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Redalyc.Escape Tactics and Effects of Perch Height and Habituation On Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica Cooper, Jr., William E. Escape tactics and effects of perch height and habituation on flight initiation distance in two Jamaican anoles (Squamata: Polychrotidae) Revista de Biología Tropical, vol. 58, núm. 4, diciembre, 2010, pp. 1199-1209 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica Available in: http://www.redalyc.org/articulo.oa?id=44918952013 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Escape tactics and effects of perch height and habituation on flight initiation distance in two Jamaican anoles (Squamata: Polychrotidae) William E. Cooper, Jr. Department of Biology, Indiana University Purdue University Fort Wayne, Fort Wayne, IN 46805, USA; [email protected] Received 23-XI-2009. Corrected 10-VI-2010. Accepted 15-VII-2010. Abstract: Escape by Anolis lizards is influenced by microhabitats and fight initiation distance increases with predation risk. Differences in microhabitat use among ecomorphs affect escape behavior, but only two studies have reported ecomorphological differences in flight initiation distance among Greater Antillean species. I studied effects of predation risk and microhabitats on escape behavior by conducting field experiments using two species of anoles, Anolis lineatopus and A. grahami, on the campus of the University of the West Indies at Mona, Jamaica. Because ecomorphological variation of anoles has evolved independently within each island of the Greater Antilles, but relationships between ecomorphs and escape behaviors are poorly known, I character- ized microhabitat use and escape tactics, and determined relationships between flight initiation distance and two risk factors, habituation to human presence and perch height, in Anolis lineatopus, a trunk-ground anole and A. grahami, a trunk-crown anole. Sample sizes for A. lineatopus and A. grahami were 214 and 93, for microhabitat use and escape destinations, 74 and 34 for human presence and 125 and 34 for perch height. The two species occurred in similar microhabitats and exhibited similar escape tactics, but exhibited key differences expected for their ecomorphs. Both species were sighted frequently on the ground and on trees, but A. lineatopus were more frequently on ground and were perched lower than A. grahami. Both species escaped from ground to trees and when on trees hid on far sides and escaped without changing climbing direction with equal frequency. The frequency of fleeing upward was greater for A. grahami than A. lineatopus. Both species exhibited habituation by having shorter flight initiation distances in areas with more frequent exposure to people. In both species flight initiation distance increased as perch height decreased because, lizards had to climb farther to be out of reach when perched lower. The relationship between flight initiation distance and perch height may apply to other anole ecomorphs that flee upward when low perched on trees. Rev. Biol. Trop. 58 (4): 1199-1209. Epub 2010 December 01. Key words: ecomorphs, escape tactics, flight initiation distance, habituation, perch height, Polychrotidae, Squamata. Escape behavior has been studied exten- of predation risk in one or more species of sively in lizards, effects of predation risk fac- Anolis. Escape strategies and responses to tors and costs of fleeing have been examined predation risk may be affected by microhabitat more thoroughly in lizards than in any other use (Losos & Irschick 1996, Regalado 1998, taxonomic group. Surprisingly little informa- Irschick & Losos 1999, Schneider et al. 2000, tion is available about effects of factors that Irschick et al. 2005, Lattanzio 2009), which affect predation risk on the escape behavior differ greatly among the nearly 400 species of of anoles. However, flight initiation distance anoles (Losos 2009). (distance between the prey and an approach- Ecomorphs of Anolis lizards in the Greater ing predator when the prey begins to flee) has Antilles differ in microhabitat use and in aspects been shown to increase with several aspects of morphology that affect locomotion in their Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 58 (4): 1199-1209, December 2010 1199 respective microhabitats (Williams 1972, 1983, species that escape by climbing out of reach, Losos 2009). Such differences might affect but increased as perch height increased in A. methods and destinations of escape. Compara- krugi and A. pulchellus, which are grass-bush tive studies to assess similarities and differenc- anoles that escape by fleeing to the ground es in escape behavior among ecomorphs have when initially perched higher (Cooper 2006a). only recently begun (Cooper 2006a, Vanhooy- In the trunk-crown anole A. stratulus perch donck et al. 2007), but few species have been height did not affect flight initiation distance, studied and many risk factors that affect escape which is presumably another indication of its have been studied little or not at all. reliance on crypsis (Cooper 2006a). Consistent Among risk factors that have been studied with the findings for Puerto Rican arboreal in anoles are conspicuousness/detectability, lizards, flight initiation distance decreased as temperature, previous attack by a predator, perch height increased in the Hispaniolan trunk perch height and possibly habituation to preda- anole A. distichus (Cope 1862) (Schneider et tors. Flight initiation distance was longer in al. 2000). A. cristatellus (Duméril & Bibron 1837), A. Other indications that anoles assess risk in krugi (Peters 1876) and A. pulchellus (Duméril making decisions about antipredatory behavior & Bibron 1837) that were in the open, fully are that duration of tonic immobility is greater exposed to the experimenter’s view, than in in A. carolinensis (Voigt 1832) when a predator individuals that were partially concealed (Coo- is closer and shorter if a refuge is near (Hennig per 2006a), suggesting that detectability influ- et al. 1976). A single study failed find an effect ences their escape decisions. Heatwole (1968) of predation risk. Lattanzio (2009) reported attributed the shorter flight initiation distance that flight initiation distance is unaffected by of A. stratulus (Cope 1862) than A. cristatellus predator approach speed in A. humilis (Peters to greater crypsis in the former. This inter- 1863) and A. limifrons (Cope 1862). However, pretation is consistent with the finding that the methods were so severely flawed that this A. stratulus had the shortest flight initiation report should be disregarded. The biggest prob- distance and was the least conspicuous among lem was that effects of occupation of different seven species of anoles in Puerto Rico [the microhabitats and perch heights were ignored twig anole A. valencianni (Duméril & Bibron despite their strong effects on escape by anoles 1837) was not studied, Cooper 2006a]. In and other lizards (Regalado 1998, Schneider et Anolis lineatopus (Gray 1840) and A. gund- al. 2000, Stankowich & Blumstein 2005, Coo- lachi (Peters 1876) flight initiation distance per 2006b, Cooper & Wilson 2007, Vanhooy- decreases as temperature increases (Rand 1964, donck et al. 2007). Furthermore, in the only Cooper 2006a). This relationship reflects the other anole for which the relationship has been slower maximum running speed and there- studied, flight initiation distance is substan- fore, reduced escape ability of lizards at lower tially greater for fast than slow approaches in A. body temperature (Bennett 1980). Rapid learn- lineatopus (Cooper 2006b, unpublished data). ing about the dangerousness of a predator is I compare aspects of escape behavior in the implied by an increase in flight initiation dis- trunk-ground anole A. lineatopus and the trunk- tance by two Cuban anoles, A. sagrei (Duméril crown anole A. grahami (Gray 1845) from & Bibron 1837), and A. homolechis (Cope syntopic populations near Kingston, Jamaica. 1864) after they were captured a single time I examine differences in substrate use related (Regalado 1998). to the ecomorphs, compare escape destina- Perch height is an important risk factor tions and movements for lizards initially on that affected flight initiation distance in six of the ground or on trees. In some lizards, flight seven species of Puerto Rican anoles (Cooper initiation distance is shorter in areas where liz- 2006a). Flight initiation distance decreased ards are habituated to human presence (Burger as perch height increased in four arboreal & Gochfeld 1990, Cooper et al. 2003a, Cooper 1200 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 58 (4): 1199-1209, December 2010 & Whiting 2007a, Cooper 2009a). I report distance are routinely confirmed for factors the relationship between habituation to people affecting both predation risk and costs of flee- and flight initiation distance for both anoles. I ing in studies using human simulated preda- also examined the difference in perch height tors (Cooper 1999, 2000a, 2009a,b, Martín & between species and ascertained the relation- López 2003, Cooper et al. 2003a). ship between perch height and flight initiation Because human beings are not natural distance in each species. predators of anoles, escape responses specific to certain natural predators might not be elicit-
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