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From Southwestern Western Australia Records of the Western Allstraliall Mllselilll 19: 57~63 (1998). Reproduction and diet in four species of burrowing snakes (Simoselaps spp.) from southwestern Western Australia l 2 N. R. Strahan ,3, R. A. How and J. DeW 1 School of Environmental Biology, Curtin University, GPO Box U 1987, Perth, Western Australia 6845, Australia 2 Museum of Natural Science, Francis Street, Perth, Western Australia 6000, Australia .1 Present address: 3/61 Anstey Street, South Perth, Western Australia 6151, Australia Abstract - The reproductive pattern and diet of four 5imoselaps species were examined from 259 museum specimens collected from southwestern Western Australia. Few reproductively active females were recorded from the four species (5. ber/holdi, 5. bimaclila/lls, 5. calon%s and 5. fasciola/us) but yolked ovarian follicles and oviductal eggs were observed between October and February with a peak in November and December. Male 5. ber/holdi had significantly larger testes in October than other months, but no significant seasonal variation was apparent in the small adult sample of other species. The prey of all species consists of scincid and pygopodid lizards. There is a pronounced separation in the diet of 5. ber/holdi and 5. calon%s with the former feeding on surface dwelling skinks and the latter on fossorial skinks and pygopodids. INTRODUCTION bertholdi (Jan), 5. bimaclIlatus (Dumeril, Bibron and The knowledge of the biology of Australian Dumeril), 5. calonotos (Dumeril, Bibron and snakes has expanded rapidly over the past two Dumeril) and 5. fasciolatus (Gunther)] that are decades (Shine 1991). A considerable amount of broadly sympatric in southwestern Australia. new information has resuIted from detailed Together with 5. semifasciatlls (Gunther) these four examination of museum specimens for both their species may occur syntopically in several sandplain reproductive and dietary information (Shine 1983, habitats on the Swan Coastal Plain (How and Dell 1984a, 1984b, 1986, 1988). 1990). Southwestern Australia has a pronounced Previous studies of burrowing snakes in the Mediterranean climate with cool wet winters and genera Neelaps, Simoselaps and Vermicella document dry hot summers; a climate that invokes a period the reproductive pattern and season, size at of pronounced reproductive activity in the spring maturity, inferred growth rates and diet (Shine and early summer in several lizard species 1984a). These data were acquired from 953 (Chapman and Dell 1985; How and Kitchener 1983; specimens of 13 species in collections at Australian How et al. 1986, 1987, 1990). It is also a region museums and were collected over extended time where the diversity of potential lizard prey items is frames from species, often with large geographic well documented (How and Dell 1993). Of the two ranges, that covered numerous climatic zones. sympatric species not considered in this study, S. Shine (1984a) found that there was a marked semifasciatlls has a markedly specialised dietary and season of reproductive activity in these species; reproductive pattern (Shine 1984a), while S. that at least five species were oviparous with larger littoralis Storr, although represented in collections, females having larger clutches; that females were is the focus of an independent study (Jennings, larger than males and that there was a clear personal communication). specialisation in the diet of many species. A study of 736 specimens of 11 species of Simoselaps and Vermicella occurring in Western METHODS Australia (Clarke and How 1995) showed that it Specimens in the collections of the Western was possible to determine the sex of each Australian Museum were selected to encompass individual by recording the tail to body ratio of the winter-rainfall dominated biogeographic specimens. Females of all species had smaller ratios regions, approximating the area of southwestern than males and these ratios did not overlap (Clarke Australia west of a line from Geraldton to and How 1995). Esperance (Figure 1). This study examines the reproductive biology Specimens were measured to the nearest and diet of four of the six species of 5imoselaps [5. millimetre for both snout to vent length (SYL) and 58 N.R. Strahan, R.A. How, J. Dell ~ 0 ... 0 0 • ... 00 00 0 0 00 0 0 0 0 Figure 1 Distribution of Simoselaps bertholdi (0), S. bimaclIlatlls (£), S. calonotos (*) and S. fasciolatlls (e) specimens from southwestern Australia examined for this study. The outline of the Interim Biogeographic Regionalisation for Australia (IBRA) regions of southwestern Australia are also depicted. tail to vent length (TVL). After dissection of the bimaculatus 52; S. calonotos 62; S. fasciolatus 28. lower abdomen with scissors the gonads were These comprised 32%, 75%, 94% and 55% of measured to the nearest 0.01 of a millimetre using individuals that had been examined previously by digital callipers. The length (1) and width (w) of Shine (1984a) for the respective species. each testis was recorded as was the length and Statistical comparisons were made using the width of all yolked ovarian follicles and oviductal Statistix (1996) program. Significant ANOVA's eggs. The volume of each testis was combined to were further examined using the Least Significant provide a measure of testes volume, while clutch Difference test (a=0.05) to compare means. volume was calculated by combining the volumes of individual oviductal eggs. Stomach contents were removed for later RESULTS examination and identification. Prey items were Reproduction identified to the species level by comparison with Maturity the extensive reference material of lizard taxa in The size range of adults and the number of adults the collections of the Western Australian Museum. of each sex of Simoselaps examined during this Calculations were made on the volume of testes, study are presented in Table 1. yolked ovarian follicles or oviductal eggs using the The sample sizes of SinlOselaps species available formulae for a prolate spheroid: from southwestern Australia were small and, as Vol. = 4/3n(w/2)2(1/2) such, probably present a very conservative The number of specimens of each Simoselaps estimate of size at sexual maturity amongst the species examined was: S. bertholdi 117; S. species of Simoselaps examined in this study. Biology of burrowing snakes 59 QI 100 .~ U 80 ::J ;:..,. 60 ................~._".:.::: "...0 Q. 40 S ber/holdi ...QI 20 ~ 0 0 :::J Ol a. u > u c .0 C month -, :::J Q) o Q) 1'0 Q) :::J ~ (j) o Z o -, u. -, 2 4 4 3 2 3 2 3 340 QI 100 > ;: ---- U 80 ::J 60 "...0 S. bimacllla/lIs Q. 40 QI ... 20 -"'---- ~ 0 0 Ol a. > u c .0 Q. >- C month "5 U 0 Q) Q) ro 1'0 -, :::J Q) 1'0 ~ :::J ~ (j) 0 z 0 -, u. ~ ~ -, N 2 3 0 2 3 3 2 2 QI 100 ;:> u 80 ::J 60 "8... Q. 40 S. calon%s QI... 20 ~ 111:1. 0 0 Ol Q. u .Cl >- c "5 0 > c ro Q. 1'0 month -, :::J CIl 0 CIl 1'0 CIl ~ ::J ~ (j) 0 z 0 -, u. ~ ~ -, N 3 4 5 0 3 3 0 0 0 3 2 100 QI ;:> 80 U ::J 60 "0 is. 40 fasciola/lis QI s. ... 20 ~ 0 0 Ol a. o > c .0 Q. C month ::J CIl o al 1'0 CIl ~ :::J ~ (j) o Z o -, U. -, o o o o 2 3 o o 2 0 Figure 2 Percentage of adult females that were reproductively active in each month of the year (N size) for )IIIIOS.['1I1/)5 spp from southwestern Australia in Western Australian Museum col!ectlOns. Consequently, adult specimens were determined these species from throughout their as those larger than the smallest reproductively range. Table 1 indicates the size range of adults of active specimen examined in either this study or both sexes determined from this study and by that of Shine (1984a), who examined the Shine (1984a). reproductive state of a larger sample of each of The SVL of the smallest female with yolked 60 N.R. Strahan, R.A. How, J. Dell Table 1 Size range and number of each sex for the adult specimens of Simoselaps species examined in this study and by Shine (1984a). STUDY S. bertholdi S. bimaculatlls S. calonotos S. fasciolatlls S. semifasciatus Adult <3 <3 147-196(44) 244-339(15) 171-280(25) 201-306(10) Adult <jl <jl 172-264(33) 237-406(22) 226-254(25) 288-363(9) Adult <3 <3 -Shine 143-225(105) 230-403(25) 182-230(30) 202-300(18) 192-336(68) Adult <jl <jl-Shine 164-312(98) 264-406(22) 186-251(32) 231-352(13) 196-334(74) ovarian follicles or eggs for each Simoselaps species (2 yolked follicles), the two reproductively active S. in our samples from southwestern Australia was fasciolatus were in November (4 eggs) and December 172 mm in S. bertlzoldi, 237 mm in S. bimaculatus, (5 eggs), while two reproductively active S. calonotos 226 mm in S. calonotos and 288 mm in S. fasciolatus. occurred in October (4 eggs) and December (3 eggs) These data are in close agreement with those of and another female had a small, slightly yolked Shine (1984a). follicle in May. Male reproductive maturity was assessed from a The volume of the testes of adult Simoselaps males plot of testes volume on SVL and determined as showed no significant variation between months, occurring at a size above the smallest male with except in S. bertlzoldi. In S. bertlzoldi testes volume markedly enlarged testes size (Tables 1, 2). These was significantly larger in October than in sizes are again in close agreement with those of November, June, September, December, January, Shine (1984a), who used the opaqueness of efferent August and April, but not other months (Table 2). ducts as an additional criterion for determining In all other species the monthly sample sizes were maturity in males.
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