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Population structure, activity pattern, and microhabitat use of Phelsuma klemmeri at Mandrozo,

Lovasoa M. S. Rakotozafy en exergue la structure de la population, le rythme Mention Zoologie et Biodiversité Animale, Faculté d’activité journalière, ainsi que le mode d’utilisation des Sciences, Université d’, BP 906 du microhabitat. Antananarivo 101, Madagascar Au total, 93 individus ont été recensés, le long du E-mail: [email protected] 600 m de transect dont 65 adultes, 23 subadultes et cinq nouvels éclos. Les résultats ont montré que la répartition spatiale de cette espèce est conditionnée Abstract par la répartition des bambous. L’abondance relative We carried out a research on the yellow-headed de ce est aussi positivement corrélée avec day gecko, Phelsuma klemmeri, in the Mandrozo le nombre de bambous morts. La sex-ratio est protected area, a recently reported new locality for fortement biaisée en faveur des mâles. Phelsuma this , to assess its population status and klemmeri possède un dimorphisme sexuel puisque propose conservation measures. Visual encounter les mâles sont plus grands que les femelles. En surveys and bamboo excavation along six transects outre, son rythme circadien est en deux modes : provided information on population structure, daily l’ est très actif le matin et vers l’après-midi, activity patterns, and microhabitat use. A total of alors que vers le midi il réduit son rythme d’activité. 93 individuals were recorded, 65 being adults, 23 Il semble par conséquent que le comportement immatures, and five hatchlings. Species abundance thermorégulateur et la prédation pourraient constituer was positively correlated with the number of dead des facteurs déterminants qui influencent le mode bamboo stems. The sex ratio is largely skewed in d’utilisation du microhabitat. En effet, cette espèce a favor of males. P. klemmeri is a sexually dimorphic été fréquemment observée sur des bambous morts species with males being significantly larger de taille variée munis de fissures. Dès qu’un danger than females. In addition, this gecko showed a se présente, elle s’incruste facilement à l’intérieur bimodal pattern of activity. It would appear that des fissures grâce à son corps aplati. Elle utilise thermoregulatory behavior and predation are également les bambous verts de grande taille pour determinant factors influencing microhabitat use. effectuer différentes activités. En étalant son corps, These results confirmed the importance of bamboo ce gecko pourrait rapidement se réfugier vers le as a microhabitat to this gecko. Bamboo is widely côté opposé du tronc de bambou car ce support harvested by villagers in this region of Madagascar, est assez grand pour qu’il puisse se cacher des resulting in a decrease of habitat for this gecko. prédateurs. Tous ces points montrent l’importance des bambous comme microhabitat de ce gecko Key words: activity, bamboo, microhabitat, Phelsuma endémique de Madagascar. Dans cette région, les klemmeri, Mandrozo bambous de grande taille sont fortement récoltés par les villageois, causant une vulnérabilité importante Résumé détaillé de l’espèce étudiée. Une investigation rapide de Phelsuma klemmeri a Bien que ce site appartienne à la catégorie V été menée dans sa nouvelle localité récemment du classement IUCN, il n’y a pas de conservation reporté qui est l’Aire Protégée de Mandrozo. Cette appropriée pour cette espèce En danger. Il y a un étude a pour but d’évaluer l’état actuel de cette déclin continu de la couverture forestière dans son population afin de fournir des bases de données aire de répartition due aux activités anthropiques. nécessaires pour une étude de suivi de cette espèce En revanche, ces dernières semblent favoriser et de proposer des mesures appropriées pour sa la prolifération des bambous. La transformation conservation. Les méthodes d’observation directe continue des bordures de forêts en zone d’agriculture et de fouille systématique des bambous le long de entraine la formation des forêts de bambous. Par six transect de 10 x 100 m ont permis de mettre conséquent, P. klemmeri bénéficie de ce phénomène

Rakotozafy, L. M. S. 2019. Population structure, activity pattern, and microhabitat use of Phelsuma klemmeri at Mandrozo, Madagascar. Malagasy Nature, 13: 88-96. Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo 89 car cette plante constitue un habitat clé pour cette the IUCN classification, and is a Ramsar Site since espèce de gecko. 5 June 2012 (Razafimanjato et al., 2012). The area is managed by The Peregrine Fund and situated at Mots clés: activité, bambou, microhabitat, Phelsuma 60 km north of , in the Region. It is klemmeri, Mandrozo administratively included within the rural communes Introduction of (in the west), (in the south) and (in the east). The protected The day- of the Phelsuma are colorful, area covers 15,145 ha of which 1800 ha is freshwater attractive lizards and among the most popular habitat (17°32’S, 44°05’E, 73 m a.s.l.). This area group for terrarium keepers (McKeown, belongs to the ecoregion of western dry forest, and 1999). The majority of Phelsuma spp. are endemic to is represented by a mosaic of ecosystems: Lake Madagascar and nearby islands, including , Mandrozo, swamps, dry patches of dense forests, Mascarenes, and (Glaw & Vences, 2007; palm forests, and savannas (The Peregrine Fund, Glaw & Roesler, 2015). All Phelsuma spp. are listed 2009). The Mandrozo dry forest is highly fragmented on the CITES Appendix II. The international pet-trade and forms several different blocks. The climate is has led to the introduction of this genus into several characterized by a warm and sub-humid bioclimate countries of the world (Meshaka, 2011). (Koechlin et al., 1974) and is divided into dry season Phelsuma klemmeri, the so-called neon day from May to October and a rainy season from October gecko or yellow-headed day gecko, is a small (body to April. The annual precipitation ranges between 500 length up to 90 mm) and colorful arboreal lizard and 2200 mm with maximum of rainfall in January. endemic to Madagascar. This species is currently The average annual temperature is between 18 and assessed as Endangered by IUCN (2016) because 32° C (The Peregrine Fund, 2009). it has an extremely restricted distribution and the population is severely fragmented, known only from Bamboo description and distribution two separate and highly isolated locations, which are the Ampasindava Peninsula (northwestern In the Mandrozo protected area, Phelsuma Madagascar) and Mandrozo (Razafimahatratra et al., klemmeri uses the bamboo Dendrocalamus sp. 2008). On the Ampasindava Peninsula this species ( Poaceae, subfamily Bambusoideae), an appears threatened by forest exploitation and introduced species, as its preferred microhabitat harvesting of bamboo (Van Heygen, 2004), and from (Razafimahatratra et al., 2010). This bamboo is its newly documented site, the Mandrozo protected commonly found in areas of high humidity (Bitariho area in western dry forest, two individuals were & Mosango, 2005), along forest edges, open areas recently reported (Razafimahatratra et al., 2008). or cleared land for cultivation, as well as along Therefore, it was deemed important to assess the riverbanks. This bamboo species is characterized by current population status of the Mandrozo population straight and smooth green culms, becoming yellow in order to provide baseline information on this gecko when dry. This bamboo was characterized into two for future monitoring effort and to suggest appropriate types: measures for its conservation. ●● Green live bamboo without cracks but holes, In this study, we aimed to determine for this which contain rainwater, and form phytotelms species (i) the population structure, including for frogs. The holes are presumably made by abundance, sex ratio and, age classes, and (ii) the and rain probably aids in the development daily activity pattern and microhabitat use in order to of larvae and frog tadpoles (Duckworth, identify its ecological requirements and the level of 1993). dependency on bamboo. We also provided general ●● Yellow dead bamboo with many cracks and considerations of threats affecting this species and without water that provide refuge and shelter for indications for its conservation. P. klemmeri.

Dendrocalamus sp. is heliophilous in nature Materials and methods and therefore competes for light and soil nutrients Study site with trees in the mixed bamboo thickets (Bitariho & Mandrozo protected area is a low-altitude dry Mosango, 2005). This is probably the reason why its deciduous forest, located in western Madagascar density in the mixed bamboo thickets often decreases (Figure 1). It is a category V protected area, following from the periphery to deeper within the forest. 90 Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo Location of the study area in Mandrozo protected area. Figure 1. Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo 91

Data collection and field sampling To avoid the double counting a given individual, each gecko was painted on its dorsum when A survey was conducted from 18 October to 2 first encountered with “white non-toxic paint” and November 2015 in the vicinity of Androy (17°34’23,9” quickly released at the site of capture. To minimize S, 44°11’15,5’’E, 36 m.a.s.l.). Six transects of disturbance, marked individuals were not recaptured. 10 x 100 m each, established from the edge to the In the field, information was collected from informal middle of the bamboo thickets were made and interviews with local people, particularly information surveyed bamboos were carefully checked for the about threats, bamboo, and the study site. presence of the gecko. Each transect was searched twice during the period this day gecko is most active: Data analysis in the morning (07:30 a.m. to 11:30 a.m.) and in the afternoon (2:30 p.m. to 5:30 p.m.) by three All statistical analyses were performed using SPSS observers. Only individuals observed within a 10 m version 17.0 (SPSS Inc., 2008). All variables were band of habitat (5 m on each side of the transect) tested for normality using Shapiro-Wilk test (Shapiro were recorded. When a gecko was spotted, data on & Wilk, 1965; Royston, 1982). Non-normal variables activity, type of microhabitat where the gecko was were log10 transformed prior to analyses (Neter et initially sighted (dead or live bamboo), perch height, al., 1996). One-way ANOVA was used to compare diameter at the breast height (DBH) of the substrate, measurements taken from males and females (SVL, light condition (sun or shade) of the specific TL, BL, and BM). The diameters of bamboo and site, and their size were recorded. The following perch heights were also examined to see if these measurements from each individual were taken with factors are related to the activity patterns of this a dial calipers: snout-vent length (SVL), tail length gecko. Associations were explored between the (TL), and body length (BL) to the nearest 1 mm, and abundance of Phelsuma klemmeri and the different body mass (BM) with a balance with a precision of types of bamboo using the Spearman correlation 0.1 g. coefficient (r).

Activity Results Population structure The activity of Phelsuma klemmeri was classified into three categories: (1) moving, which includes an A total of 93 individuals were recorded and were action accompanied by a change of location such composed of 69.9% adults, 24.7% immatures, and as jumping, walking, running, retreating, and hunting 5.4% hatchlings. The sex ratio (SR = 1.5) was skewed (including stalking bamboo ), (2) basking, active in favor of males and adult males were the most thermoregulation, specifically when they were found frequently found age-sex group in the population. in the sun on exposed bamboo, and (3) resting: For the morphometric analyses, only adults for when the individual remained in the same position on which data was recorded for SVL, TL, BL, and BM a shaded bamboo-trunk or inside bamboo canes. were taken into account. According to the one-way ANOVA result, body size (SVL, TL, and BL) differed Identification of age and sex significantly between sexes, but not the body mass (Table 1). Phelsuma klemmeri is a sexually dimorphic Three age classes were defined based on total length species with males being significantly larger than of the gecko and on external sexual characters. Sex females. was determined by the occurrence of hemipenial bulges and preanofemoral pores for males and Vertical movement and microhabitat use developed extracranial endolymphatic calcium sacs Relationship between perch height and daily for females (Glaw and Vences, 2007). The sex ratio activity pattern was calculated as the proportion between males and females. Adults were those individuals with BL ≥ 70 In the early morning (8:00 a.m to 9:00 a.m), when mm, in which the external sexual characters were sunlight has not yet reached the forest floor, well-developed, immatures were those ranging in individuals were observed at the upper levels of size from 30 mm ≥ BL < 70 mm (starting at 60 mm bamboo canes, in search of basking sites (Figure in BL, males are recognizable by not well-developed 2), where they reach their optimal body temperature. external sexual characters), hatchlings ranged in size Thereafter they usually move to lower sites either to from 22 mm ≥ BL < 30 mm. expose themselves to sun-exposed bamboo canes, 92 Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo

Table 1. Morphological measurements with mean ± standard deviation of each mensuration of adult male and adult female of Phelsuma klemmeri. *= p < 0.05; N.S. = not significant withp > 0.05.

Sex SVL (mm) TL (mm) BL (mm) BM (g) Male 39.2 ± 1.31 39.10 ± 3.94 78.26 ± 4.60 1.57 ± 0.26 Female 38.13 ± 1.22 36.82 ± 2.06 74,94 ± 2.55 1.55 ± 0.15 F and P-value F(1,45) = 7.33 F(1,45) = 5.35 F(1,45) = 8.12 F(1,45) = 0.07 p = 0.010 * p = 0.025* p = 0.007* p = 0.715 N.S. SVL: Snout-vent length, TL: Tail length, BL: Body length or total length and BM: Body mass or rest on those in the shade, or within bamboo. is segregated into two different substrates: those These activities often take place between 10:00 within cracks in bamboo canes and those on bamboo a.m. and 2:30 p.m. Although the activity of Phelsuma canes. Phelsuma klemmeri showed a bimodal activity klemmeri was significantly independent of perch pattern, with peak activities in the morning between height (ANOVA: F = 1.01; df = 2; P = 0.371; Figure 2), 8:30 a.m. and 9:30 a.m. and again in the late they usually hunt bamboo flies at lower perches. afternoon between 3:00 p.m. and 4:00 p.m. There Frequency histograms of activity were plotted for was a reduction in activity during the hotter parts of different periods during the day (Figure 3). The data the day (11:00 a.m.-1:30 p.m.).

Figure 2. Perch height preference distribution of Phelsuma klemmeri according to their activity.

Figure 3. Daily activity pattern of Phelsuma klemmeri. Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo 93

Relationship between diameter and nature of Threats faced by Phelsuma klemmeri and its bamboo habitat

Of 93 individuals observed, 55 were recorded on Based on direct field observations and interviews of local people, the principal threat for Phelsuma dead and 38 on live bamboo. Spearman correlation klemmeri is mainly the loss of bamboo stands test showed that the abundance of Phelsuma resulting from direct exploitation of bamboo for klemmeri was positively correlated with the number human use. This plant is widely harvested by villagers of dead bamboo (rs = 0.22; p = 0.034). Hence, the for house and fence construction. Local people also abundance of P. klemmeri was significantly greater practice selective exploitation of large bamboo by on dead than living bamboo. cutting down their lower part, which sections between The diameter of bamboo selected by this gecko the nodes are used to collect and store liquids, such as that of the palm Bismarckia nobilis to produce an as a substrate depends significantly on the type of alcoholic drink. Selective cut of bamboo has a direct the bamboo (One-way ANOVA: F = 14.21; df = 1; impact on P. klemmeri. Further, over the last decade, p < 0.001). When they used dead bamboo, they lowland forest at the study site has been gradually selected different diameters ranging from small to cleared for rice, tobacco, and vegetable cultivation, large, whereas for living bamboo, they mainly used with a spontaneous growth of bamboo thickets in medium sized or large bamboo (≥5 cm) (Figure 4). cleared areas along forest edges.

Figure 4. Box plot of the diameter of bamboo per type of bamboo used by the population of Phelsuma klemmeri.

Discussion temperature of eggs determines the sex of the Population structure hatchlings (Ferguson & Joanen, 1982), females are produced at lower temperatures. Therefore, an The results of this study indicate that the population explanation for the skew could be that the ambient of Phelsuma klemmeri at Mandrozo shows a temperature in this region would promote the male biased sex ratio. This situation may lead to production of males. However, this is a supposition increase male-male competition associated with and further work is needed to assess this possible female sexual selection (Grant et al. 1995; Jones relationship. et al. 2004). According to Mitchel & Janzen (2010), On the basis of this study, with both the presence climate change can skew the sex ratio of temperature of hatchlings and immatures at different growth dependent sex-determination and might lead stages present between mid-October to early to local extirpation, specifically if the temperature November, female P. klemmeri probably have an change resulted in a preponderance of males (Dale, extended breeding period, including laying eggs 2001; Clout et al., 2002). In where incubation during the dry season (August-September). 94 Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo

threatened these geckos could quickly retreat and Vertical movement and microhabitat use hide at the opposite side of these bamboo. They also used small-to-large-sized dead bamboo with Possible factors that influence microhabitat use in cracks and could easily retreat into the fissures when reptiles are thermoregulatory behavior and predation threatened. Apparently, this behavior constitutes an avoidance (Huey, 1974; Gillis, 1991; Sanchez & anti-predator strategy. Parmenter, 2002; Dias & Rocha, 2004). Regarding On the Ampasindava Peninsula, P. grandis, one thermoregulatory behavior, this study indicates that of the largest Phelsuma spp. on Madagascar, co- Phelsuma klemmeri shows a bimodal activity pattern. occurs with P. klemmeri, P. seippi and P. vanheygeni Small geckos are highly susceptible to overheating in the same bamboo thickets, but no aggression was and desiccation (Stevenson, 1985; Steinberg et al., noticed towards smaller species of the genus even 2007; Allen & Powell, 2014). This species has a small when they occupy the same bamboo trunk (Van body size (up to 90 mm in BL) and seems to reduce Heygen, 2004). Nevertheless, P. kochi, another large its activity during the hottest part of the day to avoid species, can be aggressive towards P. klemmeri at overheating. Thus, using shaded bamboo stems or Mandrozo. Razafimahatratra et al. (2010) reported cracks, both sheltered from direct sunlight might be that P. kochi lived in sympatry with P. klemmeri but a tactic to avoid overheating and desiccation. Moving did not occupy the same microhabitat. During this between sun and shade (Cowles & Bogert, 1944) in study, direct predation of P. kochi upon P. klemmeri lizards is one example of behavioral thermoregulation was observed (Figure 5) and both species were often and this may be an important factor in the use of found on the same bamboo stems. bamboo by P. klemmeri. Predation avoidance is often considered as Pressure and threat the most important factor determining differential microhabitat use and activity patterns by many The main threat to endemic Malagasy reptiles is species (Lagos et al., 1995; Kronfeld-Schor & the loss of original habitat from different range Dayan, 2003). Due to its brightly colorful body and of anthropogenic activities including severe relatively small size, P. klemmeri might be subjected fragmentation by fire, illegal logging, and deforestation to important levels of predation. Throughout all their for agriculture (Ganzhorn et al., 2001; D’Cruze et daily activities, they were found on medium-sized al., 2007; Glaw & Vences, 2007; Rakotondravony, green bamboo. While exposing their bodies, when 2007). Primary forests are considered to be the

Figure 5. Phelsuma kochi preying upon P. klemmeri. (Photo by Lovasoa M.S. Rakotozafy.) Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo 95 major habitat of geckos. This is not always the case regarding Phelsuma klemmeri. On one hand, this Acknowledgements bamboo-dwelling species suffers from excessive This work was supported by the Madagascar exploitation of bamboo by villagers, which will likely Biodiversity Fund and The Peregrine Fund. Achille lead to population decline of this gecko. On the other Raselimanana provided helpful advice during hand, the species seems to benefit from increased this project. I am grateful to The Peregrine Fund, bamboo growth in human-modified habitats. During particularly Gilbert Rakotomanana for his logistic this study, it was noticed that bamboo thickets support and all the local technicians for their valuable are denser in cleared open areas or along forest assistance in the field. Particular thanks go for Steve edges. This finding corroborates Van Heygen’s Goodman, Hery Rakotondravony, Markus Roesch, (2004) observation that bamboo thickets are rapidly and Kevin Wallace for their helpful comments on this established in human disturbed areas, which in turns manuscript. constitute an advantage for P. klemmeri. Hence, P. klemmeri are also distributed and more abundant in References areas where bamboos are very dense. Allen, K. E. & Powell, R. 2014. Thermal biology However, in the Mandrozo protected area no and microhabitat use in Puerto Rican particular regulation is present with respect to the eyespot geckos (Sphaerodactylus macrolepis macrolepis). Herpetological Conservation and Biology, exploitation of bamboo, which leads to overharvesting 9(3): 590-600. by local communities, and possibly creates a Bitariho, R. & Mosango, M. 2005. Abundance, distribution, negative impact on the abundance of P. klemmeri. utilization and conservation of Sinarundinaria alpina Conservation attention is required to protect this in Bwindi and Mgahinga forest National Parks, South species in the long term. West Uganda. Ethnobotany Research and Applications, 3:191-200. Clout, M. N., Elliot, G. P. & Robertson, B. C. 2002. Effect Conclusion of supplementary feeding on the offspring sex ratio of This study provides the first baseline reference kakapo: A dilemma for the conservation of a polygynous parrot. Biological Conservation, 107:13-18. for future monitoring of an endangered gecko, Cowles, R. B., & Bogert, C. M. 1944. A preliminary study Phelsuma klemmeri, in the Mandrozo protected area. of the thermal requirements of desert reptiles. Bulletin of This study shows that P. klemmeri is dependent on the American Museum of Natural History, 83: 265-296. bamboo, which is used for shelter, hunting, basking, Dale, S. 2001. Female biased dispersal, low female resting, and nesting sites. The presence of cracks recruitment, unpaired males, and the extinction of small in dead bamboo could be one of the key factors and isolated bird populations. Oikos, 92: 344-356. determining the presence of this gecko, as based on D’Cruze, N., Sabel, J., Green, K., Dawson, J., Gardner, its compressed body is able to use this microhabitat. C., Robinson, J., Starkie, G., Vences, M. & Glaw, F. 2007. The first comprehensive survey of amphibians Hence, the presence and distribution of this bamboo and reptiles at Montagne des Français, Madagascar. appears to be related to relatively higher population Herpetological Conservation and Biology, 2: 87-99. densities of this gecko. Apart from P. klemmeri, other Dias, E. J. R. & Rocha, C. F. D. 2004. Thermal ecology, gecko species, insects and phytotelm-breeding activity patterns, and microhabitat use by two sympatric frogs depend on this plant. This demonstrates the whiptail lizards (Cnemidophorus abaetensis and Cnemidophorus ocellifer) from northeastern Brazil. importance of bamboo to Madagascar’s highly Journal of Herpetology, 38: 586-588. endemic herpetofauna for the viability of several Duckworth, J. W. 1993. Feeding damage left in bamboos, species. probably by Aye-Ayes (Daubentonia madagascariensis) An extensive survey of non-investigated areas in Marojejy National Park, Madagascar. International in Mandrozo is crucial to identify the presence of Journal of Primatology, 14: 927-931. new populations of this gecko and further details Ferguson, M. W. J. & Joanen, T. 1982. Temperature of egg on its local distribution. Genetic assessment of the incubation determines sex in Alligator mississipiensis. Nature, 296: 850-853. P. klemmeri populations from the Ampasindava Ganzhorn, J. U., Lowry II, P. P., Schatz, G. E. & Sommer, Peninsula, 600 km northwest of Mandrozo is needed S. 2001. The biodiversity of Madagascar: One of the to understand the evolutionary relationship between world’s hottest hotspots on its way out. Oryx, 35: 346- them. 348. 96 Rakotozafy: Ecology of Phelsuma klemmeri at Mandrozo

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