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Bull. Chicago Herp. Soc. 26(9):197-198, 1991 Cannibalism in the Ctenosaur Lizard, Ctenosaura similis, in Costa Rica José M. Mora Welder Wildlife Refuge P.O. Box 1400 Sinton, TX 78387 Cannibalism is a natural behavioral phenomenon well ‘known in many groups (Crump, 1990; Fox, 1975; Polis, 1981), ‘but has been examined only recently in reptiles (Mitchell, 1986), Numerous descriptions of this behavior have been ‘published, including many observations of lizard species in both field and captive environments, which suggest that can- nibalism is not an aberrant isolated occurrence in lizard pop- lations (Sugerman and Applegarth, 1980). In a review of cannibalism in reptiles, Mitchell (1986) mentioned the oceur- rence ofthis behavior in the field in the ctenosaur, Crenosaura “Sinilis. His information was based on a report in Fitch and Henderson (1978). These authors found a partly digested tail ‘ofa small ctenosaur in the stomach of an adult. They also found one adult ctenosaus with a ctenosaur egg in its stomach and another adult which had eaten three such eggs. Although stomach contents often serve as the primary evi dence for cannibalism, the problem with such data is in deter- ‘mining how the animals came to be ingested (Schwartz, 1985). ‘They may have been dead when encountered or ingestion ‘may have resulted from an active attack by one animal on the ‘other (Schwartz, 1985), Both situations, the eating of a dead conspecific and the killing and eating of a conspecific have been referred to as cannibalism in the literature (McNichol, 1977), The first situation, however, should be termed scar enging (McNichol, 1977). (On the other hand, as Mitchell (1985) pointed out, the be- havioral act of oophagy (the consumption of eggs or embryos ‘ya conspecific) often has been called cannibalism. However, “intraspecific oophagy isa specialized behavior involving df= ferent stimuli and occurs most often in female reptiles eating their own eggs or embryos" (Mitchell, 1986). Cannibalism is applicable only in cases of intraspecific predation (Fax, 1975). Following these definitions, only the finding of the ‘tenosaurtal by Fitch and Henderson (1978) could be called Cannibalism, although even this could bea case of carrion eating. Here [report that cannibalism may bea common behay- forin etenosaurs, based on observations made atthe Rafael Lucas Rodriguez Wildlife Refuge (Palo Verde), Guanacaste Province, Costa Rice (10°21°N; 85°21° W). At least three instances of cannibalism by ctonosaurs under diferent con- ditions ae discused. ‘A group of ctenosaurs used to spend the night underneath the floor of an old house in Palo Verde. The lizards were ‘normally found around the house (Figure 1) and fled under it when people approached. Probably some females nested there since many hatchlings emerged fom this place during the 1984 hatching season, Unfortunately the house burned in January 1985 and thereafter the concitions were not favor- able for ctenossurs. In May 1984 I observed an adult ctenosaur catching and eating what I presumed to be a hatchling ctenosaur that had ‘emerged from under this house. A bit later I released a hatch- ling etenosaur close to an adult female who caught it and ate it (Figure 2), A second hatchling ctenosaur was offered to ‘another adult female who captured and ate it immediately. Even though many hatchlings as well as adults were observed at this site, no other etenosaurs were observed catching hatch- lings. I offered a skink (Mabuya uoimarginata) to an adult ‘male ctenosaur but it did not take it, nor did an adult female ctenosaur feed upon a hatchling green iguana (guar iguana) that was offered. Lizards are known to be part of the cteno- saur’s diet (Fitch and Henderson, 1978). In May 1985 ob- served a young adult ctenosaur pursuing, attacking and eating, a hatchling ctenosaur in che wild, In June 1986, while con- ‘Cucting a study of social behavior in ctenosaurs, I placed three Figure 1. The old house at Palo Verde, with ctenosaurs basking in the foreground, Figure2. An adult female ctenosaur, Cenosaura sis, with the head of a conspecific hatchling protruding from its mouth. 197 cone year-old ctenosaurs in an enclosure with 10 hatchling, crenosaurs. Three hatchlings were missing a week later, but they may have escaped. Three more hatchlings were missing the next week. I flushed the stomach contents from the three ‘yearling ctenosaurs and found remains of hatchlings in two of the three animals. Petzold (1982) discussed cannibalism from the perspective of overerowding in captivity. In my captive study the canni- balism observed was more likely de to overcrowding rather than a food shortage, since food was apparently sufficiently abundant. A similar situation may have occurred under natur- al conditions atthe old house in Palo Verde, where overcrowd- ing was potentially present. Ctenosaurs probably were attract- ed to the house because they could find abundant retreat and resting sites there. In the wild, the yearing ctenosaur was observed to attack its victim. This suggests that cannibalism by etenosaurs may also represent opportunistic predation Normally, there is habitat separation of young and adult ‘tenosaurs that would reduce predation pressure on young by adults Fitch and Henderson, 1978). In terms of inclusive fitness, it might be disadvantageous 10 eat a close relative (Crump, 1983), However, only 2 very ‘small increase in individual fitness of the cannibalizing ind vidual would be necessary t0 maintain cannibalism, even if {ull siblings are eaten (Cramp, 1983; Bickwort, 1973). ick- ‘wort (1973) pointed out that cannibalism must be favored especially during the time in the life eycle when mortality ‘and nutritional benefits are high. The ctenosaur’s hatching 198 season coincides with the beginning of the rainy season, a time of year when food is probably still scarce. Hatchlings ‘would be a good altemative food source for some adult ‘tenosaurs, especially when the dry season is extended. Hatch- lings that emerge at this time have lower survival probabili- ties, mainly because the insects available as prey decline in density during the dry season (Janzen, 1983). Insects are ‘their main food source during that critical stage (Mora, 1986). ‘These two conditions, nutritional benefit for adults and a ‘high hatchling mortality, would favor the occurrence of ean- nibalism in ctenosaurs. Later, the probability of cannibalism decreases due to the habitat separation already mentioned. Cannibalism is a natural phenomenon that is beneficial to some members of a population (Crump, 1990), and is subject to natural selection (Fox, 1975). This behavior could offer a number of advantages for a predator in natural populations by, among other things, eliminating potential competitors (Gogerman and Applegarth, 1980). It could also benefit some ‘members of the population during lean periods, when mortal- ity of young animals is high anyway. These factors may ex plain the occurrence of cannibalism in the ctenosaur. [My thanks £0 A. C. Chaves for her assistance in the fel M. DiMare, C. MeDanough and W. Grant made helpful com- rents on the manuscript. Financial support was provided by the Universidad Nacional (UNA), Heredia, Costa Rica, the ‘World Wilelife Fund (WWF), and the Orgenization for Trop- ical Studies (OTS). Literature Cited ‘Crump, M.L. 1983. Opportunistic cannibalism by amphibian larvae in temporary aquatic environments. Am, Nat. 121(2):281-287. 1990, Possible enhancement of growth in tadpotes through cannibalism. Copeia 1990(2) 560-64. icovort, KR. 1973. Cannibalism and kin selection in Labdomera clivicols (Coleoptera: Chysomelidae). Am. Nat 10(955):452-453, Fitch, H.S, and R, W. Henderson, 1978, Ecology and exploitation of Crenasaura similis. Uni. Kansas Si. Bull. 51(15):483500. Fox, LR. 1975. Cannibalism in natral populations. Ann, Rev. Heol. Syst. 687-106 Janzen, D.H. 1983, Insects, Pp. 612-645, fn D. H Jenzen, editor, Costa Rican natural history. Chicago: University of Chicago Press. MeNicholl,M.K, 197, Usage ofthe terms cannibalism” and “scavenging” in ecological iterture. Can, Fielé-Noturalist 1:6 Mitchel J.C. 1986. Cannibalism in repiles: A worldwide review. SSAR Herp. Circular 15 Mora, J.M. 1986. Alimentacion y crecimiento corporal del garrobo, Ctenosaure similis Gray, en su primer afo de vida. Unpubl. MS ‘Thesis. University of Costa Rica Petzold, H-G. 1982. Aufgaben und Probleme der Tiorgartnerei bei der Erforschung der Lebenstusserungen der niederen Amnioten (Reptilien). Milu 5(4/5):485-786, Polis, G.A. 1981, ‘The evolution and éynamies of intraspecific predation. Ann. Rev. Heol. Syst. 12:725-251 ‘Schwartz, J.M. 1985, Life history notes: Thannophissrtalissinalis (eastern garter snake) cannibalism. SSAR Herp Rev. 16(4):112. Sugerman, R.A.and JS. Applegarth. 1980, An instance of natural cannibalism by Uma n, notata Baird, SSAR Herp. Rev. 11(4)90.

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