Reproductive Biology of the Peninsula Dragon Lizard, Ctenophorus fionni

Greg Johnston

Journal of Herpetology, Vol. 33, No. 4. (Dec., 1999), pp. 694-698.

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694 SHORTER COMMUNICATIONS

to the knowledge of hemoglobin evolution in turtles. C 0 2 and organic phosphates on oxygen affinity of
Comparison of functional differences in isohemoglo- sea turtle hemoglobins. Respir. Physiol. 48:75-87.
bins of S. odoratus and other species also would con- MAGINNISS, L. A., Y. K. SONG,AND R. B. REEVES. 1980.
tribute to an understanding of the adaptations facili- Oxygen equilibria of ectotherm blood containing
tating underwater aerobic hibernation. 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 Uni- transport 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 electrophore- Clzrysetnys 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. Electro-
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BRAMAN, 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, ox-
bow trout, Saln~ogairdi~eri,and cutthroat trout, Sal- ygen, 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 tur-
DUNSON,W. A. 1960. Aquatic respiration in Trioi~yx tles (Sternotl~cr~is odoratus) during simulated hiber-
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, tur-
20:341-352. 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 Biochem- freshwater turtles from the northern portions of
istry and Molecular Biology, Vol. 1, part I, p p 1- their 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 Amer-
physical factors influencing aquatic animal respi- ican 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 pa- 57(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
studies of the king salmon, O i ~ c o r ~ i ~ c l tslzm~lyts-
zus
clza, hemoglobins. Comp. Biochem. Physiol. 848:
111-116. WI. 33, No. 4, pp. 694498, 1999
/olirr~nlof H~r~ii.tology,
HATTlNGH, J. 1976. Haemoglobins in Labeo umbratus: Copyright 1999 Society for the Study of Amph~b~ans and Reptlles
the influence of temperature and oxygen. Sth. Afr.
J. Sci. 72:27-28.
H u n o ~K. , E., AND C. 1. GOODNIGHT. 1957. Variations Reproductive Biology of the Peninsula
in blood chemistry of turtles under active and hi- Dragon Lizard, Ctenophorus fionni
bernating conditions. Physiol. Zool. 30:198-207.
INGERMANN, R. L. 1992. Structure-function relation- GREG JOHNSTON",' School of Biological Scieilces, Flillders
ships of the ectothermic vertebrate hemoglobins. In Ui~iilersityof Soutlz Australia, GPO Box 2100, Adelaide
C. P. Magnum (ed.) Advances in Comparative and 5001, Australia.

Environmental Physiology, Blood and Tissue

Oxygen Carriers, Vol. 13, pp. 411-431. Springer- Dragon 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 (In-
spiratory surfaces of the chelid turtle Elseya latis- ger and Greenberg, 1966; Allison, 1982), or lay clutch-
tert114ti1.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 Ecol-
Biochem. 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: gregj@bgumail.bgu.ac.il
 SHORTER COMMUNICATIONS 695

1989). This variation in reproductive and life-history flywire lids and overhead, broadspectrum heat lamps.
traits emphasises the need to collect data on each in- While in captivity females were offered food (meal-
dividual species, rather than relying on inference from worms, crickets, or cockroaches) every day and water
the relatively few species for which data are available.was available ad libitum. Dampened paper towelling
The Peninsula Dragon Lizard (Ctenoplrorusjonni) is was placed on the floor of the containers to prevent
endemic to the Eyre Peninsula in South Australia dehydration of any eggs. Each female was given week-
(Houston, 1974).It is a medium-sized (596 mm snout- ly intraperitoneal injections of Oxytocin (HERIOT
vent length) diurnal, rock dwelling lizard. This paper AGVET, Rowville, Victoria) at a dosage of 6.0mllkg
presents data on reproduction in C. jonizr for the firstuntil they laid a clutch of eggs (a maximum of two
time, and provides information on body size differences weeks). The SVL and weight after laying was record-
between sexes, size at maturity, reproductive cycles, ed for each female, along with clutch size, egg lengths,
clutch sizes and frequency of reproduction in this spe- egg widths, and egg weights. Not all measurements
cies. were taken from all clutches, so sample sizes varied
The study of reproductive cycles was based on 120 for different statistical analyses. After oviposition, all
(74 males, 46 females) C. fionni collected from Iron live females were returned to the site from which they
Duke (33"15'S, 137O07'E) and deposited in the South were captured. All lengths were measured with ver-
Australian Museum. This collection was made be- nier callipers to the nearest 0.lmm. Weights were
tween April, 1985 and April, 1986 just prior to Iron measured to the nearest 0.1 g with a Mettler top-load-
Duke being cleared for an extensive open cut mine. ing electronic balance. A mark-recapture study at
Over this period the area was visited in most months. Midgee Rocks between 1991 and 1994 (Johnston,
During each visit, between four and fourteen lizards 1997) provided additional information on the frequen-
were collected by hand, and killed within 12 h of cap- cy of reproduction. During this field study females on
ture. They were fixed in 10% formalin and stored in a study plot were captured at least once every two
70% alcohol. weeks between August and December in all years ex-
For each lizard, snout-vent length (SVL) was mea- cept 1991. The site was not visited during November
sured to the nearest mm and the reproductive tract of 1991.
was inspected through a small, lateral incision into the Iron Duke, Midgee Rocks, and Siam Station are all
peritoneal cavity. Body size of mature males and fe- on Eyre Peninsula and share a Mediterranean climate,
males was compared using a oneway analysis of var- with hot dry summers and cool, moist winters. Iron
iance (ANOVA; Sokal and Rohlf, 1981). In females, the Duke is the southernmost peak in the Middleback
number of vitellogenic follicles and corpora lutea were Ranges, a meridional ridge of Ironstone. A description
counted for each ovary. The maximum diameter of the of the Iron Duke area is provided by Johnston (1982).
largest follicle was measured and any oviductal eggs Midgee Rocks and Siam Station are a series of granite
were counted. The presence of vitellogenic follicles or domes, with flat sandy planes between the rock out-
corpora lutea were taken as an indication of repro- crops.
ductive maturity. In males the maximum width (w) At Iron Duke, mature males were significantly larg-
and length (1) of the right testis were measured and er (SVL: 2 = 75.9mm; SD = 5.64) than mature females
used to calculate the testicular volume (V) in ml using (SVL ?, = 70.3mm; SD = 5.92)(F,,,, = 20.01, P <
the standard formula for a prolate spheroid: 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.
opaque (containing sperm). All males above the min- Enlarged, vitellogenic ovarian follicles were present
imum size at which opaque epididymides were found in females from Iron Duke in September and Novem-
were regarded as mature. Mature male lizards were ber (Fig. 2). Oviductal eggs were present in November
pooled into collections from spring (August-October), (2 of 8 females) and December (4 of 6 females).
summer (November-January), autumn (February- Among mature males from Iron Duke, the GSI was
April) 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 semi-
males above the smallest mature size were included niferous tubules during August and September. In all
in this analysis. other months the testes were flaccid and seminiferous
Observations of the reproductive cycle based on the tubules were not visible on gross dissection.
specimens from Iron Duke were supplemented with Clutch size at Iron Duke, based on the number of
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
ber from Siam Station (32"33'S, 136"43'E; 110 km = 4.43, SD = 1.27, N = 7). Females from Siam also
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
 SHORTER COMMUNICATIONS

SNOUT-VENT LENGTH (mm)

1, size frequency distribution of male and fe-
/
Lll
i.d
J
a
F M A M J J A S O N D

MONTH
0 .

FIG.3. Annual reproductive cycle in male Cterlo-

male Ctt,nqdzorus fio~znicollected at Iron Duke. phorus fionni from Iron Duke. Each point may repre-
sent more than one lizard.

maternal SVL. so clutches were pooled. Using these from Midgee Rocks and Siam Station to obtain clutch
pooled data, there was no significant relationship be- data died shortly after producing a clutch of eggs. The
tween clutch size and maternal SVL (F, = 0.99, P =
jj
arcsine transformed values of RCM were compared
0.158)(Fig. 4). However, when the two outlying large between females who survived and females who died
females who produced small clutches were excluded using a oneway ANOVA. RCM was significantly
from the analysis, the regression was significant (Fl,,j smaller for females who survived after producing a
= 7.65, P = 0.009). Both outlying females were col-
clutch (RCM = 0.30-0.45; i= 0.35; N = 10) than for
lected from Iron Duke, one in November and one in
those females who died shortly after producing a
December. Corpora lutea were present in one of them. clutch (RCM = 0.51-0.86; 2 = 0.66; N = 7; F, ,; =
This suggests that these small clutches may have been 63.56, P = 0.0001).Females who died weighed less (i
the second clutch produced by these females in that = 8.9, SD = 2.59) than females who survived (= i
year. Measurements of freshly laid eggs and hatchlings 11.0,SD = 2.39), but this difference was not significant
from Midgee Rocks are given in Table 1. (ANOVA F, ,: = 3.07, P = 0.1).
Relative clutch mass (RCM) was expressed as the Among the 20 adult females collected from August
clutch mass divided by the post-ovipositional female to December at Iron Duke, most (85%)had vitellogen-
mass. Several females (N = 7) brought into captivity ic follicles, oviductal eggs, or corpora lutea. This in-
dicated either that they would produce a clutch of
eggs, or that they had already produced a clutch.

J F M A M J J A S O N D

MONTH
FIG. 2. Annual reproductive cycle in female Cteiz- SNOUT-VENT LENGTH (mm)
ophorus fionni from Iron Duke. = ovarian follicles,
0 = oviductal eggs (not to scale of y-axis). Each point FIG. 4. Relationship between clutch size and fe-
may represent more than one lizard. male Snout-Vent Length in Ctet~ophorusfionnl.
 SHORTER COMMUNICATIONS 697

TABLE1. Measurements of eggs and hatchling Females at Midgee Rocks may reproduce in two
Cte~zqlzorusfionni from Midgee Rocks. consecutive years, although most captured in more
than one year were found gravid only once. In captiv-
Measurement N x SD Min-max ity females may lay sequential clutches at 40 d inter-
vals (Johnston, 1997). This is presumably the time nec-
Eggs essary for vitellogenesis, fertilization, and formation of
Length (mm) 45 20.3 1.52 16.8-24.0 new eggs, and places a limit on the ability of females
Width (mm) 45 11.0 0.81 9.0-12.7 to produce multiple clutches in most years.
Weight (g) 45 1.4 0.28 0.7-1.8 Like other dragon lizards (Bradshaw et al., 1991),
Hatchlings Peninsula dragons are able to respond to appropriate
SVL (mm) 23 30.9 1.68 28.0-34.8 environmental conditions by extending their repro-
Weight (g) 23 1.3 0.22 0.9-1.7 ductive season. Two female Peninsula dragons that
were held in captivity under conditions of excess food
availabilitv during the svrine and summer of 1991-
1992 produced mLltiple ;lut&es (Kranz, 1992). Wild
Repeated captures of adult females in the field at females from the same population produced no more
Midgee Rocks showed that the proportion of females than a single clutch in the same year (unpublished
found gravid at least once within in a breeding season data). Females that invested heavily in producing a
ranged from 19 of 30 (63%) in 1991 to 1 of 15 (7% in clutch (RCM > 0.5) died, whereas females who sur-
1994. Most mature females found gravid (43 of 45 = vived invested less heavily in reproduction (RCM 5
96%) at Midgee over the entire four year period pro- 0.46). These observations suggest that food availability
duced only one clutch in a breeding season, implying may limit the ability of females to reproduce, and al-
that multiple clutches were unusual. The two females most certainly limits their ability to produce multiple
that produced two clutches in a season were recorded clutches.
gravid in late September and again in late November. In summary, male and female C. fionni matured at
Most females (41 of 45 = 91%) were recorded gravid about the same size and reached a similar maximum
in only one breeding season during the study, but four size, but on average mature males were larger than
females were gravid in two successive breeding sea- females. The reproductive cycles of males and females
sons. No females were recorded gravid for more than were highly synchronised. Testes and ovarian follicles
two successive breeding seasons at Midgee Rocks. were largest during the austral spring. Most females
The results presented here show that C. fionni re- produced a single clutch of between two and six eggs,
produce sexually. Males had enlarged testes in Au- and many females died after laying eggs. Those fe-
gust and September. Females were gravid between males who survived had invested less in reproduction
September and December, and normally produced a than females who died. The ability to produce mul-
single clutch of between two and six eggs. The eggs tiple clutches appears to be related to the availability
incubated over the summer, and hatchlings appeared of food during spring.
in the field in January and February (Johnston, 1997).
Thus Peninsula dragons have a similar reproductive Acknmu1ed~ments.-This paper was partially funded
season to most other Australian dragon lizards (Brad- by an Australian Postgraduate Research Scholarship
shaw, 1981; James and Shine, 1985; Greer, 1989). held at Flinders University of South Australia. The
Males are generally larger than females among Broken Hill Proprietary Company Limited allowed ac-
Agamid lizards (Fitch, 1981). This pattern is thought cess to Iron Duke. The Carmody and Pumpa families
to be a result of sexual selection brought about by allowed access to Midgee Rocks and Siam Station, re-
male competition (Stamps, 1983; Johnston, 1997). In C. spectively. Craig Martins and Barbara Levings helped
fiolzl~ithe males and females matured at similar body collect lizards at Iron Duke. Mike Bull and Paula John-
sizes, and reach similar maximum body sizes. How- ston commented on the manuscript.
ever, mature males were generally larger than females
(Fig. 1). This difference is presumably a result of dif-
ferences between the sexes in growth pattern or sur-
vivorship. ALLISON,A. 1982. Distribution and ecology of New
The reproductive cycles of male and female C.fion~zi Guinea lizards. 111J. L. Gressitt (ed.), Biogeography
at Iron Duke coincided, so that the testes were pro- and Ecology of New Guinea, pp. 803-813. Dr W.
ducing sperm at the same time vitellogenesis occurred Junk, The Hague.
(Figs. 2 and 3). Mating presumably followed shortly BRADSHAW, S. D. 1981. Ecophysiology of Australian
afterwards in August and September. Thus sperm desert lizards: studies on the genus Amplzibolurus.
storage by females, which occurs in some lizards at 111 A. Keast (ed.), Ecological Biogeography of Aus-
similar latitudes (Cuellar, 1966; Smyth and Smith, tralia, pp. 1394-1434. Dr W. Junk, The Hague.
1968), may not occur in Peninsula dragons. No obvi- BRADSHAIV, S. D., H. SAINTGIIIONS,AND F. J. BRAD-
ous sexual activity was observed in the field after No- SHAW.1991. Patterns of breeding in two species of
vember (Johnston, 1997). However, adult males main- agamid lizards in the arid subtropical Pilbara re-
tained opaque epididymides throughout the year. In gion of Western Australia. Gen. Comp. Endocrinol.
so far as this indicates the presence of spermatozoa, it 82:407324.
appears that males store sperm in the epididymides CUELLAR, 0 . 1966. Oviducal anatomy and sperm stor-
even when the testes were not enlarged (i.e., actively age structures in lizards. J. Morphol. 119:7-20.
producing sperm). This may be advantageous for FITCH,H. S. 1981. Sexual size differences in reptiles.
males if females that produce a second clutch mate Misc. Pub. Mus. Nat. Hist. Kansas. 70:l-72.
between clutches. GREER, A. E. 1989. The Biology and Evolution of Aus-
698 SHORTER COMMUNICATIONS

tralian Lizards. Surrey Beatty, Norton, New South food habits, or locomotion of the animal. . . [and for
Wales. this reason it] may give better information on the ge-
HALL,W. P. 1970. Three probable cases of partheno- netic relationships than some habit- or habitat-corre-
genesis in lizards (Agamidae, Chamaeleontidae, lated characteristics" [our brackets]. Such a belief is
Gekkonidae). Experientia 26:1271-1273. here viewed with caution inasmuch as increasing
HOLSTOK,T. F. 1974. Revision of the Amphibolitrits de- knowledge about interspecific variation in snake taxa
cresii complex (Lacertilia: Agamidae) of South Aus- is showing just the opposite situation, wherein hemi-
tralia. Trans. Roy. Soc. S. Aus. 98:49-60. penial structures show highly complex transformation
INGER, R. F., AND GREENBERG, B. 1966. Annual repro- series within moderately diversified assemblages at
ductive patterns of lizards from a Bornean rain for- low taxonomic levels (e.g., Myers, 1974; Zaher, 1994).
est. Ecology 47:1007-1021. Similarly, there is no reason to assume that hemi-
JAMES, C. D., A N D SHINE,
R. 1985. The seasonal timing penial morphology in snakes is highly conservative
of reproduction: a tropical-temperate comparison intraspecifically, although potential intraspecific vari-
in Australian lizards. Oecologia 67:464-474. ation has been documented rarely (e.g., Inger and
JOHNSTON, G. R. 1982. The herpetofauna of the Mid- Marx, 1962; Keiser, 1974; McDowell, 1979; Cole and
dleback Range area, South Australia 1. An anno- Hardy, 1981). Here we describe the hemipenial vari-
tated checklist. Herpetofauna (Australia) 14:52-60. ation found in Triparnirgos coit~yressusand in the six
-. 1997. Behavioural Ecology of the Peninsula species comprising the genus Sipi~lophis. The termi-
Dragon Lizard, Cte1lopi:orus fiorlrli. Unpubl. Ph.D. nology used follows Zaher (in press).
Thesis, Flinders Unit: South Australia. In a phylogenetic analysis of the tribe Pseudoboini
KRAKZ,C. 1992. Results of breeding the peninsula (Zaher, 1994, 1996), Tripnr~urgoscort~pressus appeared
dragon, Cteilopkorus Fiorlni. Monitor 4:54-57. to be nested within Sipillopilis, as the sister-group of a
SMYTH, M. AND SMITH,M. J. 1968. Obligatory sperm clade composed of S. cerz>ir:us,S. yitlclwv, and S. Iciico-
storage in the skink Henliergis peroilii. Sclence 161: ccyi~nlus.For this reason, Triynrnirgos will be consid-
575-576. ered herein as a synonym of the genus Siyi~ioyll~s (in
SOKAL,R. R. AND ROHLF,F. J. 1981. Biometry. W. H. conformity with Article 24 of the International Code
Freeman, San Francisco. of Zoological Nomenclature). Additionally, Tri;~nilur-
STAMPS,J. A. 1983. Sfxual selection, sexual dimor- g o s c o n : p r e s ~ t s s h a r e swith all the species of Sipklopilis
phism and territoriality. I11 R. B. Huey, E. R. Pian- at least four unambiguous synapomorphies within the
ka, and T. W. Schoener (eds.), Lizard Ecology: pseudoboine radiation (Zaher, 1994). These features
Studies of a Model Organism, pp. 169-204. Har- include (plesiomorphic condition in parentheses): (1)
vard Univ. Press, Cambridge, Massachusetts. maxillan process of the prefrontal reduced or absent
(a well-detreloped process is present in all other pseu-
Accepted: 4 July 1999 doboines); (2) Meckelian canal closed from the ante-
rior region of the splenial to the tip of the dentary (all
other pseudoboines have an open canal); (3) postero-
ventral region of the nasal gland sharp-edged (this
/ourrral o Hrrprtolop, Vol. 33, KO. 4, pp. 698-i02, 1999 region is rounded in all the other pseudoboines); (4)
Copyridt 1999 so'hetyfor the Study of Amphibians and Reptiles 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 pseudo-
Intraspecific Variation of the Hemipenis boines). Zaher (1994) did not study S. ~~nroritzo?cli, and
in Siphlophis and Tripanurgos the only available specimen of S. leucocq~ilalusis in a
poor state of preservation. Recently, Prudente (unpubl.
HUSSAMZAHERAND ANA LUCIAC. PRLDENTE, De- data) reanalyzed Zaher's data, adding newr informa-
pnrtmeilto de Zoologin, UrlirursiiiniC de Srio Pnulo, Cnixn tion and including 5. ztoroiltzmc~i, and confirmed the
Postal 11461, Slio Pniilo, SP 05422-970, Brail, E-mail: phylogenetic position of T coit~yressuswithin the ge-
/:;n/ur@ib. us;,. br; alcgritdet~te@itsp.br nus Sipi~lopkis.
Intraspecific variation was found among specimens
The use of hemipenial characters in systematic stud- representing all the species of Sipi~loyl:is,each species
ies of snakes has increased substantially in the past showing the same two distinctly different types of
few years (see Savage, 1997 for a review). Such re- hemipenes. The first type, the "Y-shaped condition,"
newed interest is due mostly to the discovery of a clearly represents the primitive state among pseudo-
large number of potentially informative characters boines. The second type, which we designate the "T-
and to the improvement of techniques that facilitate shaped condition," is present also in Oxyrilopus clnth-
the preparation of uneverted organs (Pesantes, 1994). rntus, and it may represent a synapomorphy of a more
Hemipenial variation among species and higher cat- inclusive clade of pseudoboine snakes (Zaher, 1994).
egories have been acknowleged as an important However, it is not the purpose of the present paper to
source of characters of potential phylogenetic value, to discuss the possible optimization schemes for this
the extent that some workers have taken the more ex- character.
treme position of considering hemipenial features to A total of 27 fully everted and inflated hemipenes
be more reliable than other morphological character- were available for the following species (Table 1): S.
istics. This view have been expressed by Dowling cerainus (6) (Fig. I), S. pulcizer (7), S, longicnudatits (5),
(1967:138) who he argued that "the squamate hemi- S. leitcoctyhalus (2), S. itoroiltzoicli (2), S. con:pressits (5)
penis is a structure that (unlike teeth, skull, or scutel- (Fig. 2). All species hatre both "T- and Y-shaped" hem-
lation) has no obvious correlation with the ecology, ipenial types. We reject the possibility that these dif-
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Reproductive Biology of the Peninsula Dragon Lizard, Ctenophorus fionni
Greg Johnston
Journal of Herpetology, Vol. 33, No. 4. (Dec., 1999), pp. 694-698.
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Annual Reproductive Patterns of Lizards from a Bornean Rain Forest

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Obligatory Sperm Storage in the Skink Hemiergis peronii

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