The Female Reproductive Cycle of the Lizard, Cordylus Polyzonus Polyzonus (Sauria: Cordylidae) in the Orange Free State
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S. Afr. J. Zoo!. 1989,24(4) 263 The female reproductive cycle of the lizard, Cordylus polyzonus polyzonus (Sauria: Cordylidae) in the Orange Free State J.H. van Wyk* National Museum, P.O. Box 266, Bloemfontein, 9300 Republic of South Africa Received 13 November 1987; accepted 24 February 1989 Winter reproductive activity is exhibited by females of the vIviparous lizard, Cordy/us p.po/yzonus. Vitellogenesis commences in May (autumn), continuing throughout the winter months with ovulation in October. Females are pregnant during summer and give birth in late summer (February). Clutch size is positively correlated to female body size. During embryonic development, ovarian follicles remain small and translucent and the corpora lutea progressively decrease in size until parturition occurs. Embryonic growth was characterized by a concomitant decline in embryonic yolk mass. Although considerable variation was observed in the fat body mass, maximal fat body mass peaked in winter (June) and in mid-summer (December). The onset of vitellogenesis corresponded to declining photoperiod and environmental temperatures. Die lewendbarende akkedis, Cordy/us p. po/yzonus toon voortplantingsaktiwiteit gedurende die wintermaande. Dooierneerlegging begin gedurende Mei (herts) en duur voort deur die wintermaande tot ovulasie in Oktober. Wyfies is verwagtend gedurende die somer en kleintjies word in die laat-somer (Februarie) gebore. Eiergetal is positief gekorreleerd met die liggaamsgrootte van die wyfie. Gedurende embrionale ontwikkeling bly die ovariumfollikels klein en deursigtig en die corpora lutea verklein voor geboorte in Februarie. Embrionale groei gaan gepaard met 'n afname in die embrionale dooiermassa. Alhoewel aansienlike variasie in die massa van die vetliggame voorkom, is maksimum vetliggaam-massa gedurende winter- (Junie) en somermaande (Desember) waargeneem. Die aanvang van dooierneerlegging val saam met die verkorting van dagliglengte en dalende omgewingstemperature. 'Present address: Department of Zoology, University of Stellenbosch, Stellenbosch, 7600 Republic of South Africa . ) 0 Studies examining the reproductive strategies of South It is widely accepted that separate phylogenies of vivipa 1 0 African reptiles are uncommon. An examination of the rity must have occurred (Guillette 1981; Shine 1985). 2 d reproductive cycle and the possible proximal factors Shine(1985) suggested that viviparity evolved only once e t influencing or controlling this cycle forms the basis of the in the family Cordylidae, however, the specific evolu a d ( analysis of the reproductive strategy of a species tionary pathway remains to be determined for this r e (Guillette & Casas-Andreu 1987). Information on the group. h s reproductive cycles of South African reptiles is anecdotal Cordylus polyzonus is a rupicolous lizard known from i l b in nature, consisting primarily of observational data the southern parts of the Orange Free State, westwards u P rather than detailed seasonal studies. throughout the karroo to western Cape Province and e h Most studies have examined the reproductive biology southern South West Africa (FitzSimons 1943; Love t y of North American or European lizard species (see Fitch ridge 1944; De Waal 1978). This species is viviparous, b 1970; Duvall, Guillette & Jones 1982; Licht 1984). These but as for all the other cordylid species, little is known d e t studies have demonstrated that variation in reproductive regarding the reproductive cycle of C. p. polyzonus. n a cyclicity varies among geographically isolated popUla Hewitt & Power (1913) suggested that courtship takes r g tions of the same species (Guillette & Casas-Andreu place in September with young born in January. e c 1987). Moreover, field data and laboratory experiments Loveridge (1944) observed fully developed in utero n e c have demonstrated that several environmental factors young in a female collected in April. Fitch (1970) i l influence reproductive cyclicity in reptiles (Duvall et al. reported mating in early spring and birth, usually two r e d 1982; Licht 1984.) As current life history theory is based young, in mid- or late summer. De Waal (1978) found n u on data derived primarily from temperate northern 2-3 well-developed embryos in specimens collected in y hemisphere reptiles, additional studies are needed January and February. a w examining species inhabiting the tropics and southern The purpose of this study is to describe the reproduc e t a hemisphere. tive cycle of C. p. polyzonus in the Orange Free State G The family Cordylidae is endemic to Southern Africa using material housed in the herpetology collection of t e and Madagascar. All the species included in the six the National Museum, Bloemfontein. Previous studies n i b genera of the subfamily Gerrhosaurinae are known to be have demonstrated that data obtained from museum col a S oviparous. Except for the genus Platysaurus, the species lections represent an important source of information y b of the three remaining genera (Cordylus, Pseudocordylus concerning the reproductive biology of reptiles (see d and Chamaesaura) of the subfamily Cordylinae are Shine 1980,1982; Guillette & Casas-Andreu 1987). e c u known to be viviparous. (FitzSimons 1943; Fitch 1970). Because C. p. polyzonus is widespread in its distribution, d o r p e R 264 S.-Afr. Tydskr. Dierk. 1989,24(4) this study is also the first of a series of investigations of the Orange Free State region (De Waal 1978) and throughout the distribution range to eventually analyse housed in the collection of the National Museum, geographic variation in reproductive timing in order to Bloemfontein. Additional collecting in the study area understand the role of biogeographic history in shaping was undertaken during 1983 to obtain larger sample the reproductive strategy of this species. sizes. Since the smallest female exhibiting vitellogenesis (therefore sexually mature) had a snout to vent length Materials and Methods (SVL) of 87 mm, only lizards larger than 90 mm SVL Study area (101± 0,54 mm) (n = 165) were dissected. The maximum SVL size reported for this species is 124 mm The specimens examined were collected in the southern (FitzSimons 1943; Loveridge 1944; De Waal 1978). Fat region of the Orange Free State (O.F.S.) within the bodies, embryo mass, yolk mass and stomach contents boundaries of 28°-31°S latitude and 24°30'-27°E were dried to constant mass at 60°C in an oven. The longitude. Vegetation predominantly represents karroo following pertinent measurements were taken during or karroid-veld (Acocks 1975). Dolerite outcrops are a autopsy: SVL (nearest 0,1 mm), diameter of the three common phenomenon in the study area and De Waal largest ovarian follicles (nearest 0,01 mm), diameter of (1978) suggested that C. p. polyzonus may occur all corpora lutea (nearest 0,01 mm) and longest and exclusively on dolerites in the study area. The study area shortest diameter of in utero eggs (nearest 0,01 mm). is a semi-arid region with an anrttial rainfall of 400 mm or Measurements were made either by vernier caliper or an less. This region is characterized by summer rains with a ocular micrometer. Additionally, clutch size was distinct dry season during the winter months (Figure 1). obtained for all pregnant females and embryonic Most of the study area falls in the moderately severe development was staged using the embryonic sequence frost zone. Weather records were obtained for several defined by Defaure & Hubert (1961) for the viviparous weather stations in the region from the South African lizard Lacerta vivipara. Weather Bureau. A principal component analysis (PCA) All similar data for a female were averaged and all was conducted on the climatic variables whereafter means for a given month were combined to calculate another PCA was conducted on the various weather monthly means and standard errors. Analysis of variance stations using the environmental variable with the (ANOVA) was performed to determine significant highest PCA score. The Fauresmith weather station changes in monthly samples. Clutch size was regressed (29°46'S /25°19'E) explained 99,8% of the climatic against SVL and linear and exponential models fitted. variation in the study area and was therefore used as the All means are presented as mean ± lSE (standard . ) error). A Student's t test was used for comparisons of 0 reference weather station. Using daily records, maxi 1 two groups (BIOSTAT: Pimentel & Smith 1968). 0 mum and minimum air temperatures and total rainfall 2 were calculated on a monthly basis for the period 1972- Probability values smaller than 0,05 were recognized as d e t 1976 (Figure 1). Photoperiod regimes were calculated significant. a d using the formula in Van Leeuwen (1981). ( r Results e h Reproductive cycle s Reproductive cycle i l b The majority of the C. p. polyzonus specimens studied The reproductive cycle of female C. p. polyzonus is u P were collected during an extensive survey of the reptiles presented in Figure 2. Females examined during late e February and early March contained small translucent h t ovarian follicles (1,78 ± 0,06 mm). Variability was y b RAlNFALL(mm) 80 evident in follicular size during vitellogenesis, indicative d e t of asynchrony among females during this period of n 30 70 follicular growth. However, a significant seasonal trend a r g 80 was observed (F ,151 48,96; P < 0,0001). Vitello l1 = e c genesis began in May (autumn) with 50% of the sample n 20 50 e containing follicles larger than 3 mm. During the winter c i l 40 15 months, June and July, 10% contained follicles larger r e than 3 mm and 36% had follicles larger than 5 mm. The d 10 30 n onset of vitellogenesis and the early part of follicular u 5 20 y growth is negatively correlated with decreasing ambient a temperatures (r -0,9203; P < 0,05), rainfall (r w o 10 = = e t -0,9012; P < 0,05) (Figure 1) and photoperiod (r = a ~ 0 G -0,8473; P < 0,05) (Figure 3) in the study area.