Reproductive Biology of the Peninsula Dragon Lizard, Ctenophorus fionni Greg Johnston Journal of Herpetology, Vol. 33, No. 4. (Dec., 1999), pp. 694-698. Stable URL: http://links.jstor.org/sici?sici=0022-1511%28199912%2933%3A4%3C694%3ARBOTPD%3E2.0.CO%3B2-K Journal of Herpetology is currently published by Society for the Study of Amphibians and Reptiles.
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to the knowledge of hemoglobin evolution in turtles. Comparison of functional differences in isohemoglobins of S. odoratus and other species also would contribute to an understanding of the adaptations facilitating underwater aerobic hibernation.
C 0 2 and organic phosphates on oxygen affinity of sea turtle hemoglobins. Respir. Physiol. 48:75-87. L. A., Y. K. SONG,AND R. B. REEVES. MAGINNISS, 1980. Oxygen equilibria of ectotherm blood containing multiple hemoglobins. Respir. Physiol. 42:329-343. -, S. S. TAI'I'ER,A N D L. S. MILLER. 1983. Effect of Acki~mvledgn~ei~ts.-Thisstudy was part of P. King's chronic cold and submergence on blood oxygen doctoral research project at North Carolina State Unitransport in the turtle, Cllryseiiys picta. Respir. versity and funded by a North Carolina Agricultural Physiol. 53:15-29. Research Service grant to H. Heatwole and Sigma Xi MUSACCHIA, X. J., AND M. L. SEIVERS. 1956. Effects of Grants-in-Aid of Research to P. King. We would like induced cold torpor on hemoconcentration in to thank C. Sullivan for assistance with electrophoreClzrysetnys picta. Fed. Proc. Amer. Soc. Exp. Biol. 15: sis equipment and advice. 134. SULLIVAN, B., AND A. RIGGS.1967. Structure, function and evolution of turtle hemoglobin - 11. Electrophoretic studies. Comp. Biochem. Physiol. 23:449BRAMAN, J. C., C. B. STALNAKER, T. M. FARLEY, AND 458. G. T. KLAR.1977. Starch gel electrophoresis of rain- TUN,N., A N D A. H. HOUSTON. 1986. Temperature, oxbow trout, Saln~ogairdi~eri,and cutthroat trout, Salygen, photoperiod, and the hemoglobin system of n ~ oclarkii, hemoglobins. Comp. Biochem. Physiol. the rainbow trout, Salrno gairdneri. Can. J. Zool. 64: 56B:435-437. 1883-1888. CARR,A. 1952. Handbook of Turtles. Cornell Univ. ULTSCH,G. R. 1988. Blood gas, hematocrit, plasma ion Press, Ithaca, New York. concentrations, and acid base status of musk turDUNSON,W. A. 1960. Aquatic respiration in Trioi~yx odoratus) during simulated hibertles (Sternotl~cr~is spiliifer i*sper. Herpetologica 16:277-283. nation. Physiol. Zool. 61:78-94. ERNST,C. H. 1986. Ecology of the turtle, Steri~otiwrus -. 1989. Ecology and physiology of hibernation odoratus, in southeastern Pennsylvania. J. Herpetol. and overwintering among freshwater fishes, tur20:341-352. tles, and snakes. Biol. Rex.. 64:435-516. GORDON,A. H. 1969. Electrophoresis of proteins in -, A N D D. C. JACKSON. 1995. Acid-base status polyacrylamide and starch gels. In T. S. Work and and ion balance during simulated hibernation in E. Work (eds.), Laboratory Techniques in Biochemfreshwater turtles from the northern portions of istry and Molecular Biology, Vol. 1, part I, p p 1their ranges. J. Exp. Zool. 273:482-493. 149. North-Holland, Amsterdam. -, C. V. HERBERT, AND D. C. JACKSON. 1984. The GRAHAM,J. B. 1990. Ecological, evolutionary, and comparative physiology of diving in North Amerphysical factors influencing aquatic animal respiican freshwater turtles. I. Submergence tolerance, ration. Amer. Zool. 30:137-146. gas exchange, and acid-base balance. Physiol. Zool. GRAHAM, T. E. 1995. Habitat use and population pa57(6):620-631. rameters of the spotted turtle, Clenln~ysglittata, a species of special concern in Massachusetts. Chel. Accepted: 2 July 1999. Conserv. Biol. 1:207-214. HARRINGTON, J. P. 1986. Structural and functional zus studies of the king salmon, O i ~ c o r ~ i ~ c l tslzm~lytsclza, hemoglobins. Comp. Biochem. Physiol. 848: 111-116. /olirr~nlof H~r~ii.tology, WI. 33, No. 4, pp. 694498, 1999 and Reptlles HATTlNGH, J. 1976. Haemoglobins in Labeo umbratus: Copyright 1999 Society for the Study of Amph~b~ans the influence of temperature and oxygen. Sth. Afr. J. Sci. 72:27-28. Reproductive Biology of the Peninsula H u n o ~K. , E., AND C. 1. GOODNIGHT. 1957. Variations in blood chemistry of turtles under active and hiDragon Lizard, Ctenophorus fionni bernating conditions. Physiol. Zool. 30:198-207. GREG JOHNSTON",' School of Biological Scieilces, Flillders INGERMANN, R. L. 1992. Structure-function relationUi~iilersityof Soutlz Australia, GPO Box 2100, Adelaide ships of the ectothermic vertebrate hemoglobins. In C. P. Magnum (ed.) Advances in Comparative and 5001, Australia.
Environmental Physiology, Blood and Tissue
Oxygen Carriers, Vol. 13, pp. 411-431. SpringerDragon lizards (family Agamidae) are a diverse,
Verlag, Berlin. widely distributed family which show considerable
KING,P. 1995. Aquatic respiration of aquatic turtles. variation in their reproductive biology. They may be Unpubl. Ph.D. Diss., North Carolina State Univ., sexual or parthenogenetic (Hall, 1970), oviparous or Raleigh. o\~o\~iviparous (Greer, 1989). They may lay clutches of -, AND H. HEATWOLE. 1994. Non-pulmonary re- one or two eggs continuously throughout the year (Inspiratory surfaces of the chelid turtle Elseya latis- ger and Greenberg, 1966; Allison, 1982), or lay clutchtert114ti1.Herpetologica 50:262-265. es of 2-35 eggs in a strongly seasonal pattern (Greer, K o u s s o u r ~ ~ o S., s , S. KAPAROS, AND D. STATHAKOS. 1986. Multiple hemoglobins in Triturus cristatus: their study by analytical isoelectrofocusing. Comp. "resent Address: Mitrani Centre for Desert EcolBiochem. Physiol. 83B:475-481. ogy, Ben-Gurion University of the Negev, Sede Boqer, LUTZ,P. L., A N D G. N. LAPENNAS. 1982. Effects of pH, 84990, ISRAEL. E-mail:
[email protected]
SHORTER COMMUNICATIONS 1989). This variation in reproductive and life-history traits emphasises the need to collect data on each individual species, rather than relying on inference from the relatively few species for which data are available. The Peninsula Dragon Lizard (Ctenoplrorusjonni) is endemic to the Eyre Peninsula in South Australia (Houston, 1974).It is a medium-sized (596 mm snoutvent length) diurnal, rock dwelling lizard. This paper presents data on reproduction in C. jonizr for the first time, and provides information on body size differences between sexes, size at maturity, reproductive cycles, clutch sizes and frequency of reproduction in this species. The study of reproductive cycles was based on 120 (74 males, 46 females) C. fionni collected from Iron Duke (33"15'S, 137O07'E) and deposited in the South Australian Museum. This collection was made between April, 1985 and April, 1986 just prior to Iron Duke being cleared for an extensive open cut mine. Over this period the area was visited in most months. During each visit, between four and fourteen lizards were collected by hand, and killed within 12 h of capture. They were fixed in 10% formalin and stored in 70% alcohol. For each lizard, snout-vent length (SVL) was measured to the nearest mm and the reproductive tract was inspected through a small, lateral incision into the peritoneal cavity. Body size of mature males and females was compared using a oneway analysis of variance (ANOVA; Sokal and Rohlf, 1981). In females, the number of vitellogenic follicles and corpora lutea were counted for each ovary. The maximum diameter of the largest follicle was measured and any oviductal eggs were counted. The presence of vitellogenic follicles or corpora lutea were taken as an indication of reproductive maturity. In males the maximum width (w) and length (1) of the right testis were measured and used to calculate the testicular volume (V) in ml using the standard formula for a prolate spheroid:
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flywire lids and overhead, broadspectrum heat lamps. While in captivity females were offered food (mealworms, crickets, or cockroaches) every day and water was available ad libitum. Dampened paper towelling was placed on the floor of the containers to prevent dehydration of any eggs. Each female was given weekly intraperitoneal injections of Oxytocin (HERIOT AGVET, Rowville, Victoria) at a dosage of 6.0mllkg until they laid a clutch of eggs (a maximum of two weeks). The SVL and weight after laying was recorded for each female, along with clutch size, egg lengths, egg widths, and egg weights. Not all measurements were taken from all clutches, so sample sizes varied for different statistical analyses. After oviposition, all live females were returned to the site from which they were captured. All lengths were measured with vernier callipers to the nearest 0.lmm. Weights were measured to the nearest 0.1 g with a Mettler top-loading electronic balance. A mark-recapture study at Midgee Rocks between 1991 and 1994 (Johnston, 1997) provided additional information on the frequency of reproduction. During this field study females on a study plot were captured at least once every two weeks between August and December in all years except 1991. The site was not visited during November of 1991. Iron Duke, Midgee Rocks, and Siam Station are all on Eyre Peninsula and share a Mediterranean climate, with hot dry summers and cool, moist winters. Iron Duke is the southernmost peak in the Middleback Ranges, a meridional ridge of Ironstone. A description of the Iron Duke area is provided by Johnston (1982). Midgee Rocks and Siam Station are a series of granite domes, with flat sandy planes between the rock outcrops. At Iron Duke, mature males were significantly larger (SVL: 2 = 75.9mm; SD = 5.64) than mature females (SVL ?, = 70.3mm; SD = 5.92)(F,,,, = 20.01, P < 0.0001) (Fig. 1). The largest male had a SVL of 84.lmm, whereas the largest female had a SVL of 83.8mm. The smallest female from Iron Duke with A gonadosomatic index (GSI) was calculated as the vitellogenic follicles or corpora lutea had a SVL of ratio between testis volume and SVL. The epididy- 60mm. The smallest male with opaque epididymides mides were classified as translucent (empty) or had a SVL of 55mm. Enlarged, vitellogenic ovarian follicles were present opaque (containing sperm). All males above the minimum size at which opaque epididymides were found in females from Iron Duke in September and Novemwere regarded as mature. Mature male lizards were ber (Fig. 2). Oviductal eggs were present in November (2 of 8 females) and December (4 of 6 females). pooled into collections from spring (August-October), Among mature males from Iron Duke, the GSI was summer (November-January), autumn (FebruaryApril) and winter (May-July) to allow statistical anal- significantly different between seasons (F, j, = 22.25, ysis of the testicular cycle. A oneway ANOVA was P < 0.001). The spring mean was significantly higher used to test the null hypothesis that the GSI of mature than the means in all other seasons (Scheffe P < 0.05) males did not differ between these four seasons. All (Fig. 3). The testes were enlarged with obvious semimales above the smallest mature size were included niferous tubules during August and September. In all other months the testes were flaccid and seminiferous in this analysis. Observations of the reproductive cycle based on the tubules were not visible on gross dissection. Clutch size at Iron Duke, based on the number of specimens from Iron Duke were supplemented with observations made at two other locations that were vitellogenic follicles or the number of oviductal eggs, each visited several times between 1991 and 1994. varied from 2 to 6 (2 = 4.15, SD = 1.09, N = 20). Gravid females were collected in October and Novem- Females from Midgee Rocks laid between 3 and 6 (2 = 4.43, SD = 1.27, N = 7). Females from Siam also ber from Siam Station (32"33'S, 136"43'E; 110 km north west of Iron Duke) and Midgee Rocks (33"25'S, laid between 3 and 6 (2 = 4.67, SD = 1.12, N = 9). 137"05'E; 12 km south of Iron Duke) in 1992 to deter- There was no significant variation in clutch size mine clutch size. These females were taken to the lab- among the three populations (Kruskal-Wallis ANOVA, oratory, where they were housed individually in clear X' = 1.872, P = 0.599). Nor were there any differences plastic containers (250mm x 200mm x 100mm) with between the populations in slopes of clutch size on
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.d / L l l i 0 .
a
J
SNOUT-VENT LENGTH (mm)
F M A M J J A S O N D
MONTH
1, size frequency distribution of male and female Ctt,nqdzorus fio~znicollected at Iron Duke.
FIG.3. Annual reproductive cycle in male Cterlophorus fionni from Iron Duke. Each point may represent more than one lizard.
maternal SVL. so clutches were pooled. Using these pooled data, there was no significant relationship between clutch size and maternal SVL (F, = 0.99, P = 0.158)(Fig. 4). However, when the two outlying large females who produced small clutches were excluded from the analysis, the regression was significant (Fl,,j = 7.65, P = 0.009). Both outlying females were collected from Iron Duke, one in November and one in December. Corpora lutea were present in one of them. This suggests that these small clutches may have been the second clutch produced by these females in that year. Measurements of freshly laid eggs and hatchlings from Midgee Rocks are given in Table 1. Relative clutch mass (RCM) was expressed as the clutch mass divided by the post-ovipositional female mass. Several females (N = 7) brought into captivity
from Midgee Rocks and Siam Station to obtain clutch data died shortly after producing a clutch of eggs. The arcsine transformed values of RCM were compared between females who survived and females who died using a oneway ANOVA. RCM was significantly smaller for females who survived after producing a clutch (RCM = 0.30-0.45; i= 0.35; N = 10) than for those females who died shortly after producing a clutch (RCM = 0.51-0.86; 2 = 0.66; N = 7; F, ,; = 63.56, P = 0.0001).Females who died weighed less (i = 8.9, SD = 2.59) than females who survived (= i 11.0,SD = 2.39), but this difference was not significant (ANOVA F, ,: = 3.07, P = 0.1). Among the 20 adult females collected from August to December at Iron Duke, most (85%)had vitellogenic follicles, oviductal eggs, or corpora lutea. This indicated either that they would produce a clutch of eggs, or that they had already produced a clutch.
jj
J F M A M J
J A S O N D
MONTH FIG. 2. Annual reproductive cycle in female Cteiz= ovarian follicles, ophorus fionni from Iron Duke. 0 = oviductal eggs (not to scale of y-axis). Each point may represent more than one lizard.
SNOUT-VENT LENGTH (mm) FIG. 4. Relationship between clutch size and female Snout-Vent Length in Ctet~ophorusfionnl.
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N
Length (mm) Width (mm) Weight (g)
45 45 45
SVL (mm) Weight (g)
23 23
x
Eggs 20.3 11.0 1.4 Hatchlings 30.9 1.3
SD
Min-max
1.52 0.81 0.28
16.8-24.0 9.0-12.7 0.7-1.8
1.68 0.22
28.0-34.8 0.9-1.7
Repeated captures of adult females in the field at Midgee Rocks showed that the proportion of females found gravid at least once within in a breeding season ranged from 19 of 30 (63%) in 1991 to 1 of 15 (7% in 1994. Most mature females found gravid (43 of 45 = 96%) at Midgee over the entire four year period produced only one clutch in a breeding season, implying that multiple clutches were unusual. The two females that produced two clutches in a season were recorded gravid in late September and again in late November. Most females (41 of 45 = 91%) were recorded gravid in only one breeding season during the study, but four females were gravid in two successive breeding seasons. No females were recorded gravid for more than two successive breeding seasons at Midgee Rocks. The results presented here show that C. fionni reproduce sexually. Males had enlarged testes in August and September. Females were gravid between September and December, and normally produced a single clutch of between two and six eggs. The eggs incubated over the summer, and hatchlings appeared in the field in January and February (Johnston, 1997). Thus Peninsula dragons have a similar reproductive season to most other Australian dragon lizards (Bradshaw, 1981; James and Shine, 1985; Greer, 1989). Males are generally larger than females among Agamid lizards (Fitch, 1981). This pattern is thought to be a result of sexual selection brought about by male competition (Stamps, 1983; Johnston, 1997). In C. fiolzl~ithe males and females matured at similar body sizes, and reach similar maximum body sizes. However, mature males were generally larger than females (Fig. 1). This difference is presumably a result of differences between the sexes in growth pattern or survivorship. The reproductive cycles of male and female C.fion~zi at Iron Duke coincided, so that the testes were producing sperm at the same time vitellogenesis occurred (Figs. 2 and 3). Mating presumably followed shortly afterwards in August and September. Thus sperm storage by females, which occurs in some lizards at similar latitudes (Cuellar, 1966; Smyth and Smith, 1968), may not occur in Peninsula dragons. No obvious sexual activity was observed in the field after November (Johnston, 1997). However, adult males maintained opaque epididymides throughout the year. In so far as this indicates the presence of spermatozoa, it appears that males store sperm in the epididymides even when the testes were not enlarged (i.e., actively producing sperm). This may be advantageous for males if females that produce a second clutch mate between clutches.
697
Females at Midgee Rocks may reproduce in two consecutive years, although most captured in more than one year were found gravid only once. In captivity females may lay sequential clutches at 40 d intervals (Johnston, 1997). This is presumably the time necessary for vitellogenesis, fertilization, and formation of new eggs, and places a limit on the ability of females to produce multiple clutches in most years. Like other dragon lizards (Bradshaw et al., 1991), Peninsula dragons are able to respond to appropriate environmental conditions by extending their reproductive season. Two female Peninsula dragons that were held in captivity under conditions of excess food availabilitv during the svrine and summer of 19911992 produced mLltiple ;lut&es (Kranz, 1992). Wild females from the same population produced no more than a single clutch in the same year (unpublished data). Females that invested heavily in producing a clutch (RCM > 0.5) died, whereas females who survived invested less heavily in reproduction (RCM 5 0.46). These observations suggest that food availability may limit the ability of females to reproduce, and almost certainly limits their ability to produce multiple clutches. In summary, male and female C. fionni matured at about the same size and reached a similar maximum size, but on average mature males were larger than females. The reproductive cycles of males and females were highly synchronised. Testes and ovarian follicles were largest during the austral spring. Most females produced a single clutch of between two and six eggs, and many females died after laying eggs. Those females who survived had invested less in reproduction than females who died. The ability to produce multiple clutches appears to be related to the availability of food during spring. Acknmu1ed~ments.-This paper was partially funded by an Australian Postgraduate Research Scholarship held at Flinders University of South Australia. The Broken Hill Proprietary Company Limited allowed access to Iron Duke. The Carmody and Pumpa families allowed access to Midgee Rocks and Siam Station, respectively. Craig Martins and Barbara Levings helped collect lizards at Iron Duke. Mike Bull and Paula Johnston commented on the manuscript.
ALLISON,A. 1982. Distribution and ecology of New Guinea lizards. 111J. L. Gressitt (ed.), Biogeography and Ecology of New Guinea, pp. 803-813. Dr W. Junk, The Hague. S. D. 1981. Ecophysiology of Australian BRADSHAW, desert lizards: studies on the genus Amplzibolurus. 111 A. Keast (ed.), Ecological Biogeography of Australia, pp. 1394-1434. Dr W. Junk, The Hague. BRADSHAIV, S. D., H. SAINTGIIIONS,AND F. J. BRADSHAW.1991. Patterns of breeding in two species of agamid lizards in the arid subtropical Pilbara region of Western Australia. Gen. Comp. Endocrinol. 82:407324. CUELLAR, 0 . 1966. Oviducal anatomy and sperm storage structures in lizards. J. Morphol. 119:7-20. FITCH,H. S. 1981. Sexual size differences in reptiles. Misc. Pub. Mus. Nat. Hist. Kansas. 70:l-72. GREER, A. E. 1989. The Biology and Evolution of Aus-
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tralian Lizards. Surrey Beatty, Norton, New South Wales. HALL,W. P. 1970. Three probable cases of parthenogenesis in lizards (Agamidae, Chamaeleontidae, Gekkonidae). Experientia 26:1271-1273. HOLSTOK,T. F. 1974. Revision of the Amphibolitrits decresii complex (Lacertilia: Agamidae) of South Australia. Trans. Roy. Soc. S. Aus. 98:49-60. INGER, R. F., AND GREENBERG, B. 1966. Annual reproductive patterns of lizards from a Bornean rain forest. Ecology 47:1007-1021. JAMES, C. D., A N D SHINE, R. 1985. The seasonal timing of reproduction: a tropical-temperate comparison in Australian lizards. Oecologia 67:464-474. JOHNSTON, G. R. 1982. The herpetofauna of the Middleback Range area, South Australia 1. An annotated checklist. Herpetofauna (Australia) 14:52-60. -. 1997. Behavioural Ecology of the Peninsula Dragon Lizard, Cte1lopi:orus fiorlrli. Unpubl. Ph.D. Thesis, Flinders Unit: South Australia. KRAKZ,C. 1992. Results of breeding the peninsula dragon, Cteilopkorus Fiorlni. Monitor 4:54-57. SMYTH, M. AND SMITH,M. J. 1968. Obligatory sperm storage in the skink Henliergis peroilii. Sclence 161: 575-576. SOKAL,R. R. AND ROHLF,F. J. 1981. Biometry. W. H. Freeman, San Francisco. STAMPS,J. A. 1983. Sfxual selection, sexual dimorphism and territoriality. I11 R. B. Huey, E. R. Pianka, and T. W. Schoener (eds.), Lizard Ecology: Studies of a Model Organism, pp. 169-204. Harvard Univ. Press, Cambridge, Massachusetts. Accepted: 4 July 1999
/ourrral o Hrrprtolop, Vol. 33, KO. 4, pp. 698-i02, 1999 Copyridt 1999 so'hetyfor the Study of Amphibians and
Reptiles
Intraspecific Variation of the Hemipenis in Siphlophis and Tripanurgos HUSSAMZAHERAND ANA LUCIAC. PRLDENTE, Depnrtmeilto de Zoologin, UrlirursiiiniC de Srio Pnulo, Cnixn Postal 11461, Slio Pniilo, SP 05422-970, Brail, E-mail: /:;n/ur@ib. us;,. br;
[email protected] The use of hemipenial characters in systematic studies of snakes has increased substantially in the past few years (see Savage, 1997 for a review). Such renewed interest is due mostly to the discovery of a large number of potentially informative characters and to the improvement of techniques that facilitate the preparation of uneverted organs (Pesantes, 1994). Hemipenial variation among species and higher categories have been acknowleged as an important source of characters of potential phylogenetic value, to the extent that some workers have taken the more extreme position of considering hemipenial features to be more reliable than other morphological characteristics. This view have been expressed by Dowling (1967:138) who he argued that "the squamate hemipenis is a structure that (unlike teeth, skull, or scutellation) has no obvious correlation with the ecology,
food habits, or locomotion of the animal. . . [and for this reason it] may give better information on the genetic relationships than some habit- or habitat-correlated characteristics" [our brackets]. Such a belief is here viewed with caution inasmuch as increasing knowledge about interspecific variation in snake taxa is showing just the opposite situation, wherein hemipenial structures show highly complex transformation series within moderately diversified assemblages at low taxonomic levels (e.g., Myers, 1974; Zaher, 1994). Similarly, there is no reason to assume that hemipenial morphology in snakes is highly conservative intraspecifically, although potential intraspecific variation has been documented rarely (e.g., Inger and Marx, 1962; Keiser, 1974; McDowell, 1979; Cole and Hardy, 1981). Here we describe the hemipenial variation found in Triparnirgos coit~yressusand in the six species comprising the genus Sipi~lophis. The terminology used follows Zaher (in press). In a phylogenetic analysis of the tribe Pseudoboini (Zaher, 1994, 1996), Tripnr~urgoscort~pressus appeared to be nested within Sipillopilis, as the sister-group of a clade composed of S. cerz>ir:us,S. yitlclwv, and S. Iciicoccyi~nlus.For this reason, Triynrnirgos will be consid(in ered herein as a synonym of the genus Siyi~ioyll~s conformity with Article 24 of the International Code of Zoological Nomenclature). Additionally, Tri;~nilurg o s c o n : p r e s ~ t s s h a r e swith all the species of Sipklopilis at least four unambiguous synapomorphies within the pseudoboine radiation (Zaher, 1994). These features include (plesiomorphic condition in parentheses): (1) maxillan process of the prefrontal reduced or absent (a well-detreloped process is present in all other pseudoboines); (2) Meckelian canal closed from the anterior region of the splenial to the tip of the dentary (all other pseudoboines have an open canal); (3) posteroventral region of the nasal gland sharp-edged (this region is rounded in all the other pseudoboines); (4) dorsal region of the maxilla with a well-detreloped, ossified notch at the level of the articulation with the prefrontal (no notch is present in the other pseudoand boines). Zaher (1994) did not study S. ~~nroritzo?cli, the only available specimen of S. leucocq~ilalusis in a poor state of preservation. Recently, Prudente (unpubl. data) reanalyzed Zaher's data, adding newr information and including 5. ztoroiltzmc~i, and confirmed the phylogenetic position of T coit~yressuswithin the genus Sipi~lopkis. Intraspecific variation was found among specimens representing all the species of Sipi~loyl:is,each species showing the same two distinctly different types of hemipenes. The first type, the "Y-shaped condition," clearly represents the primitive state among pseudoboines. The second type, which we designate the "Tshaped condition," is present also in Oxyrilopus clnthrntus, and it may represent a synapomorphy of a more inclusive clade of pseudoboine snakes (Zaher, 1994). However, it is not the purpose of the present paper to discuss the possible optimization schemes for this character. A total of 27 fully everted and inflated hemipenes were available for the following species (Table 1): S. cerainus (6) (Fig. I), S. pulcizer (7), S, longicnudatits (5), S. leitcoctyhalus (2), S. itoroiltzoicli (2), S. con:pressits (5) (Fig. 2). All species hatre both "T- and Y-shaped" hemipenial types. We reject the possibility that these dif-
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Literature Cited Annual Reproductive Patterns of Lizards from a Bornean Rain Forest Robert F. Inger; Bernard Greenberg Ecology, Vol. 47, No. 6. (Nov., 1966), pp. 1007-1021. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28196611%2947%3A6%3C1007%3AARPOLF%3E2.0.CO%3B2-Q
Obligatory Sperm Storage in the Skink Hemiergis peronii Michael Smyth; Meredith J. Smith Science, New Series, Vol. 161, No. 3841. (Aug. 9, 1968), pp. 575-576. Stable URL: http://links.jstor.org/sici?sici=0036-8075%2819680809%293%3A161%3A3841%3C575%3AOSSITS%3E2.0.CO%3B2-I
