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Reciprocal Sexual Size Dimorphism and Rensch's Rule in Toad-Headed

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Reciprocal Sexual Size Dimorphism and Rensch's Rule in Toad-Headed SALAMANDRA 52(3) 261–268 30 October 2016 SSDISSN in a lizard0036–3375 Reciprocal sexual size dimorphism and Rensch’s rule in toad-headed lizards, Phrynocephalus vlangalii Li Zhao1, Yin Ji Chen2, Shang Ling Lou1, Yan Huang1, Robert Jehle3 & Wen Bo Liao1 1) Key Laboratory of Southwest China Wildlife Resources Conservation (Ministry of Education), China West Normal University, Nanchong, 637009, P.R. China 2) National Engineering Laboratory of Food Storage and Transportation, Nanjing University of Finance and Economics, Nanjing, 210023, P.R. China 3) School of Environment & Life Sciences, University of Salford, Salford M5 4WT, UK Corresponding author: Wen bo Liao, e-mail: [email protected] Manuscript received: 4 December 2014 Accepted: 18 May 2015 by Edgar Lehr Abstract. Rensch’s rule describes a pattern of allometry whereby sexual size dimorphism (SSD) increases with body size when males are the larger sex, and whereby SSD decreases with body size when females are larger by intraspecific compari- son. In groups of related species or sets of conspecific populations, Rensch’s rule has so far largely been confirmed with fe- males being the larger sex in small-sized populations and males the larger sex in large-sized populations. The toad-headed lizard Phrynocephalus vlangalii is a small viviparous agamid endemic to China. It exhibits either male- or female-biased SSD depending on the population, and is therefore a good model species to test Rensch’s rule relative to SSD direction. Using morphological data from 38 populations across the species’ range, we studied whether populations with male- and female-biased SSD both exhibit isometric relationships consistent with Rensch’s rule. For populations with male-biased SSD, we reject the hypothesis that SSD consistent with Rensch’s rule is driven by allometries in tail length (in this species the tail is used for visual signalling during territory defence). In populations with female-biased SSD we find significant evidence for fecundity selection favouring large females, but Rensch’s rule is not supported in this species. Our findings suggest that the underlying evolutionary forces (sexual and fecundity selection) do not promote the direction of SSD con- sistent with Rensch’s rule in both male- and female-biased SSD across populations within a species. Key words. Squamata, Agamidae, Phrynocephalus vlangalii, fecundity selection, Rensch’s rule, sexual selection, sexual di- morphism. Introduction Brock et al. 1977), and across a small range of taxa with female-biased SSD (Stuart-Fox 2009, Fairbairn 1997, Conspecific males and females differ significantly in body Teder & Tammaru 2005, Stephens & Wiens 2009). In size in many animal taxa (Andersson 1994). The phenom- contrast, Rensch’s rule is not supported in some taxa with enon, known as sexual size dimorphism (SSD), contin- female-biased SSD (Jannot & Ke rans 2003, Tubaro & ues to attract considerable research efforts (e.g., Monnet Bertelli 2003, Webb & Freckle ton 2007, Lindenfors & Cherry 2002, Matějů & Kratochvíl 2013, Liao et al. et al. 2007, Liao et al. 2013, De Lisle & Rowe 2013, Cabre- 2013, Liao et al. 2015 , Liao et al. 2016, Jin et al. 2016). In his ra et al. 2013). During recent years, intraspecific studies on seminal work, Rensch (1960) proposed that SSD increases Rensch’s rule have focused on wild (Pearson et al. 2002, with overall body size in species where males are the larg- Roitberg 2007, Herczeg et al. 2010, Liao & Chen 2012, er sex (hyper-allometry), and SSD decreases with increas- Kelly et al. 2013) and domestic animals (Polák & Frynta ing body size in species where females are larger (hypo- 2009, 2010, Remeš & Székely 2010, Frynta et al. 2012). allometry). Over recent decades, Rensch’s rule has been The variations in SSD across populations for most of these confirmed across a wide range of taxonomic groups with do not conform to Rensch’s rule (Herczeg et al. 2010, male-biased SSD (insects: Fairbairn 2005, Wolak 2008, Liao 2013). Lengkeek et al. 2008, Walker & Mccormick 2009; fish: Sexual dimorphism has attracted considerable attention Young 2005; reptiles: Iverson 1985; turtles: Cox et al. 2007, since Darwin (1871) proposed the principles of sexual se- Stuart-Fox 2009, Frýdlová & Frynta 2010, Ceballos lection (Andersson 1994). Rensch’s rule generally implies et al. 2013; birds: Dale et al. 2007; mammals: Clutton- that the evolution of SSD is mainly driven by selection, © 2016 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany All articles available online at http://www.salamandra-journal.com 261 Li Zhao et al. favouring large males as a consequence of sexual selec- Materials and methods tion stemming from a large-male advantage in male–male Study species competition (Berry & Shine 1980, Fairbairn & Prezio- si 1994, Abouheif & Fairbairn 1997, Smith & Cheve- The toad-headed lizard P. vlangalii is a small viviparous rud 2002, Cox et al. 2003, Székely et al. 2004, Fairbairn agamid endemic to China and typically found in arid or 2005, Dale et al. 2007, Blanckenhorn et al. 2007, Lisle- semiarid regions. Its distributional range covers Qinghai, vand et al. 2009, Serrano-Meneses et al. 2009). For taxa Gansu, Xinjiang, and northwestern Sichuan, at altitudes with male-biased SSD, Rensch’s rule results from the ac- ranging from 2,000 to 4,500 m above sea level (Zhao tion of sexual selection where intense sexual selection & Adler 1993). The species has an activity season that fuels the evolution of larger body size in males, accom- stretches from May through October. Female reproduc- panied by a weaker correlated selection that favours in- tion and sexual dimorphism as well as altitudinal varia- creased body size in females (Fairbairn 1997, Abouheif tion of morphological characters of P. vlangalii have been & Fairbairn 1997). In species with female-biased SSD, described previously (Huang & Liu 2002, Wu et al. 2005, SSD might evolve through different selection pressures Zhang et al. 2005, Jin & Liu 2007, Jin et al. 2006, Qi et on males and females. As proximate causes, sexual selec- al. 2011). We collected data on average reproductive invest- tion may favour smaller male size to increase mobility or ments of females in 15 populations (i.e., from 6 male- and agility (Fairbairn 1997, Székely et al. 2004, Stuart-Fox 9 female-biased populations). During courtship, males use 2009). In contrast, fecundity selection favours larger fe- tail-curling to attract potential mates, with tail length be- males with increased reproductive potential, and thus has ing positively correlated to the frequency of tail-curling as been used to explain SSD patterns consistent with the in- well as territory size (Zhang et al. 2005, Qi et al. 2011). verse of Rensch’s rule (Cox et al. 2003, Liao et al. 2013, Lu Females will take note of the length of males’ tails, the de- et al. 2014, Jin et al. 2015). gree of curling, and the frequency of swinging, and then Previous studies on reptiles have confirmed the gener- approach a male that advertises himself with a longer tail, al pattern of SSD following Rensch’s rule applying to taxa intense curling and swinging (Qi et al. 2011). As a result, with male-biased SSD (Cox et al. 2007, Frýdlová & Fryn- males with longer tails, intense curling and swinging are ta 2010, Cabrera et al. 2013). In Phrynocephalus lizards, more likely to mate with females, confirming that a male’s larger male individuals have relatively longer tails so that tail length plays a more important role than body size and these individuals have an advantage of improved visibil- resulting in greater variation in SSD. ity when engaging in the important behaviour of tail curl- ing and thus stand a better chance of attracting mates. Hence, sexual selection favours an increase in male body Data collection size, resulting in an SSD that follows Rensch’s rule (Cox et al. 2007). In the present paper, we describe the patterns For the present work, we combined published information of SSD in the toad-headed lizard Phrynocephalus vlanga­ (data taken from Zhang et al. 2005, Jin et al. 2006, Jin & lii, a species characterized by an inconsistent SSD (male or Liu 2007, Qi et al. 2011) with our own data based on the female-biased) depending on the population. The aim of measurements of 362 museum specimens at the Zoologi- this study was to investigate patterns and possible causes cal Museum in Chengdu Institute of Biology, Chinese Aca- of variation in reciprocal SSD found within a single species demic of Science, collected between the 1950s and 2000s (P. vlangalii) and in particular, conduct an intraspecific test during the breeding season between July and August (Sup- of Rensch’s rule in the species. To this end, we analysed a plementary table 1). All museum specimens were stored large dataset of male and female body and tail sizes from in 10% neutral buffered formalin, which did not result in 38 populations collected in the Hengduan Mountains, Chi- shrinkage based on the fact that there is no difference in na. In addition to testing for Rensch’s rule, we also tested body and tail sizes between data recorded at the time of hypotheses regarding whether sexual selection favouring sampling and the same individuals in preservative (Huang increased body size in males, which was tied to longer tail et al. 2014). Snout–vent length (SVL) and tail length (TL) length, led to variation in SSD consistent with Rensch’s rule were measured to the nearest 0.1 mm, and SVL was used across male-biased populations. Moreover, we investigated as the body size variable to investigate Rensch’s rule. Indi- whether fecundity selection favouring increased reproduc- viduals larger than 48 mm in SVL were considered adults, tive investment in larger female size resulted in variation because these males and females have the potential to re- in SSD consistent with the inverse of Rensch’s rule across produce (Jin et al.
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