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Male Courtship Display in Two Populations of Anolis Heterodermus (Squamata: Dactyloidae) from the Eastern Cordillera of Colombia

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Male Courtship Display in Two Populations of Anolis Heterodermus (Squamata: Dactyloidae) from the Eastern Cordillera of Colombia Herpetology Notes, volume 12: 881-884 (2019) (published online on 15 August 2019) Male courtship display in two populations of Anolis heterodermus (Squamata: Dactyloidae) from the Eastern Cordillera of Colombia Iván Beltrán1,2,* and Leidy Alejandra Barragán-Contreras3 Animal displays are generally associated with Anolis heterodermus (Duméril, 1851) is a medium-size territoriality, predator avoidance and courtship arboreal lizard that inhabits shrubs and small trees of high behaviour, in which visual cues transmit a large amount Andean forests in Colombia and Ecuador (Moreno-Arias of information (Alcock and Rubenstein, 1989). Visual and Urbina-Cardona, 2013). Their aggressive and sexual cues can vary in type and frequency depending on behaviour have been described mainly as occasional several factors such as habitat structure, environmental observations in the field and laboratory (Jenssen, 1975; temperature and density of conspecifics (Endler, 1992; Guzmán, 1989; Beltrán, 2019). This species belongs to Candolin, 2003). Visual displays usually convey the heterodermus complex of species from which its information about species identity and/or physiological phylogenetic relations are not well established (Lazell, status of the signaller. Moreover, since an effective 1969; Castañeda and de Queiroz, 2013). Recently, it communication will determine the reproductive was suggested that there are at least three genetically success of the individual and ultimately its fitness, the distinct clades within the complex (Vargas-Ramírez and information must be quickly comprehended by the Moreno-Arias, 2014). However, there is no evidence receiver (Sullivan and Kwiatkowski, 2007). Variations that these genetic differences are backed by behavioural in the signalling pathway constitute a prezygotic changes that could act as a prezygotic barrier. Here barrier that might promote reproductive isolation and we report behavioural observations that are consistent potentially influence the speciation processes (Shuster with the existence of more than one lineage within A. and Wade, 2003; Zamudio and Sinervo, 2003). heterodermus. In lizards, courtship communication includes different We captured animals from two populations across the body movements, postures, and changes in coloration Cordillera Oriental of the Colombian Andes between (Greenberg and Crews, 1990). In lizards in the genus February-July 2016. The populations were named A Anolis, variation in sexual behaviour depends highly on and C following the classification made by Vargas- the type of habitat and density of males, with displays Ramírez and Moreno-Arias (2014). Current evidence being more frequent and with higher duration in areas suggests that there is a continuous occurrence of Anolis with dense vegetation and more males (Bloch and heterodermus between the two studied populations Irschick, 2006, Dufour et al., 2018). (Vargas-Ramírez and Moreno-Arias, 2014). However, the effect of the increasing habitat fragmentation due to human activity (i.e. urbanization and agriculture) on the species population dynamics needs to be considered in future studies (Moreno-Arias and Urbina-Cardona, 2013) (see Figure 1). 1 Department of Biological Sciences, Universidad de Los Adult males from population A (n = 2) were collected Andes. Carrera 1 No. 18A-12. A.A. 4976. Bogotá, Colombia. in Guasca (4.794 ºN, 73.908 ºW) and Bogotá D.C. 2 Department of Biological Sciences, Macquarie University, (4.675 ºN, 74.015 ºW), both populations above 2800 Sydney 2109, New South Wales, Australia. (current address) m.a.s.l and with similar vegetation structure. Adult 3 Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia. Carrera 45�26-85 males from population C (n = 3) and adult females (n Bogotá D.C. 111321. = 3) were collected in the municipality of Tabio (4.921 * Corresponding author. E-mail: [email protected] ºN, 74.070 ºW, approximately 2600 m.a.s.l) (Figure 1). 882 Iván Beltrán & Leidy Alejandra Barragán-Contreras frequency, iii) duration of dewlap extension, iv) duration of body sagittal expansion and v) total duration of the courtship display. Graphical analyses were performed in R software version 3.4.1. At the end of the experiments, all animals were released at their respective sites of capture. To reduce dimensionality and because all behavioural variables were highly correlated, we conducted a Principal Component Analysis (PCA) (Sih et al., 2004). The first two principal components explained more than 75 % of covariation in the data and separated the groups as shown in Figure 2. The first principal component (PC1) was negatively correlated with the latency to the first display and positively correlated with the Figure 1. Geographic location of the two populations of duration of dewlap extension and the total duration of Anolis heterodermus used in this study. Individuals from the courtship display. The PC2 was positively correlated population A (in red) were captured in Bogotá D.C. (4.675 ºN, with the head-bobbing frequency and negatively with the 74.015 ºW) and Guasca (4.794 ºN, 73.908 ºW). Individuals duration of body sagittal expansion. As shown in Figure from population C (in blue) were captured in Tabio (4.921 ºN, 2, males from population C performed a more elaborate 74.070 ºW). courtship. They displayed faster (low latency) and for a longer time (long courtship duration) compared to males from population A. Also, males from population Unfortunately, it was not possible to capture females C extended their dewlap for a longer time and showed from population A, which could have given us a more complete idea of the differences in courtship behaviour between populations. We acknowledge that this limitation, together with our small sample size, does not allow us to make strong conclusions. However, these observations are relevant for the ecology and behaviour of the species and we consider they will encourage further and more rigorous research. Animals were transported to the University of Los Andes (Bogotá D.C) where they were kept in individual glass terraria (60 × 50 × 60 cm), with a 12L:12D photoperiod. Environmental temperature and relative humidity varied within 20 ± 2.73 °C (mean ± SD) and 56.4 ± 13.7 %, respectively. Each terrarium was provided with small logs, leaf litter and natural wooden sticks. Lizards were fed ad libitum twice a week with crickets and mealworms. Animals were kept in these conditions for at least four weeks before the beginning of the experiments. Each observation started when a female was transferred from her original enclosure into the male’s enclosure and male courtship behaviour Figure 2. Principal Component analysis summarizing differences in the courtship behaviour of males of Anolis was video-recorded for 30 minutes. Each male was heterodermus from two populations (population C in blue: paired only once with each female, then the male had 3 males; population A in red: 2 males). Males from both at least one week of rest between each observation. We populations were paired only with females from population measured the following variables based on Guzmán C (n = 3). Each male was observed three times, one time with (1989): i) latency to the first display, ii) head-bobbing each female. Male courtship display in two populations of Anolis heterodermus 883 a higher duration of body sagittal expansion but a lower References head-bobbing frequency. Alcock, J., Rubenstein, D.R. (1989): Animal behavior. Sunderland, Our observations suggest that male courtship behaviour MA, USA: Sinauer Associates. towards females was different between populations. Beltrán, I. (2019): Diurnal colour change in a sexually dimorphic These differences between populations could be trait in the Andean lizard Anolis heterodermus (Squamata: caused by abiotic factors, such as habitat structure Dactyloidae), Journal of Natural History 53(1-2): 45–55, doi: 1 (Bloch and Irschick, 2006). Lizards from population C 0.1080/00222933.2019.1572245. Bloch, N., Irschick, D.J. (2006): An analysis of inter-population inhabit denser forests with medium-sized trees and big divergence in visual display behavior of the green anole lizard shrubs. In contrast, small shrubs are more common in (Anolis carolinensis). Ethology 112(4): 370–378. doi: 10.1111/ population A (personal observations). Although to our j.1439-0310.2006.01162. x. knowledge there are no published studies examining Candolin, U. (2003): The use of multiple cues in mate the ecomorphs of this species; most species from the choice. Biological reviews of the Cambridge Philosophical heterodermus complex are best classified as twig Society. 78(4): 575–595. Castañeda, M.R., de Queiroz, K. (2013): Phylogeny of the ecomorphs (Jonathan Losos and Rafael Moreno-Arias, Dactyloa clade of Anolis lizards: New insights from combining personal communication). Further research is needed to morphological and molecular data. Bulletin of the Museum of test if populations A and C belong to the same ecomorph Comparative Zoology 160(7): 345–398. doi: 10.3099/0027- class. 4100-160.7.345 Differences in courtship behaviour have been used Dufour, C.M.S., Herrel, A., Losos, J.B. (2018): The effect of recent to separate species in the Anolis genus. For instance, competition between the native Anolis oculatus and the invasive A. cristatellus on display behavior. PeerJ. 6:e4888. doi: 10.7717/ the A. brevirostris complex was divided into three peerj.4888 species based on differences in head bobbing displays Duméril, A.M.C., Duméril, A.H.A. (1851):
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