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Sexual Dimorphism in Trapelus Ruderatus Ruderatus (Sauria: Agamidae) with Notes on the Natural History

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Sexual Dimorphism in Trapelus Ruderatus Ruderatus (Sauria: Agamidae) with Notes on the Natural History Copyright: © 2011 Fathinia and Rastegar-Poutani. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any Amphibian & Reptile Conservation 5(1):15-22. medium, provided the original author and source are credited. Sexual dimorphism in Trapelus ruderatus ruderatus (Sauria: Agamidae) with notes on the natural history BEHZAD FATHINIA1 AND NASRULLAH RASTEGAR-POUYANI1, 2 1Department of Biology Faculty of Science, Razi University, 67149 Kermanshah, IRAN Abstract.—We studied sexual dimorphism and some aspects of natural history and behavior of the Persian agama (Trapelus ruderatus ruderatus) from a population in Dehloran Township, Ilam Province, southwestern Iran. Findings were obtained by personal observations and using SPSS 13 statistical package. Based on the analyses, some characters show differences between males and females. All findings forT. ruderatus in this paper are reported for the first time. Key words. Agamidae, Trapelus ruderatus, statistical analyses, Ilam, Iran, dichromatism, sexual selection, natural selection Citation: Fathinia, B. and Rastegar-Pouyani, N. 2011. Sexual dimorphism in Trapelus ruderatus ruderatus (Sauria: Agamidae) with notes on the natural history. Amphib. Reptile Conserv. 5(1):15-22(e22). Introduction velopment of sexual dimorphism in various animal taxa. However, most of these mechanisms can be summarized The genus Trapelus Cuvier, 1816, comprises four species by three major forces differentially acting on males and on the Iranian Plateau as follows: T. agilis (Olivier 1804), females of a population: sexual selection, fecundity, and T. lessonae (De Filippi 1865), T. ruderatus (Blanford natural selection (Olsson et al. 2002; Cox et al. 2003). 1881) (sensu Rastegar-Pouyani 2000) and T. megalonyx In many taxa, competition between males over resources (Günther 1865). The distribution of T. ruderatus in Iran characteristically produces an asymmetry in body size is limited to southern and southwestern regions of the between the sexes. Thus, the advantages of larger size for Iranian Plateau (Anderson 1999; Rastegar-Pouyani 2000; males typically results in sexual size dimorphism (SSD) Fathinia 2007; Rastegar-Pouyani et al. 2008). Among the (Terry et al. 2001). Sexual selection acts on competi- Iranian species of the genus Trapelus the study of sexual tion between males, often resulting in larger body size dimorphism has already been carried out in Trapelus and in larger sizes of morphological structures related to agilis (Rastegar-Pouyani 2005). In this relation, study fight (Darwin 1874; Verrastro 2004). Anderson and Vitt of sexual dimorphism, coloration and color pattern, and (1990) suggest that the causes of sexual dimorphism in natural history of the Persian agama (Trapelus ruderatus) size could be related to several factors: competition be- is of interest and importance. tween males; differential mortality between sexes due to As genetic correlation between the sexes is very differences in longevity; larger amount of energy allo- high for most morphological traits, it is often believed cated by females for reproduction; males are more active that long periods of time are required to overcome ge- because they need to search for females and thus present netic constraints and to evolve sexually dimorphic mor- a larger predation risk. phological traits (e.g., Lande 1980; Hedrick and Temeles In this paper, the patterns of sexual dimorphism in 1989; Kratochvíl et al. 2003). Moreover, the evolution of the Persian agama, T. ruderatus, in relation to environ- sexual dimorphism may be limited by physiological and mental issues are discussed. ecological constraints as well (Kratochvíl et al. 2003). In agamid lizards, both sexual selection and natural selection influence the form of dimorphism in secondary Materials and methods sexual traits (Stuart-Fox and Ord 2004). Sexual dimor- phism (SD) in body shape as well as overall body size This survey was carried out in Dehloran area at an eleva- is a widespread and common trait among animals (Ji et tion of 202 m, approximately 5 km around the city of De- al. 2006; Kaliontzopoulou et al. 2007), most species be- hloran, Ilam Province. The coordinates of study site are ing dimorphic rather than monomorphic (Schoener 1977; 33.5°39΄N, and 45°18΄E. The information was accessed Mouton and van Wyk 1993; Andersson 1994). Different by a GPS model Etrix. The study area has an annual pre- evolutionary mechanisms have been proposed for the de- cipitation of 244.2 mm, and an annual average maximum Correspondence. 2Email: [email protected] Amphib. Reptile Conserv. | amphibian-reptile-conservation.org 015 September 2011 | Volume 5 | Number 1 | e22 Fathenia and Rastegar-Pouyani Figure 1 (left). Dorsal view of a male (right) and a female (left) of Trapelus r. ruderatus. Figure 2 (above). Ordination of the individual males and females of Trapelus r. ruderatus on the first two princi- ple components. Note the relative degree of isolation of males and females. Figure 3. The color pattern of an adult male T. ruderatus during the hottest hours of the day. Amphib. Reptile Conserv. | amphibian-reptile-conservation.org 016 September 2011 | Volume 5 | Number 1 | e22 Sexual dimorphism in Trapelus ruderatus ruderatus Figure 4. An adult male T. r. ruderatus capturing a spider while foraging. Figure 5. The occurrence of T. r. ruderatus with Uromastyx loricatus in the same hole. Amphib. Reptile Conserv. | amphibian-reptile-conservation.org 017 September 2011 | Volume 5 | Number 1 | e22 Table The comparison 1. of 16 characters in males and females of Trapelus. SEM: standard r. ruderatus error of mean; D. of d.: Direction of difference; M. of d.: Mean of differences. All measurements in millimeters (mm). SEX IL IN NP SDL SVL TL HL RP SL SBEH LFL LHL LFH VL HW CT ♂ Mean 16.84 5.76 8.48 21.48 85.95127.25 21.66 1.16 18.08 15.4831.6844.9142.648.75 19.10 15.92 N 25 SEM 0.27 0.240.780.362.754.400.660.11 0.24 0.481.351.191.8 0.37 0.610.282 ♀ Mean 16.06 5.73 2.86 21.33 76.05106.819.530.40 16.40 15.2027.3239.5937.576.54 17.54 13.86 N 15 SEM 0.37 0.360.760.422.822.380.660.1393 0.45 0.350.870.971.720.3 0.6 0.29 Fathenia andRastegar-Pouyani M. of d. 0.78 0.035.620.159.9 18.85 2.13 0.761.680.284.365.325.072.211.562.06 D. of d. M>F 018 farian 2008). of Ilam”(Mozaf based on“Flora were determined cies Plant spe as “exposed.” body regionswereconsidered whereas theremaining were regardedas“concealed,” regions gions ofthehead,throat,chestandventral re lateral According toStuart-FoxandOrd(2004)the carefully. were considered type andvegetation habitat and of concealedandexposedbodyregions,behavior, changes and color color pattern species including the of study ofsexualdimorphism,someaspectstheecology the to In addition analyses. statistical out the carrying for The SPSS software version13wasused were employed. matrix) (PCA: correlation Analysis Component Principle of sexual dimorphism, the test ANOVA as well as the Gulf and/or stereomicroscope. To test the significance model Shoka caliper the nearest0.01mmusingdigital to in mm measured were characters and meristic metric The scales. preanal rows ofcallose RP: scales; callose toe; IN: number of internasals; NP: number of preanal the fourth under lamellae SDL: subdigital across head; eyes between scales SBEH: scales; supralabial of number SL: scales; number of infralabial IL: dorsal sideoftail; VL: ventlength. hind limb,fromaxiltogroin; and limb fore between LFH: length hind limb; of length LHL: of forelimb; length LFL: head width; maximum border ofearopening;HW: end ofrostraltoanterior length, from HL,head to tipof tail; from cloaca length, TL: tail fromendofmentaltocloaca; snout-vent length, and (2007). Torki The metric characters included: SVL: (1999,2005) istic characters,basedonRastegar-Pouyani and mer 16metric Measurements included study area. the in and released measured, were caught, specimens The mm 17 and remaining 92.72 mm SVL, respectively. were 111.76 and female male The largest further studies. sex andthenpreservedfor were dissectedtodetermine (12 malesand 8females) Many ofthespecimens area. study 2009 inthe 2008 toSeptember during September specimens werecollected shah) whilethe37remaining Kerman Museum, Zoological University RUZM (Razi three wereborrowedfrom Ofthese, in thissurvey. amined mens of totalof40adultspeci A fan area. semi-desert, alluvial a which is and Ord (2004) Stuart-Fox based on habitat tively (Abdali 2009). The study area classified as open 17.8°C, respec 31.6°C and of temperature minimum and the males have greater values than females. Based onthis values thanfemales. the maleshave greater characters significant the all For CT. and VL, LHL, LFL, SL, TL, HL, RP, SVL, and femalesfor 10 characters NP, males between There areobviousdifferences 1. Table is shownin characters summaryofthe16measured A Statistical analysis Results The meristic characters included: CT: crossbars on CT: included: characters meristic The T. r. ruderatus r. T. (25 males, 15 females) were ex 15females) (25males, - - - - - - - - Amphib. Reptile Conserv. | amphibian-reptile-conservation.org September 2011 | Volume 5 | Number 1 | e22 Sexual dimorphism in Trapelus ruderatus ruderatus Table 2. Extraction of principle components 1-4 using IL and CT, compared with individuals at the other end the component matrix. with relatively small IN and SBEH and high values for IL and CT. Characters PC1 PC 2 PC 3 PC 4 The fourth axis (PC4) contains only 6.94% of the IL -0.239 -0.575 -0.355 -0.239 IN -0.402 -0.450 -0.586 -0.032 total difference, highlighting a meristic axis showing in- NP -0.715 -0.370 -0.175 -0.322 dividuals at one end with relatively large SDL and small SDL -0.086 -0.410 -0.180 -0.666 CT, SBEH, and NP, compared with individuals at the SVL -0.960 -0.206 -0.096 -0.022 other end with small SDL and large CT, SBEH, and NP.
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