Parthenogenetic Lizards Research Articles

Instability of (GATA)n microsatellite loci in the parthenogenetic Caucasian rock lizard Darevskia unisexualis (Lacertidae).

Mini- and microsatellites, comprising tandemly repeated short nucleotide sequences, are abundant dispersed repetitive elements that are ubiquitous in eukaryotic genomes. In humans and other bisexual species hypervariable mini- and microsatellite loci provide highly informative systems for monitoring of germline and somatic instability. However, little is known about the mechanisms by which these loci mutate in species that lack effective genetic recombination. Here, multilocus DNA fingerprinting was used to study M13 minisatellite and (GATA)n microsatellite instability in the parthenogenetic Caucasian rock lizard Darevskia unisexualis (Lacertidae). DNA fingerprinting of 25 parthenogenetic families, from six isolated populations in Armenia (comprising a total of 84 siblings), using the oligonucleotide (GATA)4 as a hybridization probe, revealed mutant fingerprinting phenotypes in 13 siblings that differed from their mothers in several restriction DNA fragments. In three families (8 siblings), the mutations were present in the germline. Moreover, the mutant fingerprint phenotypes detected in siblings were also present in population DNA samples. No intrafamily variations in DNA fingerprint patterns were observed with the M13 minisatellite probe. Estimates of the mutation rate for (GATA)n microsatellite loci in D. unisexualis showed that it was as high as that seen in some bisexual species, reaching 15% per sibling or 0.95% per microsatellite band. Furthermore, in one case, a somatic (GATA)n microsatellite mutation was observed in an adult lizard. These findings directly demonstrate that mutations in (GATA)n microsatellite loci comprise an important source of genetic variation in parthenogenetic populations of D. unisexualis.

Genetic Homogeneity between Populations of Aspidoscelis rodecki, a Parthenogenetic Lizard from the Yucatán Peninsula

Abstract We undertook skin-grafting between populations to determine whether Aspidoscelis rodecki originated from single, or multiple, parthenogenetically capable hybrids. We transplanted 292 skin grafts within and between the two most geographically distant populations, considering only grafts in animals surviving more than 45 days. Two hundred fifteen grafts were analyzed. Histocompatibility within (100%) and between (97.9%) populations suggests that A. rodecki was derived from a single, parthenogenetically capable, hybrid.

The Whiptail Lizard Reconsidered

Harry Collins and Trevor Pinch's introductory text, The Golem: What Everyone Should Know About Science (1993), includes a controversy about the significance of pseudosexual behavior in the parthenogenetic whiptail lizard. Collins and Pinch, basing their account on the work of Greg Myers (1990), claim that “in this area of biology, experiments are seldom possible” and that the debate has “battled to an honorable draw.” I argue that a closer look at the publications of the scientists involved shows that, at least by the late 1980s, it was widely accepted that pseudosexual behavior is important for reproduction in these lizards. Moreover, a variety of experiments, as well as laboratory and field observations, proved decisive in this acceptance.

Hyperunstable (TCT/TCC) n Microsatellite Loci in Parthenogenetic Lizards Darevskia unisexualis (Lacertidae)

Using multilocus DNA fingerprinting, we have examined variability of (TCT) n microsatellite and M13 minisatellite DNA repeats in populations, families, and tissues of parthenogenetic Caucasian rock lizards Darevskia unisexualis (Lacertidae). It has been shown for the first time that population and family DNA samples of D. unisexualis (75 samples in total) have individually specific DNA fingerprinting patterns of (TCT) n fragments. Analysis of inheritance of (TCT) n microsatellites in 46 first-generation progeny in 17 parthenogenetic D. unisexualis families revealed their extremely high instability. Mutant TCT fingerprint phenotypes were found in virtually each animal of the progeny. Moreover, varying fragments in the progeny and their original variants in the mothers were shown to simultaneously contain (TCT) n and (TCC) n polypyrimidine clusters. At the same time, no variability of (TCT) n fragments has been detected in the tissues and organs of mature parthenogenetic lizards and in the analogous tissues of the two-week-old progeny of this year. This suggests the absence of somatic mosaicism and methylation of the corresponding loci in the samples. Along with the hyperinstability of (TCT/TCC) n polypyrimidine clusters, we have shown that the population and family DNA fingerprinting patterns of M13 minisatellites were invariable and monomorphic in the same DNA samples of D. unisexualis.Our results indicate that mutations at loci containing polypyrimidine microsatellites significantly contribute to the total genomic variability of parthenogenetic lizards D. unisexualis.

Variation of mini- and microsatellite DNA repeats in parthenogenetic lizard Darevskia armeniaca as revealed by DNA fingerprinting analysis

Population and family samples of two morphological forms (mutant and normal with respect to dorsal color) of pathogenetic lizard Darevskia armeniaca were examined by means of DNA fingerprinting using M13 mini- and (GATA)n and (TCC)n microsatellite DNA markers. The morphological forms examined were characterized by clonally inherited, species-specific patterns of the DNA markers, which were different from the species-specific DNA fingerprints of the other parthenogenetic species of the genus Darevskia (D. dahli. D. unisexualis, and D. rostombekovi). The mean index of similarity (S) obtained for a sample of 36 individuals from three isolated populations using three types of DNA markers was 0.966. This was similar to the variability level observed in D. dahli (0.962) (P > 0.05), but higher than that in D. unisexualis (0.950) (P < 0.05) and D. rostombekovi (0.875) (P < 0.01). Inheritance of M13 minisatellite and (TCC)n microsatellite DNA markers in the F1 offspring of parthenogenetic lizards was examined. It was shown that variability and clonal diversity of the fingerprint phenotypes observed in the populations and families of D. armeniaca could be at least partly explained by RFLP mutations in microsatellite repeats.

Quantitative Assessment of Gene Diversity and Between-Population Differentiation of Parthenogenetic Lizard of the Genus DarevskiaUsing Mini- and Microsatellite DNA Markers

Methods of estimating within- and between-population gene diversity in parthenogenetic species using mini- and microsatellite DNA markers and modified Wright's FST statistic are presented with special reference to model populations of lizards of the genus Darevskia(D. dahli, D. armeniaca, D. unisexualis). We used DNA fingerprinting data for several populations of these species examined earlier. The effects of variation in M13 minisatellite, (GACA)n- and (TCC)n-microsatellite loci on the formation of within-population gene diversity in parthenogenetic species D. dahli and D. armeniaca were shown to be different. The equality of the realized gene diversity Hand its maximum possible value Hmax in two populations of D. dahli (Hmax = 0.032, H = 0.031, P≪ 0.0431; Hmax = 0.024, H = 0.027, P = 0.09) and D. armeniaca (Hmax = 0.05, H = 0.053, P = 0.03; Hmax= 0.054, H = 0.055, P= 0.02) suggests that variation in (GACA)n loci substantially contributes to the maintenance of within-population genetic diversity. Analysis of between-population genetic diversity using loci M13, (GACA)n, and (TCC)n showed differentiation of D. dahli populations from northeastern and northwestern Armenia (FST = 0.0272, P = 3 × 10–13) and genetic homogeneity of the Armenian and introduced to the Ukraine populations of D. armeniaca characteristic of one clone (FST = 0, P = 1).

Natural Hybridization Between the Teiid Lizards Cnemidophorus tesselatus (Parthenogenetic) and C. tigris marmoratus (Bisexual): Assessment of Evolutionary Alternatives

Annual hybridization is taking place between representatives of the parthenogenetic lizard Cnemidophorus tesselatus (2n = 46, 47) and males of the bisexual species C. tigris marmoratus (2n = 46) in desert grassland habitats at Arroyo del Macho, Chaves County, New Mexico. This raises the question of whether a new triploid parthenogenetic species may be originating as a consequence of this activity. Hybrids were collected in each of four years (1996–1999), and 20 of 21 hybrids collected (12 males and 8 females) were available for study. Although a triploid parthenogenetic species (Cnemidophorus exsanguis 3n = 69) and a diploid bisexual species (C. inornatus 2n = 46) were also found at the hybridization site, the genealogy of the hybrids was determined unequivocally with karyotypic and electrophoretic evidence (34 loci tested). The specimens examined electrophoretically included an adult female and one of her laboratory-reared daughters, which demonstrated for the first time clonal inheritance in C. tesselatus pattern class E.The population of C. tesselatus at Arroyo del Macho is characterized by two karyotypic cytotypes. The ancestral one (2n = 46) occurs at about half the frequency of the derived cytotype (2n = 47), which apparently was produced by centric fission of the ancestral X-chromosome from C. tigris In contrast, the occurrence of the two cytotypes was reversed and strongly asymmetrical in the hybrids; only one of nine hybrids possessed the fissioned X-chromosome. This individual was significantly different in 12 meristic characters from the sample of hybrids with intact X-chromosomes. Predictably, principal components scores for this individual fell outside the 95% confidence ellipse of scores of the other eight hybrids that were karyotyped. The skewed ratio and multiple phenotypic differences suggest that hybrids inheriting a fissioned X-chromosome might be at a selective disadvantage compared to hybrids with intact X-chromosomes.All 20 hybrids closely resemble C. tesselatus in most color pattern features. However, these hybrids, like C. tigris marmoratus lack lateral stripes. Because the population of C. tesselatus at Arroyo del Macho has lateral stripes (or their remnants), hybrids can be readily distinguished from C. tesselatus by this color pattern feature. Compared to the two parental species, hybrids had a significantly lower mean number of scales around midbody, but hybrids resembled either C. tesselatus or C. tigris marmoratus in other univariate meristic characters. This mosaic pattern of resemblance was simplified to a three-dimensional depiction of variation using principal components analysis. Each of two principal components expressed the resemblance of hybrids to one of the two parental species. A third component reflected the difference between hybrids and both parental species. A canonical variate analysis of meristic characters demonstrated the multivariate distinctiveness of each group—hybrids, C. tesselatus and C. tigris marmoratus However, based on Mahalanobis D2 distances, the closest morphological resemblance among hybrids and parental species was between hybrids and the maternal species, C. tesselatusNine additional museum specimens, suspected of being C. tesselatus × C. tigris marmoratus hybrids, were identified, as such, by a canonical variate analysis using our samples of C. tesselatus, C. tigris marmoratus and hybrids from Arroyo del Macho as a priori groups. These nine individuals document hybridizations between C. tesselatus and C. tigris marmoratus at two additional localities in Chaves County, New Mexico, two localities in Sierra County, New Mexico, and a cluster of sites near Presidio, Presidio County, Texas. Previously, several of these hybrids had been misidentified as male C. tesselatusThe reproductive systems of female and male hybrids were compared histologically to those of C. tesselatus and C. tigris marmoratus respectively. Sexually mature and reproductive adults of C. tesselatu

Ecology and Behavior of Lizards of the Parthenogenetic Cnemidophorus laredoensis Complex and Their Gonochoristic Relative Cnemidophorus gularis: Implications for Coexistence

The Cnemidophorus laredoensis complex consists of two, all-female parthenogenetic lizard species, designated LAR-A and LAR-B, that commonly coexist with their gonochoristic (= bisexual) relative Cnemidophorus gularis in southern Texas. The ecology and behavior of these Cnemidophorus lizards was studied in a Texas state park where LAR-A, LAR-B, and C. gularis coexist. I used time of capture records to estimate daily activity periods and used focal animal observations of free-ranging individuals to determine what proportion of time lizards spent using different microhabitats, occupying different exposures (full sun, partial sun, shade), and performing different behaviors during the May-June breeding seasons of 1994- 1996. There was little difference among the three species in daily activity period, microhabitat use, or exposure occupancy; added to previously published data on diet similarity, these results suggest that com- petition between the two parthenogens and C. gularis potentially is severe. The greatest behavioral differ- ences were between male C. gularis and female C. gularis, LAR-A, and LAR-B: male C. gularis spent a much greater percentage of their time interacting with other lizards and moved significantly farther per unit time than did females. These results reflect the large proportion of time male C. gularis devoted to trailing and courting female C. gularis as well as parthenogenetic females, especially LAR-A. The only major difference between C. gularis and the two parthenogens was that the latter spent a greater proportion of their time searching for prey. Despite this, LAR-A and LAR-B did not achieve a greater prey capture rate, nor a greater success at locating the preferred prey of termites than did C. gularis, suggesting the two parthenogens are less efficient foragers than their gonochoristic relative. This may account for the reason why the two par- thenogens are not competitively excluding C. gularis from the park despite advantage of a much greater population size. Overall, data from this and other studies lend support to the idea that parthenogenetic Cnemidophorus are ecologically and behaviorally inferior to their gonochoristic congeners and should, therefore, be able to coexist with them only in habitats where selective forces are relaxed.

Do Timing and Pattern of Myogenesis Correlate with Life History Mode in Anurans?

cozumela. They accepted the idea that parthenogens derived from separate hybridizations should be recognized as different species, but they did not consider it as an exclusive criterion. They agreed with Echelle (1990) that postformational mutational events of sufficient magnitude can serve as points of origin for new parthenogenetic species (Taylor and Cooley, 1995a, b). We agree with this position. The karyotypic and morphological differences and the geographical barrier between C. cozumela and C. maslini appear to be sufficient elements to give species recognition to C. maslini, thereby reflecting the substantial evolutionary divergence between these two groups of parthenogenetic lizards.

Origin and Evolution of the Parthenogenetic Lizards, Cnemidophorus maslini and C. cozumela

Origin and Evolution of the Parthenogenetic Lizards, Cnemidophorus maslini and C. cozumela

Genetic homogeneity based on skin histocompatibility and the evolution and systematics of parthenogenetic Cnemidophorus laredoensis (Sauria: Teiidae)

The parthenogenetic whiptail lizard Cnemidophorus laredoensis of Texas and México was derived from one or more hybrids between the gonochoristic species Cnemidophorus gularis and Cnemidophorus sexlineatus. We analyzed clonal structure in C. laredoensis on the basis of histocompatibility reactions to reciprocal skin grafting. Individuals of C. laredoensis rejected skin grafts from both parental species and from a hybrid between C. gularis and an undescribed parthenogenetic species. One lizard initially identified as C. laredoensis was found to be a triploid C. laredoensis × C. gularis hybrid, based on its histocompatibility reactions. The hybrid accepted grafts from individuals of C. laredoensis; however, C. laredoensis rejected grafts from this and a second hybrid. Seventeen individuals of C. laredoensis from six sites located throughout the range of the species in Texas were fully histocompatible, based on a large number of reciprocal skin grafts. Only three individuals of C. laredoensis from three sites donated grafts to other individuals of the species, but they were unable to host all or most of the reciprocal grafts. The skin histo compatibility among 17 individuals of C. laredoensis from six widely separated geographic sites points to a high level of genetic homogeneity in the species due to clonal descent from one parthenogenetic C. gularis × C. sexlineatus hybrid. Unidirectional rejection of skin grafts by three lizards could be a result of postformational mutations within this lineage, though this suggestion is open to question. The name C. laredoensis should be restricted to populations in Texas and México that are diagnostically identical with the parthenogenetic species described from two sites in Laredo, Webb County, Texas, in 1973.

Reproductive Characteristics and Body Size in the Parthenogenetic Teiid Lizard Cnemidophorus tesselatus: Comparison of Sympatric Color Pattern Classes C and E in De Baca County, New Mexico

Cnemidophorus tesselatus is an allodiploid, parthenogenetic lizard comprised of a number of distinctive tokogenetic arrays (Frost and Hillis, 1990) presently identified by four color pattern class descriptors: C, Colorado D, New Mexico D, and E (Taylor et al., 1996; Walker et al., 1997). The macrogeographic relationships of these pattern classes were outlined in Zweifel's (1965) pioneering study of geographic variation in Cnemidophorus tesselatus. Pattern class C was known to exist isolated from the other pattern classes in parts of southeastern Colorado, northwestern Oklahoma, and northwestern Texas. Sympatry between pattern classes C and D was known to occur only in the vicinities of the historic townsite of Higbee, Otero County, Colorado, and Conchas Lake, San Miguel County, New Mexico. Pattern class E occurred to the south, with specimens from Fort Sumner, De Baca County, New Mexico, approximately 105 km south of the Conchas Lake locality, documenting the closest proximity of pattern class E to pattern classes C and D.

Cnemidophorus neomexicanus-Cnemidophorus perplexus Nomenclatural Problem (Sauria: Teiidae) and Its Resolution

For the past 44 yr, populations of the parthenogenetic New Mexico whiptail lizard have been allocated to either Cnemidophorus perplexus Baird and Girard 1852 or Cnemidophorus neomexicanus Lowe and Zweifel 1952. The latter name has been in general use for the past 30 yr based on an opinion that the lectotype of C. perplexus, United States National Museum (USNM) 3060, was a triploid hybrid derived from insemination of a normally parthenogenetic individual of C. neomexicanus by a male of C. inornatus. Questions concerning the genealogy of the lectotype have recently resurfaced. Therefore, we used canonical variate (CVA) and principal components analyses (PCA) of nine meristic characters and snout-vent length to test the hypothesis that the lectotype of C. perplexus is a hybrid. Because C. neomexicanus and C. inornatus are sympatric at the type locality of the former, this also provided an opportunity to verify that the type of C. neomexicanus is not a hybrid. Our reference material included two sets of geographically paired samples of the presumptive parental species, C. neomexicanus (n = 77) and C. inornatus (n = 105) from the vicinity of the restricted type locality of C. perplexus. Of 23 specimens presumed from morphological features to be C. neomexicanus x C. inornatus hybrids, 13 that were syntopic with one of the reference sets were used as our hybrid reference group. The type specimens of C. perplexus and C. neomexicanus were included in the CVA as unknowns for assignment to the hybrid group or taxon with which each shared the greatest multivariate similarity. The CVA assigned the holotype of C. neomexicanus to C. neomexicanus (Set 1: P = 0.642; Set 2: P = 0.358), and the lectotype of C. perplexus was assigned to C. neomexicanus as well (Set 2: P = 0.789; Set 1: P = 0.211). Projections of scores for both types in a principal components analysis of all specimens supported both assignments. In addition, the color pattern of the lectotype resembles C. neomexicanus and none of the 23 hybrids, and meristic scores of the lectotype are within the range limits of our reference samples of C. neomexicanus. Our analyses indicate that the lectotype of C. perplexus was misidentified as a hybrid by previous workers. However, there are two compelling reasons why we shall petition the ICZN to suppress perplexus as an available name for a species of Cnemidophorus: (1) ambiguity concerning the type locality; and (2) controversy as to whether the name C. perplexus should be associated with specimen USNM 3060. Approval of the petition will bring the issue to closure and extend the 30yr use of the name C. neomexicanus for the New Mexico whiptail.

Rapid Screening of DNA Diversity Using Dot-Blot Technology and Allele-Specific Oligonucleotides: Maternity of Hybrids and Unisexual Clones of Hybrid Origin (Lizards,Cnemidophorus)

Allele-specific oligonucleotide probes, together with dot-blot methods, can provide rapid and inexpensive screening of DNA types in large samples of organisms. Here we demonstrate their use in: (1) determining types of mitochondrial DNA in hundreds of lizards from a dynamic hybrid zone; (2) discovering intraspecific geographic variation in genes; and (3) determining and verifying the maternal ancestry of unisexual, parthenogenetic lizards in clones of hybrid origin. These methods are broadly applicable in research involving rapid screening of DNA types in large samples of specimens for any gene with sequence data from which to design specific probes.

Patterns of Meristic Variation among Parthenogenetic Teiid Lizards (Genus Cnemidophorus) of the Yucatan Peninsula and Their Progenitor Species, C. angusticeps and C. deppei

Patterns of Meristic Variation among Parthenogenetic Teiid Lizards (Genus Cnemidophorus) of the Yucatan Peninsula and Their Progenitor Species, C. angusticeps and C. deppei

The bisexual brain: sex behavior differences and sex differences in parthenogenetic and sexual lizards

The parthenogenetic lizard Cnemidophorus uniparens alternates in the display of male-like and female-like sexual behavior, providing a unique opportunity for determining the neuronal circuits subserving gender-typical sexual behavior within a single sex. Here we report a 6-fold greater [ 14C]2-fluoro-2-deoxyglucose uptake in the medial preoptic area of C. uniparens displaying male-like behavior in comparison with C. uniparens displaying female-like receptivity. The ventromedial nucleus of the hypothalamus showed greater 2DG accumulation in receptive C. uniparens than in courting C. uniparens. When a related sexual species ( C. inornatus) was compared to the unisexual species, the anterior hypothalamus in C. inornatus males exhibited significantly greater activity.

Hybrid Cnemidophorus (Sauria: Teiidae) in Ninemile Valley of the Purgatoire River, Colorado

Canonical variate analysis (CVA) of color pattern and scutellation characters in lizards collected in Ninemile Valley of the Purgatoire River at Higbee, Otero Co., Colorado graphically depicted two forms of parthenogenetic Cnemidophorus tesselatus (diploid color pattern class C and triploid color pattern class B), gonochoristic C. sexlineatus, and hybrid C. tesselatus x C. sexlineatus as four morphologically distinctive groups. A newly obtained male specimen with a C. tesselatus-like color pattern entered as an unknown was classified to the hybrid group in the CVA. It represented the fourth tetraploid hybrid C. tesselatus B(3n) x C. sexlineatus collected in the valley. A female specimen was also classified to the hybrid group by the CVA. A triploid karyotype (3n = 69) and color pattern identified this female as a putative C. tesselatus C(2n) x C. sexlineatus hybrid. Whereas male hybrids are presumably sterile and biological novelties at most, the possibility remains that a female hybrid could be the founder of a new parthenogenetic lineage. Zweifel (1965) informally described six color pattern classes of parthenogenetic lizards (A, B, C, D, E, F) in the Cnemidophorus tesselatus complex (family Teiidae) of which four (A-D) were reported from Colorado. Wright and Lowe (1967a) found that pattern classes A and B were triploid hybrid derivatives and pattern classes CF were diploid hybrid derivatives. Analyses of morphology (Walker et al., 1990) and karyology (Cordes, 1991) identified three of Zweifel's pattern classes of C. tesselatus in sympatry in Ninemile Valley of the Purgatoire River at Higbee, Otero Co., Colorado rather than two as indicated by Zweifel (1965), Densmore et al. (1989), and Wright (1993). These included diploid C. tesselatus C and D and triploid B, not triploid pattern class C as indicated by Parker and Selander (1976), Densmore et al. (1989), Wright (1993), and Leuck (1993). We found that the Ninemile Valley triploid population was morphologically indistinguishable from the pattern class B population described by Zweifel (1965) from Pueblo Co., Colorado. In samples collected in the valley since 1960 by various workers, diploid C. tesselatus outnumbered triploid C. tesselatus by a combined ratio of over 7:1. Moderate numbers of the gonochoristic species C. sexlineatus, the paternal parent of triploid C. tesselatus B (Wright and Lowe, 1967a; Parker and Selander, 1976), were also found in many parts of the area sampled. Four hybrid male C. tesselatus x C. sexlineatus were included in these collections. Three of these were described, and their probable parental forms identified as triploid C. tesselatus B x C. sexlineatus, by Walker et al. (1990). In the present study we describe the most recently collected hybrid male Cnemidophorus, infer its putative parentage on the basis of critical morphological characters, and evaluate the possibility of a hybrid origin for a female specimen displaying an ambiguous combination of classification characters. MATERIALS AND METHODS-Characters of scutellation that were quantitatively analyzed included: number of granules around midbody (GAB), number of granules from occiput to rump (OR), sum of left and right femoral pores (FP), number of subdigital lamellae on the longest toe of the left pes (SDL), sum of left and right circumorbital scales (COS), sum of This content downloaded from 207.46.13.57 on Mon, 08 Aug 2016 05:24:23 UTC All use subject to http://about.jstor.org/terms 236 The Southwestern Naturalist vol. 39, no. 3

Comparative ecology of sexual and asexual gecko species (Lepidodactylus) in French Polynesia

The asexual geckoLepidodactylus lugubris, its sexual congenerL. sp. (Takapoto) and hybrids between the two species inhabit the atoll of Takapoto, providing a natural experiment for studying co-existence and interactions between asexual and sexual populations. The range of the sexual species is confined to one section of the lagoon beach and the trees and buildings which abut it, whereas the asexual is distributed across the whole atoll and occupies many habitats. Behavioural experiments revealed no asymmetry in levels of aggression between the two species, suggesting that the confinement ofL. sp. (Takapoto) to the lagoon beach is not due to agonistic interactions. Ecological differences among the constituent clones of the asexual species exist but cannot completely account for the broader habitat use of the asexual. Within a single habitat, one clone ofL. lugubris consumes a wider range of prey items than its sexual relative. Other studies have found that the asexuals are extremely heterozygous relative to the sexuals; we hypothesize that their broad ecological tolerance may be attributable to heterosis. The co-existence of the sexual and parthenogenetic lizards on this small island seems to be stable and may be facilitated by the specialization of the sexual taxon to beach habitats.

Diet of Diploid and Triploid Populations of Parthenogenetic Whiptail Lizards of the Cnemidophorus tesselatus Complex (Teiidae) in Southeastern Colorado

nithol. Union, Washington, D.C. AVERY, M. L., AND C. VAN RIPER III. 1986. Bird community survey at Pinnacles National Monument. U.S. Natl. Park Serv., Coop. Natl. Park Resourc. Stud. Unit Tech. Rep. 24, Davis, California, 83 pp. BALTOSSER, W. H. 1989. Costa's hummingbird: its distribution and status. West. Birds, 20:41-62. BOLTEN, A. B., P. FEINSINGER, H. G. BAKER, AND I. BAKER. 1979. On the calculation of sugar concentration in flower nectar. Oecologia, 41:301-304. CARPENTER, F. L. 1983. Pollination energetics in avian communities: simple concepts and complex realities. Pp. 215-234, in Handbook of experimental pollination ecology (C. E. Jones and R. J. Riddle, eds.). Van Nostrand Reinhold Co., New York. HIXON, M. A., AND F. L. CARPENTER. 1988. Distinguishing energy maximizers from time minimizers: a comparative study of two hummingbird species. Amer. Zool., 28:913-925. JOHNSGARD, P. A. 1983. The hummingbirds of North America. Smithsonian Institution Press, Washington, D.C. LEVALLEY, R., AND K. F. CAMPBELL. 1984. The spring migration: March 1-May 31, 1984, middle Pacific Coast region. Amer. Birds, 38:952-957. SPIRA, T. P. 1980. Floral parameters, breeding system and pollinator type in Trichostema (Labiatae). Amer. J. Bot., 67:278-284. STILES, F. G. 1971. On the field identification of California hummingbirds. Calif. Birds, 2:41-54. 1972. Food supply and the annual cycle of the Anna hummingbird. Univ. Calif. Publ. Zool., 97:1-109.

Geographic variation in the parthenogenetic lizard,Hemidactylus stejnegeriOta &amp; Hikida 1989 (Gekkonidae Reptilia), with comments on recently discovered male phenotypes

A population of the triploid gecko, Hemidactylus stejnegeri Ota & Hikida 1989, was found on an off-shore island of Vietnam for the first time. Chromosomal analyses and multivariate analyses of meristic and morphometric characters revealed that Vietnamese populations are slightly but significantly differentiated from the Philippine and Taiwanese populations. The latter two, on the other hand, are poorly differentiated from each other morphologically. Male phenotypes were found in Vietnamese and Taiwanese populations, and their significance is discussed.