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Sexual dimorphism in Pomacea canaliculata (Gastropoda: Ampullariidae)

Article · December 1990

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Nestor J. Cazzaniga
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THE VELIGER
@ CMS, Inc., 1990
Thc Vcligcr 33(~):384-388 (Octobcr 1, 1990)

Sexual Dimorphism in Pomacea canaliculata

(Gastropoda: Ampullariidae)l
by

NÉSTOR J. CAZZANIGN
Departamento de Biología, Universidad Nacional del Sur,
8000 Bahía Blanca, Argentina

Abstract. The Argentine apple snail. Pomacca canaliculata (Lamarck, 1822), shows sexual di mor-
phism in the shape of the shell aperture and the operculum, whose relative widths are significantly
different between sexes. The shape of these structures is therefore a reliable means of externally sexing
the snails. ~-lales also bear a heavier shell than females. Consideration of sexual dimorphism will
imprm'e the predictive curves relating linear measuremt"nts and biomass. A test for sexual shell dimor-
phism would also enhance taxonomic undcrstanding of some conchological species.

II'\1'RODUC1'ION dissecled 10 ascertain lheir sexo Figure 1 shows lhe me;!-

While studying Pomacca canaliculata (Lamarck. 1822) a
shell dimorphism involving lhe shape of lhe apenure and
lhe operculum was observed. A similar observation was
made by HALE (1964) and DEMIAN & IBRAHIM (1972)
for lhe planorboid ampullariid Marisa cornuarietis (L.).
1'he females of bOlh species have a narrow aperture com-
pared wilh lhe more rounded one of lhe males. 1'0 an
experienced eye this dimorphism permils routine eXlernal
sexing of mosl specimens, wilh a high degree of reliabililY,
Though some males show a narrower aperture, expanded
perislomes are infrequent in females. In this paper a pre-
liminary analysis of the dimorphic condition is presenled
and its ecological and laxonomic derivations are discussed.
MA TERIALS AND ME1'HODS
Thirly-four live individuals of Pomacea canaliculata, col-
lecled al Palermo Park (Buenos Aires dt}', Argentina) in
February 1987, were used lo lesl a hypothesis of sex related
differences. AII specimens were within the size range of
mature adults (MARTIN, 1986).
Shells and opercula were measured wilh a Vernier cal-
iper lo the neareSl 0.01 mm, and ¡he animals were ¡hen
I Contribution No. 27 of the Laboratorio de Ecología Acuática,
Departamento de Biología de la Universidad I\acional del Sur.
This work was supported by a research grant from lhe Consejo
Nacional de Investigaciones Científicas y Técnicas (CONICET,
Argentina): PID 3915503.
l Researcher of the Comisión de Investigaciones Científicas de
la Provincia de Buenos Aires, Argentina.
surements taken. The aperture length was measured OJ¡
ilS largesl axis (i.e., obliquely lo lhe columellar axis) and
lhe aperture width was measured p.=rpendicular lo lhe
lenglh. This way of measuring fits .he operculum pro-
portions better than the custom of using lhe aperlure lenglh
parameter parallel to lhe largesl axis of lhe shell.
The shells were weighed lo lhe nearesl 0.01 g. For lhe
analysis of shell weight the largest female and the smallest
male were omitted from lhe calculations in order 10 study
the same size range for bolh sexes.
RESUL TS
Figure 2 shows two examples in which the relative widlh
of lhe aperture iIIuslrates lhe mentioned sexual dimor-
phism. The perimeler of lhe operculum is smoo¡h and
generally coincides wilh lhe shape of the aperlure in the
females. In the males il is often wider, showing' a sinuous
surface and an irregular ouler edge.
Table 1 summarizes the characteristics of lhe studied
malerial. There is no significanl diITerence between the
shell width-Ienglh ralio of males and females (t = 0.297,
P > 0.70, d.f. = 32) or hetween lhe apenure lenglh-shell
width ralio (t = 0.503, P > 0.60). The length of the body
whorl is almOSl equal 10 lhe shell widlh and is nol a
dimorphic characler (t = 1.300, P > 0.20). The widlh-
Icnglh ratio of lhe operculum was compared lo lhal of lhe
aperture using a paired t test, which gave a non-significant
value of 0.648 (1' > 0.40). The analysis could therefore
be carried oul eqUallY weIl on the peristome or on the
operculum.
N. J. Cazzaniga, 1990 Page 385

o
OLI

.80 .
. ..
..J
.76 . ....
1\ 1/ 11 1 .72
I 3:1
mi I "'- 11 11 I
~~
o o
~-!-i~----------------
.68
00 o o
08000 o o
.64 o o o

.60

30 40 50 60
sw shell wldth (mm)

Figure 1 Figure 3
Measurements ef Pomacea canaliculata shells. AL, apenure length; Scatter plet ef operculum width/operculum length versus shell
\ W, aperture width; B WL, body wherllength; SL. shelllength; width in the twe sexes of Pomacea canalicu/ala.
,-¡W. shell width.

The operculum width-Iength ratio is not significantly

dJ. = 2 and 28). The hypothesis of equality of lhe slopes
corrc:laled lo shel1 size (width) within this range. in either
sex: r = -0.1329 for lhe females; r = -0.2790 for the
was not rejected (F = 0.0015, P > 0.96), and it is lhus
lhe inlerCepts whieh are lhe souree of the dissimilarily.
males. 'The operculum ratio shows liule overlap between
The males have a heavier shel1 lhan the females, and
sexes (Figure 3), a critical leve! of discrimination being
the difference in shel1 weight-shel1 width ratio between
near 0.72. The comparison of the mean operculum ratios
the sexes is highly significant: l = 3.64, P < 0.01, d.f. =
yieldcd a highly significant diffcrence between sexes: l = 30).
10.45, P < 0.001.
Figure 4 shows the rcgression lines of shcll weighl (SlVe) DISCUSSION
as a funclion of shcll \Vidlh (S IV):
Males: SlVe = 0.271 36SIV - 5.72112 PO/TIllc/!acarw/rcu/ala is the sccond ampullariid species in
Females: SIV/! = 0.26807SIV- 6.71721 which sex dimorphism has becn reponed. BALE (1964)
described a similar feature in a Pueno Rican population
of Marisa comuarielis (L.), il1ustrating it by means of a
The analysis of covariance of the regressions over sexes scaller plOL She said lhat the difI'en:nce was "much lcss
indieatcd a significant diO'erence (1' = 7.694. P < 0.002, evidcnt" in a Florida populalion of lhe same species, lJut

- ......

Figur~ 2
A female (teft) and a male (righl) specimen of Pomaceacanalicu/ala illustrating sexual dimorphism in the relative
width ef the aperture.
Page 386 The Veliger, Vol. 33, No. 4

shell The expansion of lhe malc perislomc is probably relaled

welght lO lhe greal de\'elopmenl of lhe penial complex, whieh
. ¡;>;ivesrise lo an appreciable swelling under lhe righl sir:
(:j /rl
of lhe manllc cdgc in many ampullariids (HVLTON SCO'l .
9
1957; ANDREWS, 1964). Thus, lhe dimorphic conditio!l
6
I / may be common among olher NeOlropical apple snails.
When delermining sex on lhe basis of emply shells of
Pomacea canaliculala, il is useful 10 consider lhe aperlure
4 Icnglh obliquely, on its grealcst axis. The more cuslomary
vertical measuremenl would obscure lhe differences be-
lween sexes, which are clear in lhe operculum proportions.
GUEDES el al. (1981) sludied lhe correlalions among
35 45 55 linear parameters and lhe weighl of lhe shell and of the
shell wldth (mml soft parls in Pomacea canaliculala, and found lhallhe wea\.:-
est correlalions were lhose involving shell weighl. Becausc
Figure 4
they did not separale males from females. lhis low cor-
Rcgression lines of shell wcighl as a funClion of shell widlh in relalion could be due in parl lo the dimorphic condilion
Pnmacca cana/icu/ala.
of lhe shell weighl. The reproduclive eO'ort in lhis species
is higher lhan in olher gaslropods in temperale waters
(Eslebenel & Cazzaniga, unpublished data), and lhe rel-
no numcrical information was includcd to assess level of
alive weakness of lhe female shcll may therefore be relaled
signiflcance. DEMIAN& IBRAHIM(1972) carried out a more lo the expenditUre of calcium 10 build eggshells.
eXlensive sludy on lhe same species. They also found lhat Thcre are se\'eral conccivable ccological and laxonomic
shell aperture prO\'ides the mosl significant sex-rclaled ml1sequel1ces of shcll dimorphism.
differences, and delected (ess obvious signs of dimorphism
on olher shell and pigmenlalion characlers. \\'e found sex- Ecology: P07llQccncannliculatn and rclated snails are ",ilh.
rclated expanded apcrtures in P. canaliculata populations srread in lhe NeOlropical Region (CAZZANIGA,1987). The}'
from difI'erent localities. conslilule the main diel of some birds (SNVDER & KALE,
The opercular ralio for some Pomacea canaliculala from 1983; BOURNE, 1983), and are ealen by caimans (DIE-
Brazil is 61.1 lO 79.4. with a general mean of 67.36 :t FENBACH, 1979), lurlles, and other native verlebrales
3.90 (n = 87) (GUEDESel al., 1981). These values are not (RUSSELL, 1972). By assessing lheir biomass on lhe basis
differenl from lhose presented in Table 1 (t.= 1.08, P > of linear shell measurements il is possible bOlh lo evaluale
0.2, dJ. = 119), suggesting a high degree of similarit)' lhe energy budgel of live populalions Wilhoul employing
belween lhe Argentinian and Brazilian populalions. destruclive teehniques and lo eSlimate the biomass con-

Table 1
Shcll measuremenls and weighls of lhe studied specimens of POInacea canaliculala (Laman'k, 1822) from La Plala.

Females (n = 21) Males (n = 13)

Min Max Mean :t SD Min Max Mean :t SD

42.8 61.7 48.21 :t 4.53 39.0 53.8 47.85 :t 5.00

Shell 1cngth (SL)
Shell widlh (S\\') 37.7 57.8 43.13 :t 4.79 33.2 48.0 42.53 :t 4.44
36.7 59.5 42.43 :t 5.04 34.1 47.8 42.45 :t 4.21
Body whorllenglh (BWL)
31.3 49.5 35.79 :t 4.17 28.4 40.0 35.08 :t 3.37
Aperture length (AL)
2\.3 33.9 24.47 :t 2.88 20.5 29.0 25.65 :t 2.7.
Aperlure width (A W)
26.2 44.0 30.95 :t 3.87 26.1 35.:\ 30.97 :!: 2111;
Opcrcu1urTI Icnth (01.)
17.0 29.2 20.40 :!: 3.27 18.8 2(,.0 22.41 i 2.22
Uperculum widlh (UW)
2.83 6.97 4.65 :t 1.04 3.67 8.38 6.03 :t 1.38
Shell weightt (SWe)
82.68 97.91 89.40 :t 3.71 80.16 96.47 88.98 :t 4.48
100 SW /SL
92.16 103.66 98.38 :t 3.37 94.87 ]06.55 99.93 :!: 3.38
lOa BWL/S\\,
78.53 87.02 82.97 :t 2.65 77.38 88.35 82.62 :t 3.09
100 AL/SW
64.52 73.99 68.40 :t 2.47 70.14 77.60 72.99 :t 2.16
tOOAW/AL
63.64 72.22 66.78 :t 2.35 70.59 77.78 75.35 :t 2.05
100 OW /OL*
9.13 13.34 10.86 :t 1.78 9.84 20.24 13.83 :t 2.86
100 SWe/SWt

t Calculaled Irorn 20 females ami 12 males.

* General mean (n = 34): 68.30:t 5.14.
 .'J.J. Cazzaniga, 1990
surncd by prcdators who Icave the empty shells. However,
females havc heavier soft parts Ihan males (GUEDESel al.,
1981). llecause the intercepts 01' lhe biomass regressions
01'both sexes are significantly different, BOURNE& BERLIN
(1982) stated that Han overall equation, although possibly
still useable, would show a consistent bias in prcdiction
dcpending on the sex 01'the individua!." We suggest that
il is possib1e to identify reliably the sex 01' eaeh specimcn
iI1d 10 use differentiatcd rcgression equations 10 predict
;he biomass.
GUEDES el al. (1981) delined an operculum product
(lcngth x widlh) thal they used as an independent variable
in a regression on living weight. This parameter seems to
be inadequale, considering the observed sexual differenees
in opereulum widlh. A fema1e will have a lower operculum
product but a grealcr biomass than a mal e 01' the same
opereulum lenglh, and the dispersion from an overall re-
gression eurve will therefore be greater.
The identillcation 01' sex in empty shells should makc
.: possible LOeSlablish whether predalion by kites and
limpkins cxerts the same pressure on both sexes, since
random capture on the normally 1:1 sex ratio 01'Pomacea
populations (MARTIN, 1984)3 would yicld an equivalent
ratio in the mounds 01'shells al feeding sites. Should this
not be the case, then difTerential activities in either sex-
as have already bcen found in olher prosobranchs (RIDI
& ARTER, l 986)-and/or {)lher factors affecling lheir cap-
lllrability should be induded in .lhe analysis.
Taxonomy: Some delailed slwlies on lhe analOrny 01'sev-
''I'al spe!'Íes are availahlt', hnl IllOsl alllPllllariids have slill
nol been wcll-dclincd (l'AIN, 1«)72). 'l'hollgh shells are
variable, the form 01'lhe apcrtu/'e has frequently been lIsc:d
as a diagnostie charaeter. At least in two genera, some
taxonomic doubts can be dispellc:d lhrough an analysis 01'
sexual dimorphism in the shcll.
PILSBRY(1933) described lvlari:;aplaTlug)'ra, from Brazil,
as differing from Ihe type species, ¡'vI.cornuarietis, based
on, among other things, the fact that "the last whorl and
the aperlllre are Icss expanded in the upper ami hasal
parts" in the former. Taking into account the results of
HALE (1964) and DEMIAN & IBRAHIM (1972), as well as
our present findings, lhis diagnostic charaeteristic of Al.
I)lanogyra could corrcspond to female shells. OLAZARRI
(1977) suggcsted thal the two species eould be synonyms.
Also, a significant variation in the aperture was men-
lioned in the supraspecies Pomacea canaliculala (sensu
CAZZANIGA,1987). For example, P.leuior (Sowerby, 1909)
was difTerentiated by its cxpanded outer fip. GEIjSKES &
PAIN (1957), studying P. dolioides (Reeve, 1854) from Su-
rinam, found that "in some specimens the perislOme is
wider than normally as in those described by him IVem-
hout, 1914] as leuior." A similar observation was also made
} It should be nOled that the sex ratio balance was tipped in
favor of the females in the populations of Marisa cornuarielis
studied by DEMIAN & IBRAHlM (1972).
Page 387
hy PAIN (t 960) with rcgard to P. lineala (Spix in Wagner,
1827) from llrazil, in whose synonymy he placed P. leuior.
In both cases the specimens appear to have been male.
A series of similar cases is easily found in the already
confused scheme of Pomacea forms. To what extent are
many of the mentioned differences among nominal species
dlle 10 an unnOliced sex dimorphism? Knowledge of many
South American species of Pomacea is based on nothing
more than a few empty shells, without any assessment 01'
variability. A test for the presence of a dimorphic condition
in the other apple snails would improve the understanding
of the conchological species: several putat'ive species could
prove 10 be sex-based synonyms.
LITERATURE CITED
ANDREWS, E. B. 1964. The funclional anatomy and histology
of the reproductive system of some pilid gastropod molluscs.
Proceedings of the Malacological Society of London 36: 121-
140.
BOURNE,G. R. 1983. Snail kile feeding ecology: some correlates
and tests of optimal foraging. Doctoral Dissertation, Uni-
versity of Michigan. Ann Arbor. xi + 110 pp. '
130URNE, G. R. & J. A. BERL1N. 1982. Predicting Pomacea
dolioides (Reeve) (Prosobranchia: Ampullariidae) weighls
from linear measurements of their shclls. The Veliger 24(4):
367-370.
CAZZANIGA.N. J. 1987. Pomacea callulicula/a (Lamarck, 1801)
en Calamarca (Argentina) y un comentario sobre I'IlIIpullllria
ca/amarcensls Sowerby, 1874 (Gastropoda: Ampullariidae).
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DEMIAN. E. S. & A. tl.1. IORAIIIM. 1972. Sexual tlinlllrphisrn
,lIul S('Xr,uio in Ihe snail Mllllw (flIIIlIIlTlt'/II (L.). Zoologil'al
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\)IEFENBACII,C. O. DAC. 1979. Ampullariid gastropods: staple
food of Caimall la/ims/ris? Copeia 1: 162-163.
GEIjSKES, D. C. & T. PAIN. 1957. Suriname freshwaler snails
of Ihe genus P,¡macea. Studies on lhe Fauna of Suriname
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GUEDES, L. M. L. A., A. M. C. FloRI & C. O. DAC. DIEFENBACH.
1981. Biomass estimation from weight and linear parame-
ters in the apple snail, Ampullaria canaliculala (Gastropoda:
Prosohranchia), Comparative Biochemislry and Physiology
68A:285-288.
HALE, M. C. 1964. The ecology and distribution of the intro-
duced snail, .\larisa cornuarielis (Ampullariidae), in South
Florida. M.Sc. Thesis, University of Miami, Coral Gables.
115pp. .
HYLTON SCOTr. M. 1. 1957. Estudio morfológico y taxonómico
de los ampulláridos de la República Argentina. Revisla.
Museo Argentino de Ciencias Naturales "Bernardino Ri-
vadavia." Zoología 3(5):233-333.
MARTIN, S. M. 1984. Contribución al conocimiento de la biolo-
gía de la familia Ampullariidae (Mollusea, Gastropoda) en
el Río de la Plata. Doctoral Dissertation No. 431, Facultad
de Ciencias Naturales y Museo, Universidad Nacional de
La Plata, La Plata. t 49 pp.
MARTIN, S. M. 1986. Ciclo reproductivo de Ampullaria canalic-
ulala (Gastropoda: Ampullariidae) en el área rioplatense.
Neotropica 32(88): 171-181.
OLAZARRI, J. 1977. Informe preliminar sobre moluscos del área
de inOuencia de la futura represa de Salto Grande. In: Reu-
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PAIN.T. 1960. PQ7Tlacca (Ampullariidac) of the Amazon River
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luscan Studics 52:91-96.
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to Paraguay. New Mexico Slalc University. 9 pp.
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by snail kites in Colomhia. Thc Auk 100:93-97.