Sexual behaviour of immature male eastern mosquitofish: a ... - CPRG

Sexual behaviour of immature male eastern mosquitofish: a ... - CPRG Sexual behaviour of immature male eastern mosquitofish: a ... - CPRG

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sexual behaviour of immature male mosquitofish 735 adult males did not change signficantly between treatments (with one or four adult females). None the less, the body size attained at maturation by males was negatively correlated with the frequency of copulatory attempts, that, in turn, depended on the sex ratio in the aquarium where they grew. These results therefore suggest that social control mechanisms, at least in G. holbrooki, may be more complicated, and perhaps more precise, than previously thought. In fact, it could be argued that, for a maturing male, aggressive behaviour by adult males may not be a precise measure of his possible reproductive success. During the breeding season, adult males spend most of their time around females, attempting to copulate or trying to keep other males away from them (McPeek, 1992; Bisazza & Marin, 1995). For this reason, the best way for an immature male to test his potential reproductive success is by trying to copulate with females, and by gaining information about copulations attempted, frequency of successful copulations and of attacks received. Preliminary studies (Bisazza, unpublished data) suggest that immature males also show sexual behaviour in other poeciliids. In three species, Belonesox belizanus (Kner), Girardinus falcatus (Eigenmann) and Phalloceros caudimaculatus (Hensel), sexual behaviour was observed at an early stage of gonopodium development. Conversely, in Poecilia reticulata (Peters) and in Heterandria formosa (Agassiz) males started to be sexually active from 5 to 10 days after the gonopodium was completely developed. Histological examination of the gonads showed that mature sperms were present in P. reticulata and in H. formosa, but not in G. falcatus and P. caudimaculatus. Interestingly, body size is the major factor determining the outcome of male contests in G. holbrooki and in the other four species that we found to have precocious sexual behaviour (Bisazza & Marin, 1995), but not in the two species lacking it (Kodric-Brown, 1983; Zannini, 1994). We would like to thank A. Kodric-Brown for helpful suggestions to a previous version of the manuscript. This research was partially supported by a MURST 60% grant to A. Bisazza. References Andersson, M. (1983). On the functions of conspicuous seasonal plumages in birds. Animal Behaviour 31, 1262–1264. Andrew, R. J. (1966). Precocious adult behaviour in the young chick. Animal Behaviour 14, 485–500. Andrew, R. J. (1975). Effects of testosterone on the behaviour of the domestic chick. II. Effects present in males but not in females. Animal Behaviour 23, 139–155. Bisazza, A. (1993). Male competition, female mate choice and sexual size dimorphism in poeciliid fishes. Marine Behaviour and Physiology 23, 257–286. Bisazza, A., Marconato, A. & Marin, G. (1989). Male mate preferences in the mosquitofish Gambusia holbrooki. Ethology 83, 335–343. Bisazza, A. & Marin, G. (1991). Male size and female mate choice in the eastern mosquitofish (Gambusia holbrooki: Poeciliidae). Copeia 1991, 730–735. Bisazza, A. & Marin, G. (1995). Sexual selection and sexual size dimorphism in the eastern mosquitofish Gambusia holbrooki (Pisces: Poeciliidae). Ethology, Ecology and Evolution 7, 169–183. Borowsky, R. L. (1973a). Relative size and the development of fin coloration in Xiphophorus variatus. Physiological Zoology 46, 22–28. Borowsky, R. L. (1973b). Social control of adult size in males Xiphophorus variatus. Nature 245, 332–335.

736 a. bisazza et al. Clutton-Brock, T. H. (1991). The Evolution of Parental Care. Princeton, NJ: Princeton University Press. Constantz, G. D. (1975). Behavioural ecology of mating in the male Gila topminnow, Poeciliopsis occidentalis (Cyprinodontiformes: Poeciliidae). Ecology 56, 966–973. Dulzetto, F. (1931). Sviluppo e struttura del gonopodio di Gambusia holbrooki. Pubblicazioni della Stazione Zoologica di Napoli 11, 1–85. Dulzetto, F. (1933). La struttura del testicolo di Gambusia holbrooki (GRD) e la sua evoluzione in rapporto con lo sviluppo del gonopodio. Archivio Zoologico Italiano 19, 405–431. Endler, J. A. (1982). Convergent and divergent effects of natural selection on colour patterns in two fish faunas. Evolution 36, 178–188. Farr, J. A. (1980). The effect of juvenile social interaction on growth rate, size and age at maturity, and adult social behaviour in Girardinus metallicus Poey (Pisces: Poeciliidae). Zeitschrift für Tierpsychologie 52, 247–268. Farr, J. A. (1984). Premating behaviour in the subgenus Limia (Pisces: Poeciliidae): sexual selection and the evolution of courtship. Zeitschrift für Tierpsychologie 65, 152–165. Farr, J. A., Travis, J. & Trexler, J. C. (1986). Behavioural allometry and interdemic variation in sexual behaviour of the sailfin molly, Poecilia latipinna (Pisces: Poeciliidae). Animal Behaviour 34, 497–509. Ford, C. S. & Beach, F. A. (1951). Patterns of Sexual Behaviour. New York: Harper & Brothers. Gronell, A. M. (1989). Visiting behaviour by females of the sexually dichromatic damselfish, Chrysiptera cyanea (Teleostei: Pomacentridae): a probable method of assessing male quality. Ethology 81, 89–122. Ibrahim, A. A. & Huntingford, F. A. (1992). Experience of natural prey and feeding efficiency in three-spined sticklebacks (Gasterosteus aculeatus L.). Journal of Fish Biology 41, 619–l625. Kodric-Brown, A. (1983). Determinants of male reproductive success in pupfish (Cyprinodon peconensis). Animal Behaviour 31, 128–137. Krumholz, L. A. (1963). Relationships between fertility, sex ratio, and exposure to predation in populations of the mosquitofish Gambusia manni Hubbs at Bimini, Bahamas. Internationale Revue der gesamten Hydrobiologie 48, 201–256. Liley, N. R. & Seghers, B. H. (1975). Factors affecting the morphology and behaviour of guppies in Trinidad. In Function and Evolution in Behaviour (Baerends, G. B., Beer, C. & Manning, A., eds), pp. 92–118. Oxford: Oxford University Press. Marconato, A. (1991). Life history tactics and population structure in freshwater fish species. In Form and Function and Zoology (Lanzavecchia, G. & Valvassori, R., eds). Selected Symposia and Monographs U.Z.I., pp. 379–393. Modena: Mucchi. Martin, R. G. (1975). Sexual and aggressive behavior, density and social structure in a natural population of mosquitofish, Gambusia affinis holbrooki. Copeia 1975, 445–454. Mason, W. A. (1965). The social development of monkeys and apes. In Primate Behavior, Field Studies of Monkeys and Apes (DeVore, I., ed.), pp. 514–543. New York: Holt, Rinehart & Winston. McPeek, M. A. (1992). Mechanisms of sexual selection operating on body size in the mosquitofish (Gambusia holbrooki). Behavioural Ecology 3, 1–12. Schultz, R. J. (1961). Reproductive mechanisms of unisexual and bisexual strains of the viviparous fish Poeciliopsis. Evolution 15, 302–325. Sohn, J. J. (1977). Socially induced inhibition of genetically determined maturation in the platyfish, Xiphophorus maculatus. Science 195, 199–201. Stone, C. P. (1940). Precocious copulatory activity induced in male rats by subcutaneous injections of testosterone propionate. Endocrinology 26, 511–515. Zannini, L. (1994). Selezione sessuale e dimorfismo sessuale nei pecilidi. Doctoral Thesis, University of Padova.

sexual <strong>behaviour</strong> <strong>of</strong> <strong>immature</strong> <strong>male</strong> mosquit<strong>of</strong>ish 735<br />

adult <strong>male</strong>s did not change signficantly between treatments (with one or four<br />

adult fe<strong>male</strong>s). None the less, the body size attained at maturation by <strong>male</strong>s was<br />

negatively correlated with the frequency <strong>of</strong> copulatory attempts, that, in turn,<br />

depended on the sex ratio in the aquarium where they grew. These results<br />

therefore suggest that social control mechanisms, at least in G. holbrooki, may be<br />

more complicated, and perhaps more precise, than previously thought. In fact,<br />

it could be argued that, for a maturing <strong>male</strong>, aggressive <strong>behaviour</strong> by adult <strong>male</strong>s<br />

may not be a precise measure <strong>of</strong> his possible reproductive success. During the<br />

breeding season, adult <strong>male</strong>s spend most <strong>of</strong> their time around fe<strong>male</strong>s, attempting<br />

to copulate or trying to keep other <strong>male</strong>s away from them (McPeek, 1992;<br />

Bisazza & Marin, 1995). For this reason, the best way for an <strong>immature</strong> <strong>male</strong> to<br />

test his potential reproductive success is by trying to copulate with fe<strong>male</strong>s, and<br />

by gaining information about copulations attempted, frequency <strong>of</strong> successful<br />

copulations and <strong>of</strong> attacks received. Preliminary studies (Bisazza, unpublished<br />

data) suggest that <strong>immature</strong> <strong>male</strong>s also show sexual <strong>behaviour</strong> in other<br />

poeciliids. In three species, Belonesox belizanus (Kner), Girardinus falcatus<br />

(Eigenmann) and Phalloceros caudimaculatus (Hensel), sexual <strong>behaviour</strong> was<br />

observed at an early stage <strong>of</strong> gonopodium development. Conversely, in Poecilia<br />

reticulata (Peters) and in Heterandria formosa (Agassiz) <strong>male</strong>s started to be<br />

sexually active from 5 to 10 days after the gonopodium was completely<br />

developed. Histological examination <strong>of</strong> the gonads showed that mature sperms<br />

were present in P. reticulata and in H. formosa, but not in G. falcatus and<br />

P. caudimaculatus. Interestingly, body size is the major factor determining the<br />

outcome <strong>of</strong> <strong>male</strong> contests in G. holbrooki and in the other four species that we<br />

found to have precocious sexual <strong>behaviour</strong> (Bisazza & Marin, 1995), but not in<br />

the two species lacking it (Kodric-Brown, 1983; Zannini, 1994).<br />

We would like to thank A. Kodric-Brown for helpful suggestions to a previous version<br />

<strong>of</strong> the manuscript. This research was partially supported by a MURST 60% grant to<br />

A. Bisazza.<br />

References<br />

Andersson, M. (1983). On the functions <strong>of</strong> conspicuous seasonal plumages in birds.<br />

Animal Behaviour 31, 1262–1264.<br />

Andrew, R. J. (1966). Precocious adult <strong>behaviour</strong> in the young chick. Animal Behaviour<br />

14, 485–500.<br />

Andrew, R. J. (1975). Effects <strong>of</strong> testosterone on the <strong>behaviour</strong> <strong>of</strong> the domestic chick. II.<br />

Effects present in <strong>male</strong>s but not in fe<strong>male</strong>s. Animal Behaviour 23, 139–155.<br />

Bisazza, A. (1993). Male competition, fe<strong>male</strong> mate choice and sexual size dimorphism in<br />

poeciliid fishes. Marine Behaviour and Physiology 23, 257–286.<br />

Bisazza, A., Marconato, A. & Marin, G. (1989). Male mate preferences in the<br />

mosquit<strong>of</strong>ish Gambusia holbrooki. Ethology 83, 335–343.<br />

Bisazza, A. & Marin, G. (1991). Male size and fe<strong>male</strong> mate choice in the <strong>eastern</strong><br />

mosquit<strong>of</strong>ish (Gambusia holbrooki: Poeciliidae). Copeia 1991, 730–735.<br />

Bisazza, A. & Marin, G. (1995). <strong>Sexual</strong> selection and sexual size dimorphism in the<br />

<strong>eastern</strong> mosquit<strong>of</strong>ish Gambusia holbrooki (Pisces: Poeciliidae). Ethology, Ecology<br />

and Evolution 7, 169–183.<br />

Borowsky, R. L. (1973a). Relative size and the development <strong>of</strong> fin coloration in<br />

Xiphophorus variatus. Physiological Zoology 46, 22–28.<br />

Borowsky, R. L. (1973b). Social control <strong>of</strong> adult size in <strong>male</strong>s Xiphophorus variatus.<br />

Nature 245, 332–335.

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