Sexual Dimorphism and Natural History of the Western Mexico Whiptail, Aspidoscelis Costata (Squamata: Teiidae), from Isla Isabel, Nayarit, Mexico
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NORTH-WESTERN JOURNAL OF ZOOLOGY 10 (2): 374-381 ©NwjZ, Oradea, Romania, 2014 Article No.: 141506 http://biozoojournals.ro/nwjz/index.html Sexual dimorphism and natural history of the Western Mexico Whiptail, Aspidoscelis costata (Squamata: Teiidae), from Isla Isabel, Nayarit, Mexico Raciel CRUZ-ELIZALDE1, Aurelio RAMÍREZ-BAUTISTA1, *, Uriel HERNÁNDEZ-SALINAS1,2, Cynthia SOSA-VARGAS3, Jerry D. JOHNSON4 and Vicente MATA-SILVA4 1. Centro de Investigaciones Biológicas (CIB), Universidad Autónoma del Estado de Hidalgo, Carretera Pachuca-Tulancingo, Km 4.5 s/n, Colonia Carboneras, Mineral de La Reforma, A.P. 1-69 Plaza Juárez, C.P. 42001, Hidalgo, México. 2. Instituto Politécnico Nacional, CIIDIR Unidad Durango, Sigma 119, Fraccionamiento 20 de Noviembre II, Durango, Durango 34220, México. 3. Laboratorio de Herpetología, Escuela de Biología, Benemérita Universidad Autónoma de Puebla, C. U. Boulevard Valsequillo y Av. San Claudio, Edif. 76, CP. 72570, Puebla, Puebla, México. 4. Department of Biological Sciences, University of Texas at El Paso, 500 West University Avenue, El Paso, TX 79968, USA. *Corresponding author, A. Ramírez-Bautista, E-mail: [email protected] Received: 25. April 2014 / Accepted: 13. June 2014 / Available online: 16. October 2014 / Printed: December 2014 Abstract. Lizard populations found in insular environments may show ecological and morphological characteristics that differ from those living in continents, as a result of different ecological and evolutionary processes. In this study, we analyzed sexual dimorphism, reproduction, and diet in a population of the whiptail lizard, Aspidoscelis costata, from Isla Isabel, Nayarit, Mexico, sampled in 1977 and 1981. Males and females from Isla Isabel showed no sexual dimorphism in many morphological structures, such as snout-vent length (SVL), but they did in femur length (FL) and tibia length (TL). Both sexes displayed synchronous reproductive cycles. However, males had smaller SVL (47.0 mm) in average than females (59.0 mm) at sexual maturity. The average clutch size was 2.4 eggs, but there was no correlation between egg number and female’s SVL. The diet of A. costata consisted of insects, arachnids, crustaceans, and plant material. Insects were the most consumed prey item, being mostly represented by orthopterans, coleopterans, and hymenopterans. Diet breadth of males was higher (Levin´s formula, B = 0.596) than in females (B = 0.358), and diet overlap between sexes was relatively high (Pianka’s index, Ojk = 0.822), with an overlap ranging from 66.2 to 100%. The population of A. costata from Isla Isabel shows different morphological and ecological characteristics from their continental counterparts, but future studies on whiptail species with island and mainland populations are required to determine the ecological and evolutionary patterns. Key words: Aspidoscelis costata, clutch size, diet, Isla Isabel, Mexico, Teiidae. Introduction phism (SSD), in the form of anatomical differences between the sexes, has evolved due to an advan- Several ecological and evolutionary studies in- tage for larger males being selected by females, volving reproduction, diet, and sexual dimor- because large body size increases survival prob- phism conducted on island vertebrates (mammals, ability, which can be passed on to offspring birds, and reptiles) have generated great interest (Anderson & Vitt 1990, Butler et al. 2000), and 2) in the scientific community (Boback 2003, Lo- Niche divergence, which assumes that such sexual molino 2005, Meiri 2008), particularly with regard differences have evolved due to intraspecific niche to adaptive processes of life on islands (Channell divergence in nutritional requirements, or selec- & Lomolino 2000, Lomolino et al. 2006). Evolu- tion for different habitats (Stamps 1983, Shine tionary processes including morphological (e.g., 1989, Hierlihy et al. 2013). sexual dimorphism) and ecological characteristics Several studies have tested some life-history (e.g., differentiation in diet, clutch size, and snout– characteristics of populations found in insular en- vent length [SVL] at birth) are well documented in vironments. For example, Andrews (1979) found vertebrates inhabiting islands (Lomolino et al. high sexual dimorphism in lizard populations of 2006). Sexual dimorphism and diet preference are the genus Anolis, with sexual dimorphism biased central components for understanding life-history to males and mainly caused by competition for evolution of lizard species inhabiting islands (An- food resources and territory. On the other hand, drews 1979, Meiri 2008). Hasegawa (1994, 1997) found that some lizards in- With respect to lizards, sexual dimorphism habiting islands had larger males due to diet type, can be explained by two hypotheses: 1) Sexual se- as larger body and head sizes were required to fa- lection, which assumes that sexual size dimor- cilitate, with the aid of bacteria, the consumption Sexual dimorphism and natural history of the whiptail lizard 375 and digestion of cellulose associated with their hindlimbs), forearm length (FL: ± 0.1 mm; length from the vegetarian diets. Herbivory in island lizards has elbow to the pad of the foot), head width (HW: ± 0.1 mm; been associated with high availability of fruits maximum width of the head, measured as the distance between the posterior margin of the left and right ears), (Olensen & Valido 2003). Therefore, island lizards head height (HH: ± 0.1 mm; length of the earpiece open- may evolve sexual dimorphism in morphological ing height), head length (HL: ± 0.1 mm; length from tip of characters, while developing dietary differences snout to the posterior margin of the tympanic mem- between sexes, as reported for some birds (Grant brane), femur length (FEL: ± 0.1 mm; distance from the 1965). angle of the groin to the knee), and tibia length (TL: ± 0.1 The genus Aspidoscelis is one of the most di- mm; length from the knee to the pad of the foot; Olsson et verse genera in North America, including Mexico, al. 2002, Aguilar-Moreno et al. 2010). These morphologi- cal characters were measured to the nearest 0.1 mm. Be- where 43 species are currently recognized (Reeder cause these variables usually vary with SVL, we first cal- et al. 2002, Wilson et al. 2013). Species in this ge- culated regressions of log10-transformed data for all vari- nus are found in many environments, both in is- ables with log10-transformed SVL (Schulte-Hostedde et al. lands and on the mainland. However, few studies 2005). For significant regressions, we calculated residuals have evaluated the different biological aspects of from the relationship of variables on SVL to produce SVL- populations living in continental or oceanic is- adjusted variables. We used these residuals to examine lands. Currently, little information is available on sexual size differences between males and females and performed a Mann-Whitney U-test (Ramírez-Bautista & the natural history of Aspidoscelis costata (Teiidae), Pavón 2009, Ramírez-Bautista et al. 2013). The normality a species known from both the mainland and ad- of the data was verified by the Kolmogorov-Smirnov test jacent islands on the Pacific. Herein, we evaluate and Lillifors probability. We calculated a Pearson’s prod- sexual dimorphism in body size, aspects of diet, uct-moment correlations coefficient to test for relationship and reproduction of a population from Isla Isabel, between the morphological characters and the prey size Nayarit, Mexico. of males and females. The data are presented as mean ± SE, unless otherwise indicated. Reproductive characteristics. The following linear Material and methods measurements were taken to the nearest 0.1 mm on ne- cropsied lizards: SVL, length and width of right testis in Study area adult males; length and width of right and left nonvitel- This study was conducted in Isla Isabel (21° 52´ 30´´ N, logenic follicles (NVF), vitellogenic follicles (VF), and the 105° 54´54´´ W; datum: WGS84). The dominant vegetation number of ovulated eggs in adult females (Ramírez- type on the island is tropical dry forest, and the climate Bautista et al. 2000, Ramírez-Bautista & Pardo-De la Rosa has been described as tropical sub-humid with summer 2002). Additionally, the number of NVF and VF in ovaries rains (García 1973). The minimum ambient temperature and oviductal eggs in each oviduct was also recorded. has been reported as 22.6°C in January and a maximum of The length and width of gonads were used to obtain 30.3°C in September (CONANP 2005). testicular and follicular volume (V), calculated with the formula for the volume of an ellipsoid: V = 4/3 πa2b, Sampling where a is one-half the shortest diameter and b is half the A total of 121 lizards (78 males, 43 females) were captured longest diameter (Selby 1965). Testicular and follicular in April 1977 (n = 43) and May 1981 (n = 78). Both sam- volumes were used as indicators of reproductive activity ples were acquired during the spring, when the reproduc- in male and female lizards, respectively (Ramírez- tive activity of males (sperm production) and females Bautista et al. 2000, Ramírez-Bautista & Pardo-De la Rosa (vitellogenic follicles and eggs production) were at their 2002). The difference in testicular and follicular volume maximum. All specimens were euthanized in the labora- between the April and May collections was tested with a tory by lowering body temperature and fixed with a 10% Mann-Whitney U-test (z). The smallest adult female (SVL formalin solution using the methods described by = 59.0 mm) containing enlarged vitellogenic follicles or Ramírez-Bautista et al. (2000), and deposited in the herpe- oviductal eggs was used to estimate the minimum SVL at tological collection of the Centro de Investigaciones sexual maturity (Vitt & Cooper 1985, Ramírez-Bautista et Biológicas (CIB), Universidad Autónoma del Estado de al. 2000, Ramírez-Bautista & Pardo-De la Rosa 2002). The Hidalgo. minimum size of adult males at sexual maturity (SVL = 47.4 mm) was based on the size of the smallest male con- Data analysis taining enlarged and highly convoluted epididymides Sexual dimorphism. All data on morphological descrip- (Lozano et al.