation between relative apertural tooth height and relative lip thickness in 

Thais clavigera from 10 sites along the shores of Hong Kong (see Materials and 

methods for site abbreviations, r = 063, p = 0 07, Spearman P 0 11) Each 

- point represents the mean SEM along each axis for a sir gle collection site (data 

from Fig 6A, B> Note that lip thickness does not include the height of apertural 

teeth The sample from PC2 (identified by t) was excluded from the analysis 

because it was biased towards younger, more rapidly growing snails where teeth 

were less likely to develop. Note that within each region lHoi Sing Wan, Hoi Ha 

Wan and Cape d’Aguilar), tooth height was also positively correlated with lip 

thickness (dashed lines). 

centrate on characters of particular functional significance (lip thickness, apertural tooth 

height, aperture area) or characters that showed substantial variation among populations 

(shell weight, occupied volume; see Table 3) 

In Thais davigua, populations with thicker lips tended to have larger apertural teeth, 

although the association was not quite significant statistically (Fig 10) Note that the 

same part rn was evident among sites within the three regions having reliable tooth 

height measurements Thus, the positive association between tooth height and lip 

thickness doe seem valid Because these characters are geometrically independent (see 

Matcral and mcLhods), ar 1d bcaus both ae intimately asoJaLd vth1.ducing ‘ul 

nerahility to predation by shell breaking crabs (‘vermeij 1978). this correlation seem 

most likely to be a product of mechanisms (a) or (c). 

For both Ihais dasic,cia and lilonodo, Ia labu . lip thickncs was also positively 

correlated with occupied volume (a measure of relative body sue. Fig. II A) lth ugh 

these correlations could have arisen if thicker lips cramped the internal volume of the 

shell forcing the body to occupy a larger fraction of thi spa e (mechanisms (c or (d)). 

the xpected necative correlation between relatil sh 11 weight and occupied olumc 

did not materiahíe (i = 0.14. P 0.71 N = 10 for f lariçera andi 0 11 P 0 8, 

N 9 for 41 labia, both estimated at a common hell capacity of 1000 mg). Hcne 

neither mechanism (c) nor (d) seems likely to account for these correlations in addi 

tion thes’ corr lation are th ievlr e of what would be expected based rn studies of 

predatn induction, xpcrimcntallv indu ed thicker lips ire associated with adureasui 

occupied volume in both ‘Vuella lainrllosa and lapillus (Appleton and Palmer 1988.

A RICH \RD PALMER 

L8 M labia [.davigra A 

L6 

l4 

L0 - 

0.8 

06 

1 6 

14 

12 

600 700 800 

Occupied solurne 1000 mg Shell capacity 

M labia 1. claviera 

LO - 

L 

100 120 160 

Aperture sue (mm’) @ Ave age sh 11 capacuts 

Fig 11, A cor elat or betwee i relabve ip 14 kness a d re ati e occupied vo ume in 

Tha cavgeraand Monodona abiofom 1 aid9s e e per veyaongt[c 

s res of Hong Kong Eact powt rep ese ts the car P1 aong a axs for 

a 14ngle coNe hon site data rorr Figs 6B a d 8C) For T c avigera r 0 71 

P 0 031 Spearr an P 0 90 for M lab o r 0 2 P 0 0 8 Spe rrra P 

0 0531 B correlat on be ween re ahve ip th ck iess and relat ye aperture s ze 

r sar1 sampas rf T cIaCger ard hI, ahc. The Jata are fr “gs 6B 

a d 7A For T c avgera r - 0 69 P - 0 028 Spe- rr ar P 0 053 or M I b o 

r 087 P 0002 Spea man P 0044 Note t Np thckres does not m 

rude if e feig[t of ape tural bet Da fed res we tted b cy 

a ni r 1 ( I t us a corrc laiL r )r e t ( r r n n r a tim I as t d 

ith p edatior rubs meL am ci ( ) a 1 s er 1 k I , 

I 

B) 

ak t thcr tc 

it ns uc 5 tI’atlipthikrc ard.c u d uncrfcti deper ii prse 

sep ra e I Ut err Iatcd I tur’ r I if n ir rimerit in i r ism ) e a ci if arc. 

I x d a lab Ii y, and pr d in intensi 

Cu i u nat Ietwrcn lie thick e an pLrtur s it ditt d 1cm ci 

r rd 14 n ntz h, Fit B) r r ap t r d opu 1 o I 

i. uter ed haethike ip nheea i ck &rascd vii in L 

apertur sue in /ll ahi F Ii hth pen larc.ei ipertures sscr a sociated s ith ir

I/I tIS AM) tIOM)DUVT4 SHELL VARI \TlO’s 

creased ssase exposure (Fig. 7A. Table 2; see also Fig. 12 beloss), thus for some rea 

son I daszge a produced thicker lips and 14. labia thinner lips on mors. ssase exposed 

shores N4edianisms iai (et and (dt all seem unlikely causes because they cmld gen 

erate parallLl patterns in both species Thus these contrasting correlations most likely 

retlect some other differen es in the biology of these to species. Since 1 lad c,u is 

a predator of or sters and small his aRes (Jay br 1980> hile Al. labia is presumably a 

microherhiore like other trochids. and since these species has e ccx different shell 

inierostructures. most notabir extensive des elopment of nacre in 14. lal’e (personal 

ohserx ann. manr explanations for these contrasting patterns are po’sihle. 

Uoncordant morphological ariation of sinpatric species across enxironments 

\s xsith correlations among characters within species, many possible correlations could 

base been examined between species. However. I chose to analyze only four characters 

shared by [haS dan izuu and Wonod n a labia ftlr interspecific correlations, Tsso of these 

exhibited con. Istent associations across habitats ssithrn speuies (aperture i,e and lip 

thickness i and Iss o exhibited a particularly high range of s ariation ( shell ss eight and ocx 

upied s olume: l’ahle 3i. These seemed the most interesting to exanline because either 

the presence or ahsence of correlations ssould he informatis e. L ntortunate] . the third 

species. 1 Ieuieevlrmra. xx as not found an enough sites to rnclude rn these analyses. 

Table 5 

Summary of the strength and direct on of association of she I trarts 

with two major aspects o the environment n two spec es of 

rocky shore gast opods from Hong Kong. 

Wave exposure’ Crab predation ntensdy 

Trait Fig T c’av T. lout. M. iabrc T. clay T feet M. fabyc 

Adult she ength 2 - 

2 2 2 2 

Tooth ‘reight 

Lip tb ckness 

7A 

7B 

x 2 x 

Sculpture heg F 

Aperh re size area 

Apertu e s ape (area) 

Shot weghF 

Occupied vcIume 

7C 

BA 

BC 

9A 

gO 

+ 

+ 

0 

0 

Q 

0 

7 

2 

2 

2 

2 

0 

7 

Q 

x 

0 

Qattat!ve rafl1flg of resors by wave e+pos..re: Cape d Agusar > Png Cnau Ho Ha War > Ho 

Seg Wan Ta Tan. Wtrr each region seaward shea here conwdered ‘e(at’xel( more esponed. 

P’edato” nterstv essence o be propotior’a ro rcdence 0 woared shet “unec Fe 4 

At a giver apeCure ength 

At a gven see 

5 capaey 

At 50 rrg dry body weig’r 

At 100 110 shel capact, 

* ‘ o g a d con s poetve as aton ± weak o 1 a nsistert p dv ass cc 0 

1 end car dfference’ - a rg s e w e o ess orsste t nega ye a socatjo 

tror g ard c snt r gat e ocntio 2 ssoc etor b guou x r t app cebin Cova t’ s 

used fc err pare trads given ir fgures (isted

. RICHARD 

P \LMER 

Although lip thickness exhibited the greatest range of ariaton of character’ shared 

by all three species tTahle 3, excluding illometric variatione and also e\hbited a con 

sistent negatise association with ase exposure at least in ‘ttonoth eta lahio ilable 5i. 

it exhibited only a weak negative correlation betw een Thai s las- gr ; a and 4f lahi (r 

0.59, P 013. N 8, Spearman P = 0 14, data not shown. As suggested in the pre 

ccdina section. th;s negatis e correlation ma he a product ot differences bets een these 

tis e species n the biology ol feeding and growth or of shell production. Additional 

;nforrnation i needed before an conclusions can he drass n \eithei sit the othei tss o 

shared characters s’s hich shos’s ed substantial ‘s ariaton s crc signficantl’s correlated be— 

tween [ iat-igera and %f. labia shell weight at a common both dry weight ; 0. 0, 

P 0 48, N = 8 Spearman P 0 17: o copied volume at a common shell capacity r 

(1,26 P 0.54 N 8, Spearman P 1 0) 

‘- 12u 

perturearea@averageshelipacity 

+CD2 

U t 110 ,,4,, PC2 ± 

100 

• HS2 

HS>4-s 

90 ‘“ 

130 140 150 160 

Thais clavigera ‘ Aperture 

2) 

area (mm 

Fig. 12. Concordant var;aton in relahve aperture size between Thais ciav;gera and 

Monodonta labio among the etght sites sampled to Hong Kong (r = 0 72. P = 

0042. Soearman P = 0096) Each point represents the mean - 

axis (or a single collection site ;aata trom Fsg 7A). 

v 

along each 

R latise apeiture site, Ilk trait which va most eonsistcntl associated tic 055 envi 

rarimer’ts ii ohio species Table Sc ‘s’s as also ignificantl\ correlated hcts’seen species 

Fig. 12;. shores harboring large apcrtured [hats Ia; iges’a also harbored large aperiured 

tI,in,isJ,n;ra lab; i. The ocLurrence ot huger .iperturcd populazons oil niore S’s as e exposed 

shores is i conimon pattern in rock —‘hore zastropods (\ermeij 1 ‘s7: Crothcrs 1985: 

See1e I $6;. ‘the fact that these t’s’so sp cics should exhibit cone )idant ‘sariation in 

apertur’ site, despite their di tinctise taxonomy aid ecology suceest that water 

m scnicr t is an environmental for e with an influence m shcll (oim that trio cends 

laxonomic arid ecoloizical considerations

11141S AD 4/1O”v( DO\[4 SHEI L \ARl \IION 

Phenohpic correlations with incidence of shell repair 

Nithough the im.idence of repaired shell injuries ‘varied substantiall’v among populat ons 

from a low ol n ne, to a high of (>38 repairs per indixidual (Fm. 4), the oserall aver 

age incidence f repair ‘vas quite compar bi among species (mean srxi’ 0 13 t 0.033, 

0.08 0 021, ar d 0 18 (3.033 for [ha s laszçera [.lcu u ira and 41 i d ma au ), 

respectis ely) These frequencic s c )mpared favourably to those obsers d in \ tic c/la 

lap I us (Ve m ij l98a> and Littorina litt ica (ermeij 1982b from the North Atlan 

tic, Hence, relatise to other species from rocky shores, snails ftc m Hong Korg appLar 

to ha’ve a similar probability of experiencing and surviving a crab attack 

In spite of the roughly tenfold ‘variation among sites differences in predation ir 

tensity accounted for ‘very little of the obsersed interpopulation shell ‘variation, fo 

obtain an oserall index of crab predation intensity, I pooled the repair frequenLies 

across species on the assumption that, within any particular habitat, each f these 

species was equally likely to be attacked and proportionalls likely to sursise the 

attack None of the traits considered likely a pi ion to respond to ‘rab predation in 

tenity exhibited significant associations with repair frequency for either Thaiv 

or Uonodonta labto’ apertural tooth height ‘= (i 0 34, p o 37, N = 9 

Sparman P —(>99, T. cJa’viç’cra only, PC2 excluded), lip thickness (ti - (3.10, P = 077, 

N= lO,SpearmanP=08l.andi 036,P=0 34,N=9,SpcarmanP 030forJ 

c/ar gela and 41 lal’h, respectisely). shell weight ( = 055 P — 0.10 N 10, 

Spearman P 0.22, andi 0 16. P —(368, N 9, Spearman P 0.67 for T ciarigera 

and 11 /ahio, rcspecti’vely), and occupied ‘volume ft 0 48 p (>16 N 10 

Spaarma9 P g 14 and i 0 46, P 0.’6, N parn9 P 06 for!. acu’c,a 

and 11, labia respectively) Either (a) larger sample si/es are required to describe 

hc ‘variation of repair frequency among site more accurately’ b), repair frequency 

is a poor indicator of predation intcnsit’v in certain situations (Vermcij l982c), or (c), 

the bulk of shell variat on observed in I. la’vie,c a and 41 labro is not in response to 

different lesels of predation In view of the rather substantial ‘variation observed in 

these species, this topic would seem worthy of further study 

CON CLC SION S 

To my cu p isc, in spite of con iderable variation in many morpl olc meal n ads the r ly 

particularly st ikina association between shell form and environment revealed by this 

stud’v v as between aperture area and wa’vc exposure (Figs 7A, B and 12, ‘1 able 2) As 

discussed above this pattern is common in ther species of gastropods and its exist 

ence in two taxonomicalls and ecologicall’v distant species suggests that wa’ve acti n is 

a significant agent f selection n Hong Kong shores 

I find it pu//ling, howexer, that traits related to predation resistance exhibit d so little 

correlation between cpeLmes r among en’vmronments The incidence of rLpaired shell 

injur is sufficiently high to uggest that shell-breaking predators ar an important sourLc 

)t ca tropod mortalift on H mg Kong shores (1-ig, 4> In addition studies of shell an 

ation in temperate rocky shore ‘a tropods have resealed that shell form ma hange 

uit rapidly in r ponse to a cr ased prdat’on risk (Vermcij 1982a Seeley 1986) a 

large I ati rn of which ma’v be ecophenoty pie (Appleton and Palmer 1988, Palmer 1990)

A 10(1 ARt P L IER 

PrtHth en rati)nmayic Li a unpi nt Ha 

hador tree d n A/we 1 rsi ila Lit. iiiemperst 

an Bertw’ 1984, App eton and P t-u 19 8 P’ inie 

toi h m ar ir,. ites (UI n. tAph utype. Pc 

r a iu 1 U rrr I u ai1 lu 

r r aio A/dab u tie rot[ kinr d 

Ia F of ‘r sd r letwe n ifeni do t c. ma 

r uV 

r a sti 

n 0 ue 

f ri 

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F ‘I is 11 ‘i 

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pe N A rr ej 1 80 K rp 

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hap Lii i v hd to Iho 

tf Mo ohr’o 9 Aoe 

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tion ri only I at onu f thu n not 

Seuw iha uredt 

apola r iuci)a /i/ 

p. F hsouu tro hd vouidsh s srriarre pa ird r i Th icrativ 

‘elatin Lv ath s peec. lii F 1 A a 11 f ted r tory 

1’ i A rriuij 8 iii it ed ately sugg th’ as ur p ior sal I Perla h 

te for uent) ‘reiti n kthat a cvi edit mire I riy ak y duc pe 

ram itintr (hid. IaH nu p men e rmnti r I Lv es’ I s I 

h sew p torisk trw )udbno f isa 

fu Bia I to r 1. o 

a in ,rtaa icr 

iufr K c ‘atofo 

)(51 esBIMA’ahom 

t it to tul 

‘, Na N 

Oree n U,, 

H 

ii 

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K ,,r 

a 0 8)1 

d e 

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eai 5’ 

a e ifs 

‘, ,, Pta a , p “ t a a It 

F n ‘ru erp( ibis r scrub as upported F N LR( OP r ( n Ai’TS 

S A 

N 

Al r 

1985a 

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It C I 

tim, c 

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t) 

fi 

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[HAIl ND MOKODOK TA SHELL K RI K T ION .79 

[or R.J 1988h Pi,siobgicalsress and in1 dy I rlfisrsitthc i cida salK Jo 

to i Fo,tx.4.(0 ) 

FoiL I C 19, ‘ Sionifica i f h 

t cit.’ ,r 5 stir tS)S S 

I hick sr a, sr s VI astropoda CJO ida B tic 

Fr ttrr V idCri am A 1)62 B in p t anti it LondtsrR y5’ iety 

(s’bF ,P I Pascie P F andBrLO R l8S Ss. cFa, n [c iid dr 4 leA, it 

op ‘nuedh trihutvlrnfrriatftul’gpaiis /it tt4f B ia 

Ass ii i . (K68 15 31 

Hami .P V.1980 Sb I pnaoonin Vt i’ti r S p ‘a sarat. ertsti it. 

reated Ito d i otRts e 138486 

Jarsin K IAI Al / iieands ci sari non Ha man a F a Praobai La 0 

di rntdispcislabilitie Bit ala at f’t-L nor it )24 

Jrhannc str B 1986 shHI ilrphoI es I Li a. a so Is DI the rrlati a. rt a 1 

physi at a tr and pedsttor 1)0 iattt spti iot Ida Biot s ct/tnt t,’ 

102 18 95 

Kemo. P ard B rtrc s VI D 1)84 Sra 1 Faa.. and rt 

al t ne y in I I t i hit Pi , o r ‘s t N I 

I 511 P. 

2S1’ 44 

Kit hre J A Kurt L a dF(lint F 1)6 Ihc (I 

si ‘nihc r V sF cii and body form in a. to tiiii 

faBat ra ‘vi .19K Stals nobody sa saf’cti it 

5 (55 F S sitmotit 20 ) It 

Isa ill 1.1984 Icic aft sla ifriE limiets etc 

ites e ‘de it. for pla t V I 

4adrs IS a. I irIS 

Ri, aid 1 f05 Lxoi oat r not a i/i nm, ,‘ i, d Ilais Ianmlhsa so n 

aids Seattle Ciliistoir Co 

Ktl’inm.J A ndLorsssoo J 9’4 Obse ati r 1 trrmaidi e loca s nt 

Lance intlsiidgatopolstfth g rusl lair Ne /aand In si B 

o )u IreNK I ea. 

[In oIL t SI) ( 

‘ci g aides u tr I i B 

I sue t u ite n 

[ape. 1 o no’o[Fsxiiiitrt Ma’int Bohc on F ,s 1 21 

[uckrn , P 5 l/0 K tnation of hel harast tf a spcc of 0 it n (Klollusc’ Ga spi Ia 

at Asamu hi S i n t B pa s 4 It hocu I ortrsit St i its IS Bolts) 35 14 St 

Ncwk rk 0 1. and Doyle K. W 1975 Cstnctic analy is of tell hapc variation n Littoroo 

sot stut s on an enviror mental ch a. [loin t Biott ç 31) t 7 

Pal ix. A. R 199 Fish predation md the volution of a rop t sh II s Ipture pnuinerial 

P’ul 

and F° graphic evidence, Eta into 3,697 “13 

c A R 1981 Do carb mate skelets ns limit the ra sf body growth Ko or 292 Is) 52 

Painter. A R 990 Effect of crab effluent nd sc nt of da naged cc s pecuftcs on fcedtnc or sth and 

sell moupholo.y of the Atlart dogwhelk K a to to I is (F ). II di thut boo 193155 82 

P i lips B F (aTnpbell, N A and Wil n B R IT’) A null sari I study f gea.. api 

auiation ‘n th sshclk I) otT s loi not t Isp in ti M itt Btt s aol F ‘it 5 

2”’ r9. 

I a R H. 86 irtunse iatur 1 clcctit cau F d 

rrarnt sial Pit telocsr / ‘vain S oa / 

strshc dtr i tisr i i irg 

itt itt I SI 53.68)7 9) 

S ill 11 9811 tnaturalh’ an ‘or r if [mIl a iat ti in thep ri ik 

son sill it ho,, titMa, Ba ii 5 ut /1 S S 41 

S a.R R i dR hI F J laS orO ndc S n F i t c KS 13 Frc rr C 

Sig,ht1 VII96Folr.. satIs. irntrr t Ott it.2 -4 

Sruh ala. 

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I S

A RIC HARD P \I MFR 

rnietj. Id J 19’S. Rimi ai api. a d a ip zr i Pat ri a ii ii’ atnhridge. Nlass. 

liarsard Lms erOr Press 

Venneij. H. J. I 95(1. (iastropod shell gr stf rate alL tnctrx ard adult iie: ens ironrnental iniplh 

ation In Sst/t i/ Ci ott ru 

i ark. Plenum Pt ass. 

o aqutr’i ii. a a i ted. Li. ( Rhoads and R. A. Lutz. ‘-i $4. 

at ntet. Ph J I952a. Phenotr pie es olution tn a oorl disprsing ‘nail aIter art is al of a preda 

tar. \l’tt’t 29°: 49 50 

S ermeit H J I 95 ‘h. Ens lronmentai chanre and the olutionarr hstor of titr pen ‘a ink Ic 

iii;: i/tnt fill//tOt n North Anencit. Fr :ar, t. :)hi—X0 

\ crmcij. H. i. i 9X1 I. nsuecessful predation and e’ alution. dine/i at; \a[UritiO I 20:9) I dii. 

\ermetj. Id. J. l°2d. Gastropod ‘hell Ltrm. breaIaoa. and repatr tn relation to predation hr the 

rah Ca/apoa (la/a a/aria 23:1 Id. 

\ermeij. H I. I 957 1 iv /i 1te a 

Lots erstir Press. 

and a ‘a a/alit a -30 0. ‘/ i.ai a! h/train ii tit Princeton: Princeton 

Vt cOhen. H. 11. 1977. Gastropoda: Prosohranchta. in Ri prim non v/ manna ot ,tchioic ed. A. 

(7 Hiese anti .1. S. Pcarse . I 9 \e’a ark: Academic Press 

55 ellinrion. (I. SI. anti Kurts. A M. I °. Gross th and shell armatton in the tropical eastern Pa 

cific intertidal gastropod genus Pioporo. ecoloetcal and es olutionarr implications. Bo’i’ri:a! 

Ba//cnn I h4: I S 

APPENDICES 

These appendices include the descriptis e regressions used to generate standardiied s alues 

for Tltai,s rId/li/era 3 Appendix I T. leureucni’na Appendix Iii and .‘vtanac/oitta labia 

(Appendix 1113 in the ftgures and iable in the text. For each regresston. sites are or 

dered rouehl\ in order of increasing was a espoxte. For each spec ics. regresion 

5 ate 

ordered alphabeticalir 1w N and then Y variables. For each character pair examined in 

each species. a separate regression was computed for each ile i.e.. slopes difbared too 

often among sites to justify usine pooled slopes from ANCOV.\, See Materials and 

n elhods for details of measurement and Discussion for a justification of the s aniables 

analyied. 

Abbreviations: Regr. # number of the regression as referred to in the text: Site — 

see Matetials and methods for site abhrcviati )ns (All recre sion computed on data 

for all sites pooledi: N sample si/c: Mm Max, mir imum and maximum 

ititiansjoirned s alues for the X sariate; Mean mean of the X s mates actuallr used to 

compute the regrecsion (may or may not be log trsnsf )rmedf Lea t Sq. [incur Regr 

results from a least squares lineat regres ion [Slope tandard e ror of slope), 

correlation cocfiictcne] Expected Y @v Mean X Y salue predicted from the least 

squares linear regression it the mean valu’ for X (st mndard erro )f expected 11: RM 

slope reduced major axis slope for a model II regression (— lea (-squares linear re 

gression slope/i. see Statistical analyses section in M termals and methods). Is 

Allom Students r alue fot the difference betxsen the obsersed RMA slopes and those 

exp cted for isomelr\ (set.. Statistical analyses section in Materials and Methods

APPENDIX I 

Thais c/a vigera 

HMA 7 

Sop A on 

Expec.ted Y A 

Mean X 

L a t Sq Urear 3eg 

Slope a) 

X Ax a 

Mm. Max 

Ro>r 

# S3e N 

I02 

02 

02 

1 60 

1 92 

Os 

34 

(4 

76 

4 28 

4) 

11 

29 

X 01 Aoerturn length nm), I log (Aperture a mx mm 

01 HS1 38 141 245 1 296 2 062 0 0544) 0 988 2 59 0 O03 

0’ 3S 40 13.3 253 1 281 2064 00409 11993 2103 00030 

03 3H1 40 102 232 1.234 2009 1 0370 0904 046 0 033) 

04 HH? 3/ 165 285 1.327 2.16/ 00551 0989 2201 0.0040) 

105 NH 44 11.6 297 .333 2013 00341 0994 2216 00013) 

>6 HH4 13 123 262 1 240 2.029 00518 0995 2023 00050) 

07 P01 40 101 246 1.256 2153 00388) 0994 2100 00035) 

08 P0? 11 110 170 1 151 233 02343 0354 1 870 00110) 

09 CD 40 146 224 1.260 2 033 (0 0630 0.982 2 139 (0 0030 

1 C DO ho 88 22 5 1 201 2 05 (0 0298) 0 995 1.981 0.0030) 

A 

(I’ 

00 

01 

1 

7- 

.5 

3. 

01 

3. 

-1 

0 

/ 

1(86 35 

1043 15 

55 1 5 

1 5/ 642 

115 >1 

1076 1>5 

1’)2 

45 >5 

128 ‘ 1 

l’6 385 

X og Apert,rx length mm I log Aperture wdth mm 

11 H81 18 141 245 1 296 1 053 00438 0970 000 00025) 

12 HS 40 133 25.3 1 281 1 029 (0.0271 0 98/ 0 988 0 00)0 

S MH1 40 102 232 1 234 1 039 00296) 0985 )C47 30028 

14 1H2 38 16.5 28 5 1 325 1 233 0.0400) 0 981 1 008 0 0030 

1 HH3 44 116 097 1 333 1 136 00302 0985 1 019 0.0028) 

HH’ 19 12 3 262 1 240 1.064(0 0389 0 989 0 923 0 0038) 

1 P111 40 10.1 246 1.256 1 179(0 0373 098 0 976 0.0035) 

18 t 02 1 11 9 1 0 1 15 1. 4 0 1680) 0 92/ 0 837 0 0078 

1) C)) 40 146 221 1 260 1 064 00609 0343 1 (09 00028) 

20 CD2 88 22.5 1 201 I 100 0 0327 0.977 C 919 0.0030) 

C 18 

68 

3 03 

0 30 

118 

03 

/6 

317 

0 58 

X oq (Apertu e le gth mm Y log (Knob hmght mn) 

21 H 1 38 141 245 1 296 0 36 0.1595) 0 140 0013 00100 0971 

2 350 40 13.3 253 81 0829 01533 0659 0.005 00)18 1 c8 

23 HH1 40 102 232 234 0732 0.1261 0686 0119 00113 1 >6 

24 HH2 40 1 .5 285 1.323 0.021 01690 0.020 3149 00 18) OoO 

I 25 HH3 45 11 6 29 7 1.328 0.290 0 1230 0 339 0 121 0.01°3 0 85 

? HH4 20 123 ‘62 1.28 0.483 0 0.3o2 U uca u.unoa) 

27 PCi 40 10 1 246 1 56 0 /17 0 1362 0 6o0 0 028 .0123 1 1 

‘8 P02 1 119 170 1 151 2 8 05126 0822 0076 00 38 ‘638 

20 CD) 40 146 224 1 260 0 539 030 7 0 306 0 078 0.0135 1 958 

31 CD 55 88 225 1 201 0 494 3 1313 0 459 0 &8 (0.012,) 1 0/C

Appendllx (cont) 

Ixpected 9’ Ca 

M an X 

Leist Sq Lnrar Ragr 

4’lope 

X Axis 

Mn Max Mean 

X g Apr 1 ci qth, r a 9’ log L p hekres cxc dng tenth, rem) 

1 H31 38 141 24a 1 296 0983 02405) 0563 00 2 0 148) 3 18 

3? H’,’ 40 13 53 1 281 0802 01721 0603 0081 00133) 201 

‘3 1 31 40 1 2 432 1 234 211 01968 3 8,7 0.818 001 8 7 

313, 4 165 285 1 3”3 2 319 (0 2a13) 0 838 0 005 3 0175 1? 

. HHI 5 116 23 328 1 333 0 1151) 0870 0082)0 3113 

3 HH4 1 262 1.2’8 1 909)02293 0891 0092 00210) 43 

‘I ‘12 4 10.1 ‘46 1 256 1 820 00969) 0°38 0067 7 0088) /95 

18 122 11 1 9 170 I 151 3’O 01936 0.536 0206 003 8 1 

3 2 3 4 14 224 1 260 099 0 1976) 0 634 0 085 (0 3088 18 

4 7, 3’ a 88 22.5 1 ‘01 I 234 0 521 0 760 001/ 0 0143) 

X Apsiti enqth ma 9’ Tooth heiq it an 

4 HS1 38 141 245 200 0002 .0079 0035 C 98 00’?O 0 

14 1(82 40 133 253 194 0003(00049 0107 1/0 00165 23 

141 HH1 40 0.2 232 175 0017 000’4 0716 3323 00250 0166 

44 3132 4 163 295 3 0 022 0 0056 0 a34 44 3 0193 41 

1, 1H3 45 116 29 218 0.019 00049) 0370 0229 0021, C (3a 

6 13H4 12 3 262 177 0.037 (0 0084 0 715 0 34 C 0123 08’ 

4 p 40 tO 1 24.8 184 0 022 0 0060 0 523 1 38 0203) 004 

43 P120 1 1 9 /0 142 0.000 00000 1 000 0000 00000 0 300 

143 CDI 40 146 224 183 0.001 00014 0.01 0048 00138 0 59 

‘0 CD? 55 88 22.5 162 004 (0 0046) 0588 01 8 0 3150) 0 41 

X log Itody dry wt mg), 9’ og (Sic dry wt mg( 

‘ Hal 38 70 596 0635 (00533) 0908 3392 00113 C. 65 44C 

52 t1S 40 85 596 0920(00930 0976 3 315 0 00/9 0°43 1 4 

(53 HHI 40 a 39 1 192 0.0784 09? 3272 00208 1 81 36’ 

‘4 HH 37 /7 910 08 0(0 9396 0.958 .502 10115 0846 39C 

55 HH3 45 35 942 0 924 0 0342 0 972 3 509 0 120 5 1 44 

r’,4 2u ‘1 So? 1 04? (2 0634 0 cob 3. )‘4 ui00 7 

7 02 1 41 2 430 1.04? (0 0458) 0 965 3 264 (0.0140) 380 

158 P8? 1 35 189 1 019 0 1432 0 922 2 904 0073 13’ 3 73 

I 5 CDl 40 31 367 1 067 0 0725 0.922 3.296 0 0103 1 101 11 

60 CD’ 18 319 1 179 00406) 0370 31 2 00115 1 ‘15 ‘.3 

X og Pro erted xrea mm 9’ og (Apertur arex in 

(01 ((SI 38 133 443 2,404 1 004 0.0409 00/1 1 034

119 434 2 383 093 0 328 987 123 0 4) 4 4 

4 (C 380 225 0933 022 0989 24 6(0 04 094 4 

3 1 9 4 2487 0932 002 1 988 201 004 943 

4 2’ >482 0906 0 21) 098 >21 03 341 4 0 

19 82 43’ 2.257 0 3D 0349) 0 989 c.0? 00 3 9 2 6 

40 a4 364 3 .279 0 31 0 0262 0 984 10) 0 053 C 9 1 

I 82 94 20/2 028 00 90 097 870 00078 1 70 

D 43 144 31 275 0886 (0043 C 946 >13? 0 050 33 2 

> 1 41 301 2 185 0906 00 >0 0980 13 1 3 0 .>4 3 2 

(4 > (UdtA y 0>4 09 Ap> w >. a a o 

(IS 38 11 3147 619 00182 0985 2 4 00)39 

H>? 4) 4>>) 3 138 (6 0 >3) 98. 2 00(40 

4 1 4 1 2 890 0 637 (0 146) 099 044 0 004 

HI >4) 322 0535 0155 388 201 00 40 

181 44 1 3239 0.a95 0.0 >2 331 2 14 00 4 / 

144 zO 2/8 343 064 0.0 8 0994 0?4 00 a, (6 

P 40 (8 941 0(36 0.0 4 0993 2100 0. 338) 

5a >44 066 00363 3.98 (70 0 0(0 11 D 

C 1 1 4 7 2.286 0.6 0 0232 09 4 139 00 35 C 

CD >> > 99 361 00 43 3986 1 981 ( 0 a 3 

71 

25 

0 Y 0 C up> I vo ume ng( 

38 4 I 31 0 47 (86 0(3 0 0984 303 (0) 5 C 

4) 4 9 3237 3 138 0 997 0 0202 0 992 “.30° .4 18 V 

43 4 221> S9 1>”” 224> 0 >9 >73> 1> 4> 

40 9 4464 320/ 1’ 3 I> 0992 3 5 00 3 

4 43 4874 3 317 36 001 a 0 234 ‘ 6 (064 / 

‘0 213 43 89 C 0432) 0988 .440 3 25 4. 

40 08 21 1 >341 1 240 (0 025 0992 1(3 C 3084 9 4’ 

1 >55 4 9 264/ 042(0 10/) 0956 2 41 04 3 3 

0 14 “003 2.986 .013 0 0357 0 977 80 0.00a3 

a 65 52 ‘ 99 1.033 01 0 99 2 aS( 001 5 .43 

3 

0 Y 09 74 drywt T19 4’ 

38 4 1 3100 4/ 0743 00454 0939 .392 033) 4 3 

4 429 323/ 3 138 0.898 C 0233) 0 984 3 5 0 0063 9 3 32 

44 134 220 2890 118 00643 0948 ‘3> 5 1 48 336 

1 ‘2 4464 3.20 0 25 0427) 0 362 3 502 0 1) 3 34> 3 

1> 33 4874 32 7 9/a 0308 979 35 0 00 3 96 13 

2/8 321 >94 4 0599 03/1’ 91 00 5 >43 

4 08 >17 2941 051’ 00402 0374 3>65 00123 1 4 

2aa 939 2644 0°6 01474 0909 >434 (.0238 1 064 0 3 

1) 447 4009 2986 1 060 006/4 0931 236 00(9 13’ 04 

a 6a 1752 2799 1.086 00348 09 4 3 1 0 0 1 3 0 

y s>> 

81 (IS 

3? (S 

03 4 

4 I’I> 

8 H) 

86 4 

C PC 

3) CD 

1 C> 

X 1 >pa 

91 >1 

9? 2 

3 (1 

>4 HI> 

HH 

3 5(4 

‘C 

98 C 

(3 p4 

1 CD

Appendix__(cont.) 

Execteo ‘7 

RMA Os 

Mean X Sooe AScii- 

eas1 So ,near Regr 

Siope .s’.’ 

X-Ax s 

M “ Max Mean 

2 Site N 

X ‘og ‘a) length mm) ‘7 log Aperture area. mm 

l HS1 38 20.8 38.5 1 459 1.597 0.0864 0 965 2.159 0.0058 1 966 040 

‘02 HS2 40 20.0 38.2 1 452 2.’58 0.0666 0.982 2 ‘23 0 005Dm 2 ‘98 297 

‘03 HI-i’ 40 30 347 1 382 ‘4j 0.0582) 0.979 2 046 0 0060 1 780 377 

-‘04 HH2 37 24’ 440 15’C ‘866 03652i 0979 220’ (‘3Q55, 1906 ‘44 

: 135 HH3 44 ‘58 4o5 1500 ‘533 3Q47’, 0985 22’5 00050 827 

‘05 imH4 ‘9 l6 363 1384 ‘922 00635 3991 2324 03063 i39 -095 

)-‘O’ 1-01 40 128 340 1 384 837 00653) 0977 2 00 )0.0068 1 880 183 

108 P 11 156 39 1 274 205 02176 0953 1 870 001101 2 158 07’ 

09 CDI 40 86 308 1 373 1 059 0 1017) 0 936 2 119 0 0055) 1 772 4 

1 C 55 108 3 6 1 342 1 06 0.0591) 0.970 1 981 3065) 1 73 08 

X 09 m I e Igt 1, mmml ‘7 -‘ log Ape-tore enqth -nxm 

I 111 A) 369 108 45 1 415 0828 0010 ) 0974 .265 00013 1484 

Ill -IS 38 218 ‘395 1 459 9)4 0 380 0970 ‘91 0 025 1 

I 13 HS2 40 200 382 1 45’ 043 002 3 0°87 1.281 0002 08 

) 11” HH1 40 110 47 1 382 0 866 1 0”35 0 84 1 34 0 3 ‘ 4 7) 

I 11 )H 40 4 1 440 1.505 0 4 00’ 6 0.983 1 323 (0 0023 5 3 

118 3H3 45 loS 455 1 494 0896 00 82 0.991 1 328 00020 

I 17 994 20 6 5 363 1 382 0 946 00 58 0.993 1 238 (0 0025) 1 1 3 

) 118 P ‘1 40 28 343 1.384 0 850 0 0287) 0.979 ) 256 3 0030) 479 

) 113 PC1 ‘1 156 ‘3° 1 274 0.89 1 C 14) 0973 1 15) 00035) 109 

120 CD) 40 186 308 1 3’3 0.813 0410) 0.956 1 iO 0 0020) 350 

I ‘ 1 002 55 108 31 6 1 342 0.8 6 0 0226) 0.981 1 01 0 00251 654 

X=ug Se enqth urn, Y= 00 Body Or, ,v:. “9 

I 132 ALL. 369 13.8 455 1 415 2 00 0.04’S 0959 2268 ‘3 00451 2515 4.4? 

133 5’ 38 208 385 1 459 3 ‘4 0 1507, 0 960 2 396 0 3133, 3.233 ‘ 

134 HS2 43 20.0 382 1452 i’7 00883 3985 2401 00083 3124 ‘43 

1 ‘35 99’ 40 13.0 34.7 1 382 2489 0’240 0956 2.113 0.0128. 2604 320 

-‘36 992 40 24.’ 440 1 505 3 526 0 ‘3’3 0 985 2.423 0.0080 3.580 5.72 

I ‘37 990 45 158 45.o 1 494 3.154 0 0841 0 985 2.464 3.0090 3.202 2.41 

1 ‘38 HH4 20 165 36.3 1.382 3 ‘91 0 1548 0 979 2 158 0 0150 3 259 1 68 

i-39 PC’ 40 128 343 1 384 28’4 0.’094i 0974 0’86 0.0i’5 2.95’ 045 

‘47 PC2 1’ ‘56 23.9 ‘ 274 2943 031511 0952 963 00158) 3091 029 

-‘4’ CD) 40 186 308 1.373 266’ 01327 0956 22’2 00070’ 2183 -‘63 

I 147 002 55 3 8 31 6 342 2 541 0 0689’ 0 981 2.38’ ‘0 0078, 2 593 5.90

X g Shei lenqt r o ‘7 log Pro octod aroa or 

14 A 369 108 45.5 1.415 878 0010/ 0994 232 00(13) 

‘O 

3’ 

2 

2- 

5- 

62 

X g SF oIl igth mr ‘7 og (Shel copo ty nq 

144 ALL 369 108 45’ 1 415 2785 0.0306 0379 016 00033 

14’ HI? 38 208 385 1 459 3 069 0 1002 0 981 ‘ 147 0068 

1 (F (32 40 20.0 382 42 3.200 0 0696 0 991 3 138 (0 0053) 

14 HH 40 130 347 382 2613 C 0999 0973 ?890 001) 

148 HH2 40 24.1 440 1 505 3 11 00809 0987 3207 00068 

4) HH3 4 154 (4.5 1 494 3.1 2 C66 2 991 21 1 

150 HH4 20 iF 5 36 3 1 38? ‘.963)0 0853 0 993 2 943 (0.0083 

13 PCi 40 1 .8 343 1 384 2867 01 34 0976 2941 00108 

15g. PC? 1 156 239 1 274 3.13, 02408 0.374 2644 0.0123 

F C Dl 40 186 30.8 373 2 /52 0.’OSl 0 973 2 986 0 0055) 

15) C C? 5’ 10.8 31 6 3(2 2.780 0.0/04 0 983 2 800 0 30/8) 

0(3 

3 48 

0, 

3 90 

34 

1 1, 

14 

303 

44 

2 344 

3 34? 

03C 

2- 

3’ C 

183 

3 421 

3 13 

3 54 

X og Sholl lnqth orr ‘7 09 S i I dry wt Tig 

I 155 (131 38 208 385 1 459 2 00 0 1005 0 370 3 392 0.0067 

115 HS? 4 2)0 382 1 ‘52 2894 00857 0983 3 315 0.3)65) 

I lol 3H1 40 13.0 347 1 382 3292 0 00 ) 0383 3272 00103 

I 158 HH2 40 241 440 1.50o 2963 0 028 0 78 3o02 (00085 

I 139 HF? 45 158 455 .49 3016 0645 09°0 3309 0.0070 

I 0 HH4 ?) 165 363 1.382 3464) 1298) .988 3034 3.0125 

I 1 1 PC 1 128 34 3 384 3 148 0 783 0 989 3 264 0 0060 

I 152 P62 1 156 239 1 274 3223 03/4( 0.944 2904(00188) 

I 16 CD1 40 86 IC 8 1.373 3 084 0 149/) .958 3 236 0 0077 

I 164 CD? 55 108 316 1 342 3 10/ 00738) 0.985 3 12 00 80

APPFNDIXII 

Thaw leuteostoma 

,,.a 

14W 

bUCd V 

ix 

X Ms 

‘U; ice 

LcsstSq. neirReg 

do.. ‘ 

M’p I— 

* ‘1 

V aqAp tubesJtinw V og ipet sa a in 

IC ‘1 .3 6a 4 1337 1868(0.08’S 0979 5( U’ 130 1 

10 C 1, 24 ‘3 iae 2202(10595) 0996 lOS (3’S 13 34 

(3 D 1 41 286 1.31., 2)6900’88 996 2.’?.) 43 03 

lot C 1 .3 1292 204200W 0337 ‘8(1048 43 4 

.1’ 

X jr leigihimY ogAuettrtw.dttiin 

105 ‘C’ 3 16’ ‘8’ .33 orooao 0944 05’00058 U 

36 C 1 24 33 229 118600449) 0961 ‘5 004 1’ 43 

7 CL 1 86 .31. 15 5’) 7 64 3 1 

8 I 3’ 129. 1)aOO’ 995 0 3)6 II 44 

z 

‘I. 

X qAw c,igtt it V ogKo,i I 

In ,e n fl d3 uno ua’2 d 113 44 

C ‘ 1 .4 3. ‘29 12°2 0l44 0.37 )37J 0 13 3) 

I 41 ‘3 31’ 1468 33.33) 3702 38 03$ 0. 

‘ 

‘ .3 29.. 0)13 2 ‘14 32’ 01’) 371 

X cj Apire 9th uim,V 09 ptIkres mit 

II PC Ia 28.. 1337 1.106(0’38 0.714 0 3 0188 1549 2 2 

4 ‘1 24 126 09490133 3.815 0’ 58 1 79 

‘ 11 286 3 1 4(0223 12 7t00198 1.1 30 

2.1 323 2 40700729) 975 04 C 10’) 4 0 

‘C q lady I vt iq o S Idrywt ‘og) 

Il 3 1 796 2528 04 at’S’ 0.940 346.)0.’lO 16 lIC 

18 • 1 47 43 215.3 2... 36 029 3. 33* 13 61 

D? 3 a.) 135 .420 110938 0988 2860178 1 

X nP rt.Ju.iri. V 

o P a ,83 ‘4t 088(0427 09,, .“J( (3 4 

I 0 8 233 (.94300?. 993 ‘0? (070 ‘ 33 

I 1 4 4 3(100., 3376 9)0) 38a 3 

3 CL I 7 C 2382 08 00196 99 ‘Itt 0(0(3 0? 

S I ipGityng V og(Apecusirnim 

14 Pt ‘ 06 4016 326) 6310276 98 2 373 t

25 P02 3 29 ‘864 0108 -078 

2 898 0 651 0 3388’ 0 981 2 017 0 0 664 

25 002 0045 154 

21 274 69>5 3.151 0 630 (0 0100) 0 998 ° 85 (0 0 531 

X q S 1 1 apac ty mg Y log (3 c ip>ed volume rr g) 

(27 P( 1 2 706 4016 3260 0995(00378 0 787 3042 00 98 1.003 

28 P00 0692) 973 0.943 

3 229 1864 2 898 0 921 0 0 2 649 (0.0 93) 1 .73 

29 002 22 200 6925 ‘32 0088 > .036 0 0>90 0 997 2 v21 0 I 330 

X og Sheh capacty mg Y - log See’ dry WI.. 09 

I 30 PC’ 2’ 006 4>>16 04’ 0045( 0193 3 260 1 0 0 955 3 471 0 1 096 1 23 

3 3> P02 >9 1864 2 898 062 (0 3013) >> 062 3 J4> ‘0 925’ 1 194 >4 

32 CD 22 274 6925 3132 1 107 (00350) 0990 3284 (0 0160 11> 338 -y 

X -.. j (56 I 2 r g r or) 2 log (Ap’ to area m ii 

I 33 PCI 23 21 4 428 653 962 2.268 0090 1.481 1 0 1029) 0 (0 1 718 4 -‘ 

0 >4 PC? 16 >54 355 365 0983 99195 ‘.89 9: 

1 847 0.0a63> 2.321 

I) 25 CDI 22 ‘9 2 38.6 1.459 984 0:9873, 2.229 9.9080 026 

9.981 2 022 

1136 002 21 ‘58 474 1442 >7589947s1 0993 218000080. 1773 400 ‘ 

X og Shel engle rim Y og Ape lure lenqt( mm’ 

I 37 ALL 83 158 774 447 875 0147) 30? (017 1 0 0 0 989 3) 84 

I 38 PCi 3 ‘1 4 428 0.886 338 9033) 1 481 0 0361 0 983 1 9 3 

I 49 PC? 3 16.4 365 839 0366 229 (0.0040 1 

35 5 1 0 (0 0 988 1 4 

140 OD1 192 386 1 459 0.96’ (0 0320 0 989 316 (000 3 1 88 

‘4> 002 2’ >58 474 1442 0860 0.3230 0995 1292>00033. 600 — 

J 

X ‘og She.’ lv ir>” gt>>. V L09 Body ory WI. >19’ 

46 AL. 63 >58 4’ 2702 4 1 436 ‘96’’ 009. 0 977 2 393 ‘0 2 766 3 

(47 P01 72 ‘14 428 1481 262701313 0961 >528 09173 762 1 4 

(48 20’ 1 104 355 1 365 2.443 (0 ‘010) 0 356 2 163 (00 2.> 25>> 13 

49 CD? ‘2 58 474 1 415 76 . 3702 0 994 420 00 5 27 8 1 

X = 101 II >11 lengt mm) Y log (Pr0(3cted area, mm 

.150 AoL 33 >5.8 474 44’ 962 0 0>93 0096 2 3°3 0.003 1 56 

X og Stifl length CaoacIy. mm 2 og SOc mgI 

35> ALL 63 158 474 437 276a 1 9 3577 ‘25 000’S 2005 9 988 3 3.54 

1 P0 ol ‘4 428 1 482 2722.91351 0975 3261 (3ui”7) 2009 . 3 

53 PC’ 11 164 315 284> (02058 0972 .1898(991 923 

1 361 5 

,4 CD? 2 118 74 43’ 1 3) 2 777 3 619 0 35 3 7 1 38 

Shell leIgh mm) 2 log ISh II y NO mo 

log 

X . 

155 PCI 00 2>4 42(4 1481 3016,0139), 3900 3461 09103 0384 ‘>7. 

3 56 P02 15 16.4 355 1.365 0.915o 9.60 

3.041 0.19601 0 985 3 058 3 387 

I o7 CD> 22 92 386 459 3.348 ‘9.9159 2.50 

1 0 1749 0 974 3 347 3 431 

158 002 22 >58 4’4 1435 3>0900579 ‘996 32840.0095 312 2>0

APPENDIX III 

Monodonta labto 

-3 

L peeled Y a 

Man l’l a 

Least Sq. L mm Regr 

Spe 

X Axis 

Miri Mo’i Mp’rn 

Site N 

mit Y og Aperture area mm 

20 70 127 0981 2148 00604 0993 796 00053( 

3 163 

24 80 14 1 045 2138 00352( 099 1 336 (00030 

4 

21 3 4° 1 017 2043 0 560( 0992 1879 0(050( 

‘059 

30 85 178 11 2039 00305 0997 2 091 0 0023 

2 4, 

30 8 1 131 1 103 ° 16 0458 o 059 (0 00 7 

2076 00)53 

2003 0 035 

1 328 (0 0060( 

2 066 0 0033 

C94 2 / 

29 90 167 1 110 2 68 00620( 0989 

30 87 154 1 076 394(3 0525( 0990 3 4 

9 81 155 1 043 2 123 007 2( 0996 13? 

40 9? 157 1 108 2 145 0 0479 0 991 164 

X p (Ape tore lerrgtt 

10 

‘7 

op 

331 

48 

3 “ 

31) 

8 

34 

90? (15 

103 390 

04 HI-Il 

lOP HH 

106 HH3 

7 I-H4 

38 PC? 

11109 (D2 

66 

24 

3 ‘4 

I 1 

38 

2 10 

44 

I aO 

Ape I ire ength mm( V 09 (Aperture widt[ om( 

16 20 70 127 0981 1 012 00448 0983 9(9 )0040( 

HS1 ‘4 80 14a 1 045 1 066 00278 0993 1385 00023 

HS ‘4 73 149 1 017 1 094 00324 0990 0961 0 0(28 

IH1 30 8 a 1 8 17 1 003(0 0300 0 988 1 057 0 00?3( 

HH2 30 81 169 1 103 1 113 00360 0986 047 00010 

IH 29 °Q 67 110 1 070 00464( 33375 1 054 0 038 

5H4 10 82 14 076 0994 004?3( 0976 1 018 0 328 

PC’> 9 81 55 1 43 11 0090a( 098 0976 00(73 

D2 40 9? 157 1 108 1 076 0 )393( 0976 045 0) 28 

X oo 

1110 

11 

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