Chromosomes of Australian Lygosomine Skinks (Lacertilia: Scincidae) II

Home , Eugongylus, Lampropholis, Lampropholis amicula, Leiolopisma, Lygosominae

Genrtic~a 83: 223-234, 1991, 0 I99 1 K~IIIW Acadwic Publishm. Prinled in the Netherlands. 223

Chromosomes of Australian lygosomine skinks (Lacertilia: Scincidae) II. The genus Lampropholis

S. C. Donnellan School of Biological Sciences, Macquarie University, New South Wales 2113, Australia Present adress: Evolutionary Biology Unit, South Australian Museum, Adelaide, South Australia 5000, Australia

Received 28 June 1990 Accepted in revised form 11 December 1990

Abstract

Standard and C-banded karyotypes of 101 specimens of the skink genus Lampropholis, including all but one of the 10 described species, were examined to assess the value of karyotypic data for resolving current systematic problems with Lampropholis. Diploid chromosome numbers were mostly 30 except for two forms with 28 chromosomes due to a reduction in the number of microchromosomes. The majority of interspecific chromosome changes were due to pericentric rearrangements. Apart from karyotypes referable to the described species, four distinct karyomorphs were found. Individuals possessing these karyomorphs could not be readily assigned to any of the described species.

Introduction The lack of resolution in systematics of Lampro- pho/is is due in part to the paucity of phylogenetically Greer (1979) divided the Australian members of the informative morphological characters. Karyotypic scincid lizard subfamily Lygosominae into three analyses have been found to be useful in reptiles for groups: the Egernia, Eugongylus and Sphenomorphus delineating species and for interpreting relationships groups. Lampropholis Fitzinger, a genus included in at higher taxonomic levels (Gorman, 1973; King & the Eugongylus group, has had a chequered systematic King, 1975; Bickham, 1983; King, 1983a). The present history. The genus typifies the ‘intractable small study was undertaken to assess the value of karyotypic brown skinks’ that have troubled systematists since data for investigating the systematics of Lampropholis. Boulenger’s time late last century. Greer (1974) ressur- ected Lampropholis from the synonymy of Leiolopisma Dumeril and Bibron to accomodate the type species L. Materials and methods guichenoti (Dumeril and Bibron) and three other species previously residing in Leiolopisma: L. challeng- Table 1 lists the 101 specimens of Lampropholis eri (Boulenger), L. delicata (De Vis) and L. mustelina studied, their geographic origins, and the chromo- (O’Shaughnessy). A further six species have been some analysis applied to each. Of the 10 described described recently (Greer & Kluge, 1980; Ingram & species, only L. tetradactyla was unavailable for study. Rawlinson, 1981), but documentation of species di- Specimens were deposited with the Australian, versity is still inadequate with several forms awaiting Queensland and South Australian Museums or re- taxonomic appraisal (Mather, 1990). tained by the original collectors as listed in Appendix The relationships of the species constituting Lam- 1. propholis remain unresolved. Greer and Kluge (1980) Karyotypes were prepared from heart and lung recognised the challengeri and delicata groups within fibroblasts cultured in Hams FlO medium supple- Lampropholis. Subesquently Ingram and Rawlinson mented with 20% foetal calf serum. Heart and lung (1981) were unable to assign with certainty L. caligula tissue was explanted onto coverslips in Leighton to either group. tubes. Outgrowths of fibroblasts derived from tissue 224

Table I. Specimens of Lampropholis examined. S= Standard Species Male Fe- Locality karyotype. C= C-band karyotype, Qld = Queensland, N.S.W. male =New South Wales. Superscripts l-7 indicate the presence of s cs c chromosomal variants as follows I: 2 individuals heterozygous for pericentric rearrangements of pair 8 (het.peri.8). 2: 1 individual L.guichenoti - I near Glare, S.A. heterozygous for heterochromatin addition (het. add.) to pair 9 1 I 2 2 Pymble, N.S.W. and I heterozygote for het.peri.8. 3: I heterozygote for Robertso- nian rearrangement of pair I or 2 and I heterozygote for het. add. L.mirabilis - I Cape Cleveland, Qld. to 9.41 I heterozygote for het. add. to 9 and IO. 5: I homozygote for peri. 8. I heterozygote for het. add. to 9. 6: I heterozygote for het. L.mustelina - I Greenwich, N.S.W. peri. 6. 7: I het.peri. 6. 1 Bellevue Hill, N.S.W. I I - Wentworth Falls. N.S.W. Species Male Fe- Locality I near Barry, N.S.W. male s cs c L.sp. A I Kuranda, Qld. 5 I - Charmillan Ck.. Qld. L.amicuia I - Mogill S.F., Qld. 1 - Rainbow Beach., Qld. L. sp. B I 3 Mt. Bartle Frere, Qld.

L. basiliscus I I 1 I Charmillan Ck., Qld. L. sp. c 2 - Connondale Range, Qld. 2 - Granite Ck., Qld. L. caligula - 1 I Barrington Tops, N.S.W. I - Bulburin S.F., Qld. 3 3 I I Crediton Ck., Qld. L.challengeri I I 1 I Randwick, N.S.W. I 1 Neutral Bay, N.S.W. L. sp. D I I I I Mt. Glorious, Qld. I Bellevue Hill, N.S.W. I 2 Mt. Nebo, Qld. I I - near Brinerville. N.S.W. I - Mt. Warning, N.S.W. I I Alstonville, N.S.W. I I I Mt. Glorious, Qld. pieces were harvested in situ on the coverslips. Three L.czechurai 2 I - Charmillan Ck., Qld. replicate cultures were initiated on each specimen. I - Millaa, Millaa, Qld. Cultures of most species grew adequately at 3O”C, L.delicata 2 2 Lauceston, Tasmania except those of L. challengeri, L. basiliscus and L. I Marble Range, S.A. czechurai which grew successfully only at 24°C. I near Port Lincoln, S.A. Meiotic preparations were made following the I I 25km N. Avenue, S.A. I I Eltham, Victoria technique of Gorman et al. (1967), excepting that I I - Bondi S.F., N.S.W. hypotonic treatment was for 25 minutes in 0.53% KCl. I 2’ near Nowra, N.S.W. Cells were harvested according to the procedure of I - Coogee, N.S.W. Donnellan (1990). Preparations were stained with 2 I 1: Pymble, N.S.W. I I - Neutral Bay, N.S.W. 10% Giemsa for two minutes. The C-banding method 2 3’ Watagan S.F., N.S.W. of Sumner (1972) was employed. I - Doyalson, N.S.W. 2 - Myall Lakes, N.S.W. I - Werrikimbe N.P., N.S.W. I Carrai S.F., N.S.W. Results 2 24 near Brinerville, N.S.W. I - Cambridge Plateau. N.S.W. 3 2 2 2’ Wiangarie SF., N.S.W. Diploid numbers were either 30 or 28 (Table 2) with 3 - Scarborough, Qld. nine macrochromosome pairs @airs one-nine) and Ih Connondale Range, Qld. either five or six pairs of microchromosomes. One I I I I’ Bunya Mtns., Qld. 2 - Granite Ck., Qld. individual of L. delicata had 29 chromosomes due to a 1 I Rockhampton. Qld. Robertsonian rearrangement among the macrochro- I Byfield, Qld. mosomes (see below). With the exception of this 2 - Mt. Morgan, Qld. individual the six largest pairs of chromosomes were invariant with the first five being metacentric and the Table 2. Summary of karyotypic variation in Lampropholis. Only distinct karyotypes could not be assigned readily to the chromosome pairs showing variation are presented. M: any described species and herein are assigned pro- metacentric, SM: submetacentric. A: acrocentric, T: telocentric. visionally to the taxa Lampropholis sp. A to D. Chromosome pair Lampropholis basiliscus Ingram and Rawlinson. 2n=30 Pairs six and seven were submetacentric, pair eight basiliscus acrocentric, and pair nine and the largest pair of an~icula microchromosomes were telocentric (Fig. 1). The caligula remaining pairs of microchromosomes were either challengeri czrchurai acrocentric or telocentric. The fourth pair of macro- delicata chromosomes and two pairs of microchromosomes pichenoti had prominent, centromeric C-bands (Fig. 5a). There tnirabilis mustelina were 'grey' telomeric C-bands on the long arm of one sp. A of the two largest macrochromosomes. sp. B sp. C Lampropholis amicula Ingram and Rawlinson. 2n=30 sp. D The standard karyotype of L. amicula differed from that of L. basiliscus in having an acrocentric pair seven, sixth submetacentric. Variation was confined to the metacentric pairs nine and ten, and in the largest remaining chromosomes (Table 2) and is detailed microchromosome pair being metacentric (Fig. 2a). along with the C-banding pattern of the whole There was insufficient material for C-banding. karyotype in the individual accounts which follow. For comparative purposes the karyotype of each Lampropholis caligula Ingram and Rawlinson. 2n=30 species is compared to that of L. basiliscus (Fig. l), The standard karyotype can be distinguished from which has the karyotype with the commonest mor- that of L. basilicus by the telocentric pair eight and by phology for the variable chromosome pairs. the largest microchromosome pair being metacentric In addition to the karyotypes of the described (Fig. 2b). C-bands (not illustrated), were minimal and species of Lampropholis, populations with four distinct confined to the centromeres. karyomorphs were found. Individuals with these Lampropholis challengeri ( Boulenger). 2n=30 The karyotype differed from that of L. basiliscus by having an acrocentric pair nine (Fig. 2c). Centromeric C-bands were minimal but telomeric C-bands were present on all members of the complement (Fig. 5b).

Lampropholis czechurai Ingram and Rawlinson. 2n=30 In three male specimens from two localities, pair seven was heterozygous for a pericentric rearrangement (Fig. 2d). Apart from pair seven, this species differed from L. basiliscus in having an acrocentric pair nine. There were prominent centromeric C-bands in this species (Fig. 5c). The first four pairs had telomeric C-bands. C-bands on the heteromorphic seven pair were confined to the centromere and appeared tq be equal in intensity. Fig. I. Standard karyotype of female Lampropholis basiliscus from Charpillan Ck.. Queensland (Qld) The bar indicates IOfim. 226

Fig. 2. Partial standard karyotype of some Lampropholis. (a) male L. amicula from Mogill State Forest (S.F.) Qld.; - (b) female L. ca/&y/a from Barrington Tops, New South Wales (N.S.W.): - (c) female L. challfngeri from Neutral Bay, N.S.W.; - (d) male L. czrchurai from Charmillan Ck.. Qld.; - (c) female L. Micara from Watagan S.F., N.S.W.; -(f) female L. guichenoti from Pymble, N.S.W.; - (g) female L. mirahilis from Cape Cleveland. Qld.; - (h) female L. mustrlina from Greenwich, N.S.W. The bar indicates IOpm. 227

Fis. 3. PwM. ~.at&.rd k.aqW.ypevaf ~wcw Lampraphdi.. c (32 m.ak L. c+, A ham. KurcW.k,QM..;- (b) A. few.dt L.. s+. B fvxv. bf.t.. &.P.% Fqplrp_, Qld.; (c) male L. sp. C from Bulburin S.F.. Qld. Note the heteromorphic pair of microchromosomes; -(d) female L. sp. C from Crediton Ck., Qld. Note absence of heteromorphism amongst the microchromosomes; - (e) male L. sp. D from Mt. Nebo, Qld. The bar indicates IOpm.

Lampropholis delicata (De Vis). 2n=28 centric. In general, centromeric C-banding was mi- The taxonomic status of L. delicata has recently been nimal and telomeric C-bands were present on the stabilised by Mather (1990) with the nomination of a larger macrochromosomes and distally on pair nine neotype. However, the species still lacks a description (Fig. 5d). adequate for identification purposes. Mather (1990) Ten individuals from the central part of the species’ also found three other electrophoretically distinct range were heterozygous for a variety of rearrange- forms which occur within the geographic range of L. ments (Table 1). One specimen was heterozygous for a delicata, which he named Forms B, C and D. These Robertsonian rearrangement involving one of the two forms had restricted ranges, all within Queensland. largest pairs of macrochromosomes (Fig. 4a). Two Thus of the four forms which Mather (1990) identified, individuals from different locations were heterozy- the specific expithet delicata is applied to the more gous for a pericentric rearrangement of pair six (Fig. geographically widespread of these species (for ex- 4b). Three specimens from two localities were hetero- ample Cogger, 1983). zygous for a pericentric rearrangement of pair eight The standard karyotype differed from that of L. (Fig. 4c), while a fourth from a third locality was basiliscus in the diploid number and in that pair eight homozygous for the variant chromosome. In both of was usually telocentric (Fig. 2e). The five pairs of the latter mentioned rearrangements C-bands were microchromosomes were either telocentric or acro- confined to the centromeres of these pairs. 228

One of these was also heterozygous for additional material on one of the larger microchromosomes. C- banding indicated that the additional material in both cases was totally heterochromatic.

4 Lampropholis guichenoti (Dumeril and Bibron). 2n=30 The standard karyotype of this species was distin- guishable from that of L. basifiscus by an acrocentric pair seven and by the metacentric nature of pair nine and the largest pair of microchromosomes (Fig. 2f). 6 7 8 9 This karyotype is generally similar to that reported by King (1973) for animals collected near Jenolan Caves, N.S.W.. Centromeric C-bands were prominent in this 6 7 species (Fig. 5e). Both specimens C-banded were heterozygous for proximal C-bands on the fifth pair and the intensity of this band varied between individuals. Prominent ‘grey’ C-bands were seen ter- minally on the long arm of the second largest pair of 6 7 8 9 macrochromosomes. The seventh pair had a fine d5 distal C-band on the long arm.

Fig. 4. Partial karyotypes of Lampropholis delicata showing Lampropholis micrabilis Ingram andRawlinson. 2n=30 structural heteromorphisms. (a) male from Watagan S.F., N.S.W. The karyotype differed from that of L. basilicus by heterozygous for a Robertsonian rearrangement of one of the two having a telocentric pair eight and a submetacentric larger pairs of macrochromosomes; - (b) female from Connondale Range. Qld. heterozygous for a pericentric rearrangement of pair pair nine (Fig 2g). There was insufficient material for six; - (c) male from near Nowra, N.S.W. heterozygous for a C-banding. pericentric rearrangement of pair eight; -(d) female from Wianga- rie S.F., N.S.W. with a short arm on one member of the usually telocentric pair nine. C-banding (lower pairs eight and nine) Lampropholis mustelina (O’Shaughnessy). 2n=30 demonstrates that the additoinal material is heterochromatic. The The karyotype of this species differed from that of L. bar indicates 10pm. basiliscus by the largest pair of microchromosomes being metacentric (Fig. 2h). Centromeric C-bands There were no apparent differences in bivalent were minimal except for the largest pair of chromo- morphology at first prophase of meiosis between the some pair nine. homozygotes and heterozygotes for the pericentric rearrangement of pair eight. The seven largest bi- Lampropholis sp. A. 2n=30 valents routinely showed more than one chiasma, The standard karyotype can be distinguished from while the remaining bivalents each displayed a single that of L. basiliscus by the telocentric pairs seven and chiasma. In the absence of significant centromeric eight and the submetacentric pair nine (Fig. 3a). The C-band markers it was not possible to determine if specimen C-banded had prominent centromeric C- there was any chiasma localization in either the bands and fine proximal bands either side of the homomorphic or heteromorphic bivalents. All stages centromere on two pairs of the larger macrochro- of meiosis observed in the homozygotes were present mosomes (Fig. 6a). There were prominent proximal in the heterozygotes. As both of the heterozygotes for C-bands on the telocentric pairs seven and eight. the pericentric rearrangement of pair six were females, meiotic analysis was precluded. Lampropholis sp. B. 2n=30 A further four specimens were heterozygous for the The standard karyotype differed from that of L. addition of a small short arm on pair nine (Fig. 4d). basiliscus in that the seventh and tenth pairs were 229

Fig. 5. C-band karyotypes otsome Lampropholis. (a)female L. basiliscus from Charmillan Ck., Qld.; - (b) male L. challengerifrom Randwick, N.S.W.: - (c) male L. czechurui from Charmillan Ck., Qld.; -(d) male L. delicata from Bondi S.F., N.S.W.; - (e) female L. guichenoti from Pymble, N.S.W. Note the interstitial C-band on pair five; - (f) male L. musrelina from Wentworth Falls, N.S.W. The bar indicates 10pm. Fig. 6. C-band karyotypes ofsome Lampropholis. (a) male L. sp. A from Charmillan Ck, Qld.; - (b)a male L. sp. C from Crediton Ck, Qld. Note the heterochromatic nature of larger element of the heteromorphic microchromosome pair; - (c) a female L. sp. C from Crediton Ck, Qld. Note the absence of a heteromorphic microchromosome pair; - (d) male L. sp. D from Mt. Glorious, Qld. The bar indicates 10pm. acrocentric while the eight was telocentric (Fig. 3b). confined to the centromere (Fig. 6b). This hetero- There was insufficient material for C-banding. morphism was absent from the only female examined (Fig. 6c), suggesting that it may be a sex chromosome Lampropholis sp. C. 2n=30 pair but more specimens must be sampled before this The standard karyotype can be distinguished from can be confirmed. There were prominent centromeric that of L. basiliscus by the telocentric pairs seven and C-bands on pairs six, seven, eight and on three pairs of eight and the submetacentric pair nine (Fig. 3c, d). Six microchromosomes. Fine proximal C-bands appear- male specimens from three widely separate localities in ed on either side of the centromere on the two largest central and southern Queensland displayed hetero- pairs of macrochromosomes. morphism in one of the two smaller pairs of microchromosomes. C-banding revealed that the larger Lampropholis sp. D. 2n=28 telocentric element was entirely heterochromatic, while The standard karyotype differed that of L. basiliscus the heterochromatin of its presumed partner was by the possession of telocentric pairs seven and eight 231 and a submetacentric pair nine (Fig. 3e). Centromeric additional heterochromatic blocks were observed, C-bands were prominent on all pairs except the four they occurred uniquely or as wide spread polymor- larger pairs of macrochromosomes. Three of the phisms providing a further source of intraspecific largest pairs of macrochromosomes had fine proximal karyotypic variation. C-bands either side of the centromere. There was a tine A further form of karyotype variation which ap- proximal C-band on the telocentric pair seven. A male pears widely but infrequently in lizards is the occurr- specimen was heterozygous for the addition of a small ence of heteromorphic sex chromosomes (reviewed in C-band to the distal end of a pair of microchromo- King, 1977). In L. czechurai and L. sp. C. the somes which also had a fine interstitial band on both observation of males from more than one locality with homologues (Fig. 6d). a heteromorphic chromosome pair suggests that sex chromosomes have differentiated morphologically in these forms. However, in the absence of adequate data Discussion for females this interpretation remains speculative although highly suggestive. In support, the male is the Karyotype evolution heterogametic sex in other skinks where heteromorphic sex chromosomes occur (Wright, 1973; Har- Various types of karyotypic changes were observed in dy, 1979). The involvement of two different chromo- Lampropholis. Most karyotypic differences between some pairs indicates two independent origins for species can be accounted for by pericentric rearrange- morphologically differentiated sex chromosomes in ments. The most frequent types of karyotype re- Lampropholis. The occurrence of heteromorphic sex arrangement observed in vertebrates involve changes chromosomes in lygosomine skinks will be addressed in chromosome number due to fusion-fission events or more fully in a forthcoming paper. changes in centromere position due to pericentric rearrangements. The former has certainly been the predominant mode in geckos (King, 1983a) and Karyotypic variation in Lampropholis delicata iguanids (German et al., 1967) while the latter is prevalent in varanids [King & King, 1975). Various structural rearrangements contribute to the In two forms of Lampropholis, L. delicata and L. sp. karyotypic variation observed in some populations of D the number of microchromosomes is reduced. L. delicata. The presence of individuals with unique Reduction in the number of microchromosome pairs rearrangements i.e. a Robertsonian rearrangement is commonly encountered in reptiles especially in the involving a macrochromosome pair, suggests that iguanid genus Anolis (Gorman, 1973), some Gehyra these are rare mutants rather than chromosomal (King, 1983a), chamaeleons and anguid lizards (Bick- polymorphisms. As sample sizes at most localities are ham, 1983). In some cases microchromosomes fuse small there must be uncertainty about the frequency of together to create new macrochromosomes or are such rearrangements. The wide spread geographic converted into macrochromosomes by addition of distributions of the telocentric form of chromosome heterochromatin. Neither of these mechanisms has pairs six and eight are indicative of pericentric re- operated in the case of Lampropholis. Instead in arrangement polymorphisms. Where sufficient num- Lampropholis a microchromosome pair has been lost bers of individuals have been studied structural hete- or translocated onto a larger macrochromosomal rozygosity has been observed in species from a pair. It is doubtful wether G-banding would help number of lizard families (King, 1983a), but there is no decide between these two possibilities because of the evidence for meiotic problems in individuals hetero- small size of the element involved. zygous for these types of rearrangements in lizards Heterochromatin has not played a major role in (King, 1983a; Porter & Sites, 1985). karyotypic differentiation between species oflampro- pholis being rather minimal in content and confined mainly to centromeric and telomeric areas. Where 232

Systematic implications of the status of the four karyomorphs will depend on an integration of ecological and morphological data The most significant taxonomic implication of the so that comparison with the appropriate species can be chromosome variation in Lampropholis, is the status made. of the four distinct chromosome forms (L.sp. A-D) Conversely karyotypic uniformity was observed in found among populations of L. delicata group ani- three species (L. delicata, L. challenger-i, L. mustelina) mals. Three of these taxa had previously been found to in which taxonomic problems exist on morphological be distinct on biochemical and morphological grounds grounds (G. Ingram, pers. comm.). For instance, L. (Mather, 1990). Thus forms B, C and D of Mather delicata is a widely distributed species showing varia- (1990) are equivalent to L. sp. C, D and A respectively. tion in body size, tail length, colouration and the It should be appreciated that each unique karyo- extent of sexual dichromatism. In this study, karyomorph may not constitute a new species as the types of animals from 25 localities, covering a major postulated chromosome rearrangements which dis- part of the range, showed overall uniformity. Karyo- tinguish these karyomorphs may not constitute a types of specimens from the far north of the range significant barrier to gene flow (Lande, 1979), although around the Atherton Tableland remain to be ex- this remains untested in the present case. However, amined. Taxonomic investigations of L. delicata, L. numerous examples of chromosome races being even- challengeri and L. mustelina should be tackled further tually recognised as species exist (e.g. King, 1982, by a biochemical technique such as allozyme electro- 1983b; Storr, 1979). phoresis which could establish a genetic framework L. delicata and L. sp. C appear to be distinct for the resolution of the taxonomic status of morpho- biological species. At Granite Ck., where two indi- logical variants. viduals of each karyomorph were collected in sym- Phylogenetic interpretations based on these data patry, differences in morphology of pair 7 and in must await more information. Too little is known of diploid number were maintained as well as the the karyotypes of appropriate outgroup genera of presence of probably heteromorphic sex chromo- lygosomine skinks to test the monophyly and the somes in L. sp. C only. The null hypothesis under test intergeneric relationships of Lampropholis. is that the four individuals from this locality were sampled at random from a single population which is therefore in Hardy-Weinberg equilibrium. The esti- Acknowledgements mated frequenceis of the submetacentric and telocentric forms of pair seven are p=OS and q=OS. This study was made possibleby the following people Likewise the estimated frequencies of the two diploid who generously provided specimens:S. Burgin, H. numbers is p=O.5 and q=O.5. The expected propor- Cogger, J. Covacevich, H. Ehmann, R. Green, A. tion of heterozygotes for either of these two characters Greer, M. Hutchinson, G. Johnston, M. Mahony, P. is 2pq=O.5. The probability of not observing an Mather, R. Sadlier, and G. Webb. The fauna authori- individual heterozygous for the rearrangement in pair ties of the States concerned are thanked for their 7 and/or with 29 chromosomes is [l-(2X0.5X0.5)]4‘*2 cooperation. S. Burgin, H. Ehmann, A. Greer, G. = 0.39% (Richardson et al., 1986). Thus a null Ingram, and P. Mather generously shared their un- hypothesis that the sample of four individuals was published information. P. Baverstock, R. Close, A. drawn from a single random breeding population is Greer, P. Johnston, and G. Sharman reviewed the not supported. An alternative hypothesis that the manuscript. Field work for this study was supported karyomorphs represent distinct biological species is in part by an Australian Museum Postgraduate also supported by the presence of fixed allelic differ- Award. ences at 20% of their loci (Mather, 1990). No further cases of sympatry occurred in the samples of L. sp. A, B, C, D and L. delicata group species studied here. Ultimately, a taxonomic decision 233

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(genus Lampropholis) from Queensland and New South Wales. czechurui (3): Charmillan Ck., Qld., 17”43’S., 145”3l’E.. AMS Mem. Qld. Mus. 20: 31 I-317. RI l4009-010. Millaa Millaa,Qld.. 17031’S.. 145”37’E., QM 341356. King, M., 1973. Karyotypic studies of some Australian Scincidae L. c/ci;catu (50): Launceston,Tasmania, 41”27’S., 147”lO’E.. AMS (Reptilia). Aust. J. Zool. 21: 21-32. RI 15281-4. Marble Range, S.A., 34”26’S., 135”29’E., SAMA King, M., 1977. The evolution of sex chromosomes in lizards. In: R297X5. near Port Lincoln, S.A., 34”42’S., 135”49’E., SAMA J. H. Calaby and C. H. Tyndale-Biscoe (eds), Reproduction and R29786-7. 25km N. Avenue, S.A., 36”45’S, 140”1‘3’E, SAMA evolution. Australian Academy of Science: Canberra: 55-60. R35818-9. Eltham, Victoria, 37”43‘S., 145”09’E.. AMS RI 15278, King, M., 1982. Karyotypic evolution in Gehyra (Gekkonidae: I 11609.. Bondi S.F., N.S.W., 37”08’S., 149”09’E., AMS RI 12353. Reptilia). Il. A new species from Alligator Rivers region in near Nowrd. N.S.W.. 35’03’S.. 150”20’E., AMS Rl02759-61. Northern Australia. Aust. J. Zool. 30: 93-101. Coogce, N.S.W., 33”55’S., 151”15’E., AMS R104796. Pymble, King. M.. 1983a. Karyotypic evolution in Gehyra (Gekkonidae: N.S.W., 33”45’S.. 151”08’E., LADEL61, SB65. one specimen lost. Reptilia). III. The Gehyra australis complex. Aust. J. Zool. 31: Neutral Bay, N.S.W., 33”5l’S., 151”12’E., LADEL62. Watagan 723-74 I. SF., N.S.W.. 33”15’S., 151°20’E.. AMS R97827, 10478789, King, M.. 1983b. The Gehyra australis species complex (Sauria: LADEL27. Doyalson. N.S.W.. 33”13’S., 151”30’E., AMSR104203. Gekkonidae). Amphibia - Reptilia 41: 147-169. Myall Lakes, N.S.W., 32”25’S., 152”30’E., LADEL7, 15. Werri- King, M. & King. D., 1975. Chromosomal evolution in the lizard kimbe N.P.. N.S.W., 31”09’S.. 152’14’E., AMS RI1 1470. Carrai genus Varanus (Reptilia). Aust. J. Biol. Sci. 28: 89-108. S.F..N.S.W.. 3l”Ol’S.. 152”20’E.,AMSR112322. near Brinerville, Lande. R., 1979. Effective deme sizes during long-term evolution N.S.W., 30”28’S., 152”33’E., AMS R102763, 111787. Cambridge estimated from rates of chromosomal rearrangement. Evolution Plateau, N.S.W., 28”45’S., 152”45’E., AMS RII 1839. Wiangarie 33: 234-25 I. SF., N.S.W., 29”08’S., 152”07’E.. AMS RI 11472, 785-6, 474, Mathcr, P.. 1990. Electrophoreticand morphologicalcomparisons 115298. Scarborough, Qld., 27”12’S., 153”07’E., AMS Rl11837, of Lampropholis delicata (Lacertilia: Scincidae) populations 478, 480. Connondale Range, Qld., 26”33’S.. 152”35’E.. AMS from Eastern Australia, and a resolution ofthe taxonomic status Rl04196. Bunya Mtns.. Qld.,26”55’S., 151’37’E.. AMS Rll4097- of this species. Aust. J. Zoo]. 37: 561-74. 8. Granite Ck., Qld., 24”36’S., I5 1”4O’E., 545295-6. Rockhampton. Porter. C. A. & Sites, J. W., 1985. Normal disjunction in Robert- Qld., 23”22’S., 150”32’E.. AMS RI 11833. Byfield, Qld.. 22”5l’S.. sonian heterozygotes from a highly polymorphic lizard popula- 150”39’E., HE2373. Mt. Morgan, Qld.. 21”28’S., 150”22’E., AMS tion. Cytogenetics and Cell Biology 39: 250-257. RI 15268-9. L. ,pichenoti (4): near Glare, S.A., 33”5O’S., 138”37’E., Richardson. B. J., Baverstock. P. R.&Adams, M.. 1986. Allozyme HE3334. Pymble, N.S.W., 33”45’S., 151°08’E., AMS Rl11574-5. Electrophoresis. Academic Press, Sydney. SB64. 1.. mirabilis (I): Cape Cleveland, Qld., 19”l I’S, 147°01’E.. 234

AMS Rll1828. L. mustelina (4): Greenwich, N.S.W., 33”5O’S., 17’24’S., 145”49’E., QM J4@333,40036-9. L. sp. C (8): Connondale 151”l l’E., AMS R96788. Bellevue Hill, N.S.W., 33”53’S., Range. Qld.. 26’33’S., 152”35’E., AMS Rl02778-9. Granite Ck., 151”15’E., SB220. Wentworth Falls, N.S.W., 33O43’S., 150”22’E., Qld., 24”36’S., 151”40’E., QM 545297-8. Bulburin S.F., Qld., AMS R111784. near Barry, N.S.W., 31”35’S., 151”25’E., AMS 24”3O’S., 151”30’E., HE2373. Crediton Ck., Qld., 21”12’S., R102748. L. sp. A (6): Kuranda, Qld., 16”49’S., 145”38’E., QM 148”32’E., AMS R111825,829-30,834. L. sp. D (5): Mt. Glorious, 541360. Charmillan Ck., Qld., 17”43’S., 145”31’E., QM 541361-2, Qld., 27”2O’S., 152”45’E., AMS R111835,37. Mt. Nebo, Qld., AMS R94537, 538, 94545. L. sp. B (4): Mt. Bartle Frere, Qld., 27”24’S., 152”45’E., AMS R97875,77.