Epidermal differentiation in the developing scales of embryos of the Australian scincid lizard Lampropholis guichenoti

Formation of the first epidermal layers in the embryonic scales of the lizard Lampropholis guichenoti was studied by optical and electron microscopy. Morphogenesis of embryonic scales is similar to the general process in lizards, with well‐developed overlapping scales being differentiated before hatching. The narrow outer peridermis is torn and partially lost during scale morphogenesis. A second layer, probably homologous to the inner peridermis of other lizard species, but specialized to produce lipid‐like material, develops beneath the outer peridermis. Two or three lipogenic layers of this type develop in the forming outer surface of scales near to the hinge region. These layers form a structure here termed “sebaceous‐like secretory cells.” These cells secrete lipid‐like material into the interscale space so that the whole epidermis is eventually coated with it. This lipid‐like material may help to reduce friction and to reduce accumulation of dirt between adjacent extremely overlapping scales. At the end of their differentiation, the modified inner periderm turns into extremely thin cornified cells. The layer beneath the inner peridermis is granulated due to the accumulation of keratohyalin‐like granules, and forms a shedding complex with the oberhautchen, which develops beneath. Typically tilted spinulae of the oberhautchen are formed by the aggregation of tonofilaments into characteristically pointed cytoplasmic outgrowths. Initially, there is little accumulation of β‐keratin packets in these cells. During differentiation, the oberhautchen layer merges with cells of the β‐keratin layer produced underneath, so that a typical syncytial β‐keratin layer is eventually formed before hatching. Between one‐fourth distal and the scale tip, the dermis under epidermal cells is scarce or absent so that the mature scale tip is made of a solid rod of β‐keratinized cells. At the time of hatching, differentiation of a mesos layer is well advanced, and the epidermal histology of scales corresponds to Stage 5 of an adult shedding cycle. The present study confirms that the embryonic sequence of epidermal stratification observed in other species is basically maintained in L. guichenoti. J. Morphol. 241:139–152, 1999. © 1999 Wiley‐Liss, Inc.     

Costs of reproduction in a tropical invariant-clutch producing lizard (Carlia rubrigularis)

Reproduction involves costs and benefits, whereby increased expenditure into current reproduction can reduce future reproductive success. Costs of reproduction, often assessed using the reduction in locomotor speed, have become well established in temperate-zone lizard species. However, substantial differences in biotic conditions and life-history traits between temperate- and tropical-zone lizards suggest that such costs may be different or of less importance for tropical species. This study examined the effects of reproduction on locomotor speed in the tropical invariant-clutch producing lizard Carlia rubrigularis . Counter to predictions and despite a low relative clutch mass (RCM), gravid and post-oviposition females experienced a reduction in locomotor speed with a physiological basis that was unrelated to the level of reproductive investment. In addition, gravid and postoviposition females exhibited locomotor speeds that were inversely related to the timing of oviposition and approached the speed of non-reproductive females after 3 weeks of oviposition. These results suggest that in addition to RCM, selection may act to reduce the period of recovery in species with extended reproductive seasons and which make numerous bouts of reproduction, such as C. rubrigularis .     

Geographic variation in lizard phenotypes: importance of the incubation environment

Geographic variation in phenotypes can result from proximate environmental effects as well as from underlying genetic factors. Reciprocal transplant experiments, in which organisms are moved from one area to another, offer a powerful technique to partition the effects of these two factors. However, many studies that have utilized this technique have focused on the post-hatching organism only and ignored potential effects of environmental influences acting during embryonic development. We examined the phenotypic responses of hatchling scincid lizards ( Lampropholis guichenoti ) incubated in the laboratory under thermal regimes characteristic of natural nests in two study areas in southeastern Australia. Although the sites were less than 120 km apart, lizards from these two areas differed in thermal regimes of natural nests, and in hatchling phenotypes (morphology, locomotor performance). We incubated eggs from each area under the thermal regimes typical of both sites. Some of the traits we measured (e.g. hatchling mass and snout-vent length) showed little or no phenotypic plasticity in response to differences in incubation conditions, whereas other traits (e.g. incubation period, tail length, inter-limb length, body shape, locomotor performance) were strongly influenced by the thermal regime experienced by the embryo. Thus, a significant proportion of the geographic variation in morphology and locomotor performance of hatchling lizards may be directly induced by differences in nest temperatures rather than by genetic divergence. We suggest that future studies using the reciprocal transplant design should consider environmental influences on all stages of the life-history, including embryonic development as well as post-hatching life.     

Morphological correlates of life‐history variation: is lizard clutch size related to the number of germinal beds in the ovary?

Clutch size varies widely in reptiles, both intraspecifically and interspecifically. The mechanisms that generate this variation have attracted detailed study, focusing primarily on ecological factors (e.g. food availability), trade‐offs with other traits (e.g. offspring size), and physical constraints (e.g. maternal body shape). Does ovarian morphology, specifically the number of germinal beds from which ova are produced, also correlate with clutch size? Our review of published data on 58 lizard species reveals that clutch size is correlated with the number of germinal beds per ovary (more fecund species have more germinal beds), and that phylogenetic changes in germinal beds have been consistently associated with concurrent phylogenetic changes in fecundity. These correlations imply a causal connection: either clutch size is constrained by ovarian morphology, and/or ovarian morphology evolves to allow adaptive shifts in clutch size. The latter hypothesis is more consistent with available data. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94 , 81–88.     

The adaptive significance of colour pattern polymorphism in the Australian scincid lizard Lampropholis delicata

Females of Lampropholis delicata are dimorphic for colour pattern, the difference between morphs being the presence or absence of a distinct white mid-lateral stripe. A less distinct striped morph occurs also in males. We evaluated alternative hypotheses for the maintenance of this polymorphism by examining temporal and spatial variation in morph frequency, testing for differential selection among morphs using data on body size and reproductive traits from preserved specimens, and experimentally manipulating colour pattern in free-ranging lizards of both sexes, to assess the influence of the lateral stripe on survival rates. We found that the relative frequency of striped individuals varied among populations and decreased from north to south in both sexes, coincident with an increasing incidence of regenerated tails. Morph frequencies did not change through time within a population. Striped gravid females appeared to survive better and produced larger clutches than did non-striped females. In our experimental study, the relationship between survival and colour morph differed between the two sexes; males painted with a white lateral stripe had lower survival than control (brown stripe) males, but survival did not differ between striped and control females. The different response in the two sexes may be due partly to differences in temperature and microhabitat selection. We propose that the white lateral stripe decreases susceptibility to predators in gravid females but increases risk of predation in males, especially in combination with low temperatures. The polymorphism might be maintained by: (1) opposing fitness consequences of the stripe in males and females; (2) sex-specific habitat selection; and (3) gene flow in combination with spatial variation in relative fitness of the two morphs.     

The influence of hydric environments during egg incubation on embryonic heart rates and offspring phenotypes in a scincid lizard (Lampropholis guichenoti)

Extensive evidence shows that incubation conditions can affect phenotypic traits of hatchling reptiles, but the relative importance of thermal versus hydric factors, and the proximate mechanisms by which such factors influence hatchling phenotypes, remain unclear for most species. We incubated eggs of an Australian scincid lizard, Lampropholis guichenoti, at four different moisture contents ranging from -500 to 0 kPa. Drier substrates reduced water uptake of eggs and resulted in smaller hatchlings, but other phenotypic traits (incubation periods, hatchling sex, body proportions, running speeds, growth rates post-hatching) were not affected by the hydric environment during incubation. Contrary to our prediction, lower water uptake during incubation (and hence, presumably, more viscous blood) did not affect embryonic heart rates. Thus, as in many other squamate species, hatchling phenotypes and embryonic developmental rates of L. guichenoti are less sensitive to hydric conditions in the nest than to thermal regimes.     

A phylogenetic analysis of variation in reproductive mode within an Australian lizard (Saiphos equalis, Scincidae)

Saiphos equalis , a semi-fossorial scincid lizard from south-eastern Australia, is one of only three reptile species world-wide that are known to display geographic variation in reproductive mode. Uniquely, Saiphos equalis includes populations with three reproductive modes: oviparous with long (15-day) incubation periods; oviparous with short (5-day) incubation periods; and viviparous (0-day incubation periods). No Saiphos populations show 'normal' scincid oviparity (> 30-day incubation period). We used mitochondrial nucleotide sequences ( ND2 and cytochrome b ) to reconstruct relationships among populations from throughout the species' distribution in New South Wales, Australia. Under the phylogenetic species concept, phylogenetic analyses are consistent with the oviparous and viviparous populations of S. equalis being conspecific. Phylogenetic analyses suggest that the long incubation period oviparous lineage is the sister group to all other populations; and that the viviparous populations belong to a cluster of weakly supported clades basal to the short-incubation-period oviparous clade. These clades correspond to variation in reproductive mode and geographic location.     

Tyrosine phosphorylated proteins in the reproductive tract of the viviparous lizard Eulamprus tympanum and the oviparous lizard Lampropholis guichenoti

Plastic changes occur in the morphology of the uterus at various stages of the reproductive cycle in both oviparous and viviparous lizards and these may be influenced by estrogen. Estrogen driven phosphorylation of effector proteins on tyrosine residues plays a major role in the plastic modulation of uterine anatomy and physiology in vertebrates. We used electrophoresis and Western blotting to characterize the phosphotyrosine protein profiles at various stages of the reproductive pathway in an oviparous lizard Lampropholis guichenoti and a viviparous lizard Eulamprus tympanum. L. guichenoti displayed major bands in the 200-35 kDa range and a triplet of bands of molecular masses 61 kDa, 52 kDa and 48 kDa in 50% of specimens and a 38 kDa band in all specimens. In contrast, E. tympanum samples all displayed a single major band at 40 kDa, which was significantly elevated at the early pregnancy stage. Somewhat paradoxically, the viviparous species, which has the more complex uterine epithelial changes during pregnancy, has the fewest phosphotyrosine bands, so how tyrosine phosphorylation is affected during the evolution of viviparity is not clear.     

HoxA10-like proteins in the reproductive tract of the viviparous lizard Eulamprus tympanum and the oviparous lizard Lampropholis guichenoti

The gene HoxA10 and its protein product are essential for the formation of the extensions of the plasma membrane called uterodomes or pinopods in mammalian uterine epithelia. In mice, the presence of the HoxA10 protein and uterodomes is needed for uterine receptivity to blastocyst implantation. The viviparous lizard Eulamprus tympanum displays uterodomes whereas the oviparous lizard Lampropholis guichenoti does not. To explore the theory that HoxA10 is involved in the formation of uterodomes we investigated whether HoxA10 immunoreactive proteins were present in both species during their reproductive cycles. Oviduct proteins from vitellogenic, gravid or non-reproductive L. guichenoti (n=19) and E. tympanum (n=28) were separated by electrophoresis and analysed by Western blot and specific antibodies to HoxA10. E. tympanum displayed HoxA10 immunoreactive bands at 59 and 63 kDa in 20 out of the 28 samples. All of the L. guichenoti samples displayed HoxA10 immunoreactive bands, 18 had bands at 59 and 64 kDa and 1 animal had a single band at 59 kDa. There were no significant differences in the level of HoxA10 immunoreactivity between the different stages of reproductive cycle in either species. The different molecular mass of the larger band in L. guichenoti (64 kDa) compared to E. tympanum (63 kDa) indicates that the two lizards express different isoforms of the HoxA10-like proteins and it will be interesting in future studies to determine whether there are differences in the biological activity of the proteins that regulate different physiological functions in the uterus of viviparous and oviparous lizards.     

Geographic variation in clutch size and a realized benefit of aggregative feeding

We investigated one causal explanation for geographic variation in clutch size and aggregative feeding of the pipevine swallowtail, Battus philenor. Populations in California lay larger clutches than those in Texas, and larger feeding aggregations grow at an accelerated rate on the California host plant. Using reciprocal transplant experiments with larvae from California and Texas populations, we found that the benefit of increased growth rate associated with feeding in larger groups occurred only on the California host plant and was observed for larvae from both populations. These results are consistent with the hypothesis that larger clutch size and aggregative feeding are adaptations to characteristics of the California host plant. Future studies on the evolution of clutch size and aggregative feeding of herbivorous insects should consider how these life-history traits affect host plant suitability.     

Local richness and distribution of the lizard fauna in natural habitat mosaics of the Brazilian Cerrado

We investigate local lizard richness and distribution in central Brazilian Cerrado, harbouring one of the least studied herpetofaunas in the Neotropical region. Our results are based on standardized samplings at 10 localities, involving 2917 captures of 57 lizard species in 10 families. Local richness values exceeded most presented in earlier studies and varied from 13 to 28 species, with modal values between 19 and 28 species. Most of the Cerrado lizard fauna is composed of habitat‐specialists with patchy distributions in the mosaic of grasslands, savannas and forests, resulting in habitat‐structured lizard assemblages. Faunal overlap between open and forested habitats is limited, and forested and open areas may act as mutual barriers to lizard distribution. Habitat use is influenced by niche conservatism in deep lineages, with iguanians and gekkotans showing higher use of forested habitats, whereas autarchoglossans are richer and more abundant in open habitats. Contrary to trends observed in Cerrado birds and large mammals, lizard richness is significantly higher in open, interfluvial habitats that dominate the Cerrado landscape. Between‐localities variation in lizard richness seems tied to geographical distance, landscape history and phylogenetic constraints, factors operating in other well‐studied lizard faunas in open environments. Higher richness in dominant, open interfluvial habitats may be recurrent in Squamata and other small‐bodied vertebrates, posing a threat to conservation as these habitats are most vulnerable to the fast, widespread and ongoing process of habitat destruction in central Brazil.     

Morphological variation in the Cape Dwarf Chameleon (Bradypodion pumilum) as a consequence of spatially explicit habitat structure differences

An organism's phenotype is to some extent influenced by costs and benefits in terms of natural and sexual selection. The intensity of natural selection can in part be driven by habitat structure, which may result in varying levels of crypsis and/or selection on traits related to maximizing performance in that habitat. This may be countered by sexual selection, which can lead to sexual dimorphism in body size and/or the expression of conspicuous ornamentation relating to maximizing reproductive success. The intensity of these forces can also be different between the sexes, resulting in complex patterns of phenotypic variation. With this in mind, we examined morphological variation within the Cape Dwarf Chameleon, Bradypodion pumilum. The species inhabits two geographically disjunct habitat types and, in the present study, we demonstrate that chameleons from the two habitats show morphological differences. Large, conspicuous individuals inhabit closed vegetation, whereas small, drab individuals inhabit open vegetation. However, when morphological traits are size‐adjusted, the open vegetation morph displays many traits that are larger for its body size than the closed vegetation morph, especially for characters related to locomotion (limbs) and bite force (head width). Sexual dimorphism is also present, although the degree and number of dimorphic characters was very different between the two morphs, with size‐adjusted male‐biased dimorphism much more pronounced in the closed morph. Overall, our findings suggest that natural selection in open habitats limits both body size and conspicuous characters, although sexual selection in closed habitats favours the development of ornamentation related to display. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 878–888.     

Calcium ATPase expression in the oviducts of the skink, Lampropholis guichenoti

Lampropholis guichenoti is an oviparous lizard that lays eggs with a calcareous outer shell. We used immunofluorescence microscopy to describe the occurrence and distribution of Ca2+ ATPase pumps in the uterus of L. guichenoti at different stages of the reproductive and egg-shelling cycles. Ca2+ ATPase pumps were not demonstrated by immunofluorescent techniques in any uterine tissue until egg-shelling had commenced and at least partly calcified eggs were in the uterus. During egg-shelling, Ca2+ ATPase pumps occur on the apical and baso-lateral surfaces of uterine epithelial cells, and those of associated shell glands in the stroma of the uterus. We conclude that Ca2+ ATPase pumps provide a major mechanism for deposition of the calcareous eggshell of L. guichenoti and that the pumps are up-regulated when required in the reproductive cycle. Furthermore, it is likely that specific calcium glands in the stroma of the uterus are involved in the rapid transport required for egg-shelling, but the differential contribution of luminal and glandular epithelial cells is not known.     

池养条件下似刺鳊鮈早期发育阶段的摄食与生长特性

对人工培育的似刺鳊鮈（Paracanthobrama guichenoti）仔鱼、稚鱼和幼鱼的生长与摄食节律进行了研究。结果显示,在1～70日龄间,似刺鳊鮈全长平均日增长率为3．55％,体重平均日增长率为11．26％,头长／全长值随着日龄的增长而变小。全长（L,mm）与体重（M,mg）的关系式为M=O．0012L^2-0．0564L＋0．6504;体重（M,mg）与日龄（D,d）的关系式为M=O．0043e^0.0936D;全长（L,mm）和日龄（D,d）的关系式为L=0．0154D^2-0．0128D＋8．1965。在饵料充足的条件下,20日龄晚期仔鱼全天均有很高的摄食发生率,小高峰出现在8：00时;40日龄稚鱼则表现出明显的昼夜摄食节律,属于典型的白天摄食类型,日摄食率2．3376％。20日龄仔鱼和4013龄稚鱼饱食后分别经6h、5h整个消化道排空。在水温25℃及饥饿空胃状态下,10日龄仔鱼只经3d即开始出现死亡个体,经5d全部死亡;20日龄仔鱼经6d开始出现死亡个体,经9d全部死亡;40日龄稚鱼经7d才开始出现死亡个体,10d后全部个体均死亡。     

Post-hatching environment contributes greatly to phenotypic variation between two populations of the Australian garden skink, Lampropholis guichenoti

While recent experimental work on a variety of reptile species has demonstrated that incubation temperature influences hatchling phenotypes, the biological significance of such phenotypic variation remains unclear. Incubation temperature may exert significant long-term phenotypic effects. Alternatively, such influences may be temporary, or negligible relative to effects induced by genetic factors, or by the environmental conditions experienced after hatching. Even if incubation temperature exerts long-term effects on phenotype, this might occur indirectly (by influencing hatching dates) rather than by direct modifications of developmental processes. We quantified the influences of the source population, incubation temperature and rearing environment, on the phenotype of the Australian garden skink (Lampropholis guichenoti) from populations that differ in nest temperature and phenotype. Intcrpopulation differences in the phenotypes of young lizards were found to be a product of all three factors. However, the long-term effects of both population and incubation temperature operated indirectly (through variation in the date of hatching) rather than directly (through genetic or developmental factors). That is, once all temporal effects were removed, the only discernible influence on juvenile phenotypes was their rearing environment. Thus, some of the most important influences on lizard phenotypes may operate via modifications of hatching date.     

Changes in oviductal morphology of the skink, Lampropholis guichenoti, associated with egg production

We describe changes in the morphology of the oviductal epithelium of an oviparous skink, Lampropholis guichenoti, during the course of egg production and oviposition: to characterize the luminal epithelial changes; to provide a baseline for understanding uterine changes in viviparous species; and to establish whether the plasma membrane transformation of uterine epithelial cells is indeed a feature restricted to viviparous species. Oviducts from vitellogenic, gravid, and postgravid females were observed using scanning electron microscopy. Cellular characteristics of the oviductal epithelium previously used to determine the plasma membrane transformation were assessed morphologically. Three anatomically different areas were defined within the oviduct, but no plasma membrane transformation was observed in the oviparous skink, suggesting that this is a phenomenon particular to viviparity.     

长江上游圆口铜鱼幼鱼肠道内寄生小吻虫种群生态的研究

对长江上游圆口铜鱼幼鱼肠道内寄生小吻虫种群生态的研究表明,长江上游圆口铜鱼幼鱼体内小吻虫感染率为69．88％,感染强度8．18(1-47),相对密度5．72。雌鱼的感染率和感染强度均高于雄鱼,体长4．0～5．9cm的圆口铜鱼被小吻虫的感染率最高,且染虫对肥满度有显著影响。小吻虫在圆口铜鱼幼鱼种群中呈负二项分布,66％的虫体集丛分布在圆口铜鱼肠道的前段,其种群主要由5mm以下个体组成,雌虫明显多于雄虫。     

Reproduction of varanid lizards (Reptilia: Squamata: Varanidae) at the Bronx Zoo

Captivity provides many opportunities for studying the reproductive biology of varanid lizards (Family: Varanidae; genus: Varanus). This article presents reproductive data for 12 varanid species maintained in captivity at the Bronx Zoo between 1977 and 2010. Five species, V. prasinus, V. beccarii, V. melinus, V. mertensi, and V. ornatus have successfully reproduced at the zoo, for which clutch, incubation, and hatchling data are presented, and compared with previously published values. Clutch, incubation, and hatchling measurements typically fell within the known ranges for each species, or represent minor range extensions. V. mertensi was the only species that reproduced seasonally (χ(2) = 9.061, P = 0.029). Five species multiclutched; individual V. mertensi and V. prasinus females produced as many as four clutches per year. The shortest intervals between successive clutches observed for V. kordensis (61 days), V. melinus (77 days), V. mertensi (51 days), and V. prasinus (68 days) represent the shortest known values for each species. Relative clutch mass (RCM) ranged from 0.187 to 0.437. A mean RCM of 0.370 ± 0.092 for V. prasinus is substantially higher than previously published values for the species. These results illustrate that when maintained under favorable conditions in captivity, some varanids are capable of remarkably high fecundities, rapid reproductive turnover rates, and sizeable maternal investments. Periodicals, Inc.     

Geographic variation and systematic review of the lizard genus Vanzosaura (Squamata,Gymnophthalmidae), with the description of a new species

Vanzosaura rubricauda (Boulenger, 1902) is a small‐bodied gymnophthalmid lizard widespread in dry biomes of South America. Throughout its distribution, V. rubricauda populations experience contrasting environmental conditions, and a marked disjunction occurs in the central portion of the Cerrado biome. Previous studies indicate that V. rubricauda may be a species complex, and here we used mitochondrial DNA data and external morphology to test for population differentiation and its systematic implications for the genus. We found three geographically cohesive groups recovered as well resolved, and strongly supported mitochondrial DNA (mtDNA) haploclades that are differentiated with respect to morphometry, meristics, and colour pattern. In light of the observed differences, we propose a new taxonomic rearrangement of the genus, where we: (1) restrict V. rubricauda to Chaco and western Cerrado regions; (2) resurrect V anzosaura multiscutata (Amaral, 1933) comb. nov. for Caatinga populations; and (3) describe V anzosaura savanicola sp. nov. for the eastern Cerrado region of Brazil. The new species is diagnosed from its congeners by having longer limbs and tail, fewer smooth subcaudals, and genetic distances ranging from 5 to 13%. The new species, along with other recent discoveries, highlights the Jalapão‐Serra Geral region as one of the most important areas of endemism for Cerrado squamates. © 2014 The Linnean Society of London