The origin of parachuting locomotion in gekkonid lizards (Reptilia: Gekkonidae)

Several recent studies have focused on the ability of the gekkonid Ptychozoon lionatum to parachute, utilizing its enlarged lateral body folds. Little attention has been paid, however, to the evolution of such behavious and here several aspects pertaining to the origin of parachuting are integrated. Behavioural shifts and morphological modifications are considered together and evaluated in the light of ecological information. It is apparent that certain behavioural shifts were the precursors of subsequent morphological changes. The structural design of the body folds is also considered in relation to parachuting behaviour and both the gross morphology and histology are shown to be significantly modified to permit effective parachuting. The absene of musculature in the lateral body flaps of geckos is confirmed and a model for the integration of the various factors involved in effective parachuting is presented. Here the importance of the "structure-function" and "habitat-behaviour" sets are emphasized together with their new interrelationships when a protoadaptation becomes functional.     

Sexual differences in caudal morphology and its relation to tail autotomy in lacertid lizards

We hypothesized that the presence of the forked hemipenes, and associated musculature, at the base of the tail in male lizards should constrain the capacity to autotomize the tail. Thus, this hypothesis predicts that the non-autotomous base of the tail should be longer in male than in female lizards. We tested this hypothesis in four species oflacertid lizards. Males have on average one to two non-autotomous vertebrae more than females, and the sexual difference in length of the non-autotomous tail base remains constant over the entire body size range. In addition, the first functional autotomy plane in males is usually located on, or is distal to, the vertebrae from which two hemipenial muscles take origin. These observations support the view that functional demands of the male intromittent organs impose constraints on the abilities of tail autotomy. In a natural population of Lacerta vivipara , the proportion of tail breaks that occurred at very short distances from the base was highest in females, indicating that the small sexual difference in length of the non-autotomous tail part is of functional significance. Total length of the tail was largest in males. This can be interpreted as a compensation for the decline in autotomy capacities at the tail base, such that the length of the autotomous part remains similar in both sexes.     

Global diversity of lizards in freshwater (Reptilia: Lacertilia)

No lizards are strictly aquatic, but at least 73 species in 11 different families can be considered to regularly utilize freshwater habitats. There are no aquatic lizards in the Nearctic or Palearctic regions, whereas the Neotropics, Southeast Asia, and the Indo-Australian Archipelago support the greatest diversity of freshwater forms, particularly in the families Gymnophthalmidae, Scincidae and Varanidae. A number of larger aquatic lizards are harvested for food and for the reptile skin trade and several are CITES listed. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Lungs and mesopneumonia of scincomorph lizards (Reptilia: Squamata)

Mesopneumonia of 28 and lungs of 44 species of scincomorph lizards are described, representing mesopneumonia in six of the seven scincomorph families and lungs of all seven families of this taxon. Except for gymnophthalmids and scincids, a family typical organization of mesopneumonia occurs. In cordylids, gerrhosaurids, xantusiids and lacertids the complete right ventral mesopneumonium (VMp) inserts cranially on the posterior vena cava and caudally on the dorsal surface of the right liver lobe. The left VMp is attached to the ventrolateral body wall in cordylids and lacertids; in gerrhosaurids the left VMp is short and inserts on the pericardium and the posterior vena cava; in xantusiids the left VMp is reduced and the lungs are fused cranially with the pericardium. In scincids the VMp of both sides vary in length and insert on the ventral mesentery, or may be lacking completely. The visceral topology of the gymnophthalmids Calyptommatus and Notobachia differs from the general scincomorph pattern, with the liver and stomach elongated and situated on the right and left side of the body cavity, respectively. The left and the right VMp extend over the entire length of the lungs and insert on the ventral mesentery. All lungs examined in the study are single-chambered and show no major structural variability of the inner surfaces, except in lacertids, some gerrhosaurids, Cordylus, and two genera of gymnophthalmids (Echinosaura and Neusticurus). In these groups, rows of dorsomedial niches are present.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-06.htm     

The cybotoid anoles and Chamaelinorops lizards (Reptilia: Iguanidae): evidence of mosaic evolution

Recent immunological comparisons involving the Hispaniolan Chamaelinorops and Anolis suggested that Chamaelinorops might be much more closely related to Hispaniolan species of Anolis than previously thought. Electrophoretic comparisons among the relevant taxa, coupled with a review of both the morphological and immunological data, suggest mosaic evolution may be responsible for the nonconcordance between some of the data sets. An example of significant morphological change with little electrophoretic differentiation was found in the A. cybotes-A. shrevei comparison while the converse was found in the A. cybotes-A. marcanoi comparison. Many of the character states that have been interpreted as implying close genealogical relationship between Chamaelinorops and other Hispaniolan anoles are known to be primitive and may be evolving slowly in these lineages.     

Dewlap and other variation in the lizards Anolis distichus and A. brevirostris (Reptilia: Iguanidae)

The dewlap of male anoline lizards is an important component of the mate recognition system. Much attention has focussed on the role dewlap colour plays in the interactions between species of Anolis but little has been paid to the effects of intraspecific variability in this feature. Patterns of intrasubspecific variability in this feature are described for several subspecies of Anolis distichus and A. brevirostris . In most cases variability in dewlap colour appears unrelated to the amount of electrophoretic variability and the level of population differentiation, although there is some evidence for large scale geographic subdivision within three of the taxa examined. Female choice and the implications for various species concepts are discussed.     

Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology.

DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva--L. fraasi clade and L. brandli are not related to Psammodromus Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian ''Lacerta'' jayakari and ''L.'' cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. ''Lacerta'' jayakari and ''L.'' cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado & Ortega, 1984 is available for the Gallotia-Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.     

Some vaguely explored (but not trivial) costs of tail autotomy in lizards

Lizard tail autotomy is considered an efficient anti-predator strategy that allows animals to escape from a predator attack. However, since the tail also is involved in many alternative functions, tailless animals must cope with several costs following autotomy. Here we explicitly evaluate the consequences of tail autotomy for two costs that have been virtually unexplored: 1. we test whether the anatomical change that occurs after tail loss causes a reduction in the role of the tail as a distraction mechanism to predators; 2. we analyzed whether tail synthesis comprises an energetically costly process in itself, by directly comparing the cost of maintenance before and after autotomy. We found that original tails displace further and at greater velocity than regenerated tails, indicating that the anti-predation responses of a lizard probably changes according to whether its tail is original or regenerated. With regard to the energetic cost of tail synthesis, we observed a significant increase in the standard metabolic rate, which rose 36% in relation to the value recorded prior to tail loss. This result suggests that the energetic cost of tail synthesis itself could be enough to affect lizard fitness.     

Tests on the locomotion of the elongate and limbless reptile Ophisaurus apodus (Sauna: Anguidae)

A series of Ophisaurus apodus was filmed while traversing plane surfaces, fields of nails and pins at different spacings, and channels of different diameter. Small individuals can practise slow lateral undulation on very rough surfaces, but with increased speed, all shift to slide-pushing, using either constrained bends of the body or very wide swings of the tail. In fields of closely-spaced pins, they travel by undulation, often pushing at a limited number of sites and pulling and pushing the trunk among these. Amid pins of wider spacing, the undulation involves some repositioning of curves. In channels, they utilize continuous bend concertina movement with the initial bend formed anteriorly and subsequent ones added, either to the level of the cloaca or on to the tail as well. The cloacal region cannot be established on the basis of locomotor pattern, as the propulsive waves pass smoothly from head to tail. Maximum voluntary velocities (of the centre of gravity) were 13 cm/s for slide-pushing, 55 cm/s for lateral undulation, and 3 cm/s for concertina movement. Propulsion is entirely effected by bending of the trunk; tests gave no evidence that the dorsal and ventral portions of the integumentary armour show significant anteroposterior displacement relative to each other.     

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.     

Morphological, functional and evolutionary aspects of tail autotomy and regeneration in the 'living fossil'Sphenodon (Reptilia: Rhynchocephalia)

Tail autotomy and regeneration are less known in Sphenodon ('Reptilia': Rhynchocephalia) than in Squamata. We examined museum specimens, Sphenodon guntheri ( N  = 8) and Sphenodon punctatus ( N  = 172), wild Sphenodon punctatus ( N  = 19) and Sphenodon sp. skeletons ( N  = 8). In S. punctatus , unlike in typical Squamata, sexes had similar relative (intact) tail lengths, and regeneration frequencies; tail and body growth was isometric. Tail breakage was usually intravertebral, usually followed by ablation of a variably sized terminal vertebral piece, partly deviating from lizards. Hypothetically, imperfect autotomy results from sphenodon's primitiveness. As in squamates, tail-losers were morphologically more left-side dominant than tail-retainers. Individual directional asymmetries in digit morphology and in digit injury were correlated (in lizards observed only at population level); tail-losers had more fluctuating asymmetry but their exclusion did not facilitate morphological taxonomic distinctions (no 'Seligmann effect'). In S. punctatus , extents and directions of sexual dimorphism paralleled differences between tail-retainers and tail-losers, females resembling tail-losers, also accounting for character interdependence (developmental constraints; employing a method similar to phylogenetic contrasts). The variation in the location of tail injury was correlated with the continuum of variation between injured and intact (pholidotic) morphotypes. These last two phenomena remain to be explored in Squamates.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 721–743.     

Ontogeny of hemopoietic and lymphopoietic tissues in the lizard Chalcides ocellatus (Reptilia,Sauna, Scincidae)

The first and major blood-forming organ to develop in the viviparous lizard Chalcides ocellatus is the yolk sac, which exhibits prominent erythropoietic activity from as early as stage 21 through birth (stage 41). Myeloid cells and megakaryocytes are produced in the yolk sac from stage 23 onward. During lizard embryogenesis hemopoietic activity is also observed in spleen and bone marrow but in neither kidney nor liver. Cells capable of giving rise to lymphocytes both in vivo and in vitro are first found in the thymus at stage 35. Active lymphopolesis in thymus and spleen begins at stages 36 and 39, respectively. In contrast, the gut-associated lymphoid aggregates are not evident before birth.     

The comparative aspects of spatial ecology of lizards exemplified by the toad-headed lizards (Reptilia, Agamidae, Phrynocephalus)

The possibility of analysis of phylogenetic parameters of the spatial distribution of populations is discussed by an example of the agamid toad-headed lizards (Phrynocephalus). Summarizing both original and published data on the individual home ranges and the relocation of individuals of 30 populations from 12 species showed that differentiation of the type of spatial distribution is weak in toad-headed lizards. This observation confirms the idea that this clade of agamids is phylogenetically young and relatively recently radiated. At the interspecific level, positive correlation between home range size and body size was observed in the studied group. Such spatial parameters, shared by all toad-headed lizards, as relatively large size and weakly structured individual home ranges can be explained by the peculiarities of their reproduction features and their foraging mode. The individual type of space-usage in toad-headed does not fit the traditional scheme dividing all the lizards into the territorial Iguania and the nonterritorial Autarchoglossa.     

Structural characteristics of the patagium of Ptychozoon kuhli (Reptilia: Gekkonidae) in relation to parachuting locomotion

Ptychozoon kuhli is known for its parachuting/gliding capabilities. In this contribution, we document the allometric scaling properties of its patagium, accessory flaps and folds, and its total body surface area and compare them to similar attributes of the agamine lizard Draco volans. Ptychozoon kuhli has passive patagia that lack skeletal support and muscular control. Patagial area in P. kuhli is smaller than that in D. volans for individuals of identical snout-vent length, but the accessory folds and flaps compensate for this shortfall and overall P. kuhli has equivalent total body surface area to D. volans for equally sized individuals. The structure of the patagium of Ptychozoon kuhli was investigated in terms of its scalation patterns and structural integrity, its relationship to the body wall and its mechanism of erection, and the distribution of mechanoreceptive sensilla across its surface. Scalation patterns and the internal architecture of the patagium indicate how its shape and form are maintained as it erects and becomes exposed to air flow. Its cross-sectional shape, together with that of the entire body indicates how it is able to behave as an airfoil. The distribution of sensilla across the patagial surface reflect positioning indicative of the monitoring of scale-to-scale contact on the dorsal surface, and possibly air pressure and flow on the ventral surface.     

Carpus and tarsus patterns in some Arabian lizards (Reptilia: Agamidae, Lacertidae, Scincidae)

Within the families Agamidae, Lacertidae and Scincidae, the carpal and tarsal elements, the sesamoidea and the ossification pattern show considerable variability. Furthermore, carpal and tarsal structure in the lizards studied here is not unique to the genera in question and identical or very similar conditions occur in other lizards. These elements contain sufficient anatomical information to make them taxonomically valuable.     

Placental morphology in two sympatric Andean lizards of the genus Liolaemus (Reptilia: Liolaemidae)

Viviparity is a remarkable feature in squamate sauropsids and it has evolved multiple times in parallel with the formation of a placenta. One example of this repeated evolution of viviparity and placentation occurs in the species‐rich South American genus Liolaemus with at least six independent origins of viviparity. However, evolutionary studies of placentation in this genus are limited by a lack of data on placental morphology. The aim of this study is to describe and compare the microanatomy and vessel diameter (Dv, a function of blood flow) of the placenta using scanning electron microscopy (SEM) and confocal laser scanning microscopy (cLSM) in two sympatric Andean viviparous but highly divergent species, Liolaemus robustus and Liolaemus walkeri. We found interspecific differences in cell types in the chorion, allantois, and omphalopleure that may be explained by divergent phylogenetic history. Time elapsed since divergence may also explain the pronounced interspecific differences in vessel diameter, and within each species, there are strong differences in Dv between tissue locations. Both species show features to improve gas exchange in the chorioallantoic placenta including absence of eggshell, large Dv in the allantois (L. robustus) or embryonic side of the uterus (L. walkeri), and when present, microvillous cells in the allantois (L. walkeri). Both species also show features that suggest transfer of nutrients or water in the omphaloplacenta, including an almost complete reduction of the eggshell, secretive material (L. robustus), or vesicles (L. walkeri) on cell surface uterus, and when present specialized cells in the omphalopleure (L. walkeri). No statistical differences in Dv were found among stages 32–39 in each species, suggesting that a different mechanism, other than enhanced blood flow, might satisfy the increased oxygen demand of the developing embryos in the hypoxic environments of the high Andes. J. Morphol. 276:1205–1217, 2015. © 2015 Wiley Periodicals, Inc.     

Density and distribution of cutaneous sensilla on tails of leopard geckos (Eublepharis macularius) in relation to caudal autotomy

The lizard tail is well known for its ability to autotomize and regenerate. Physical contact of the tail by a predator may induce autotomy at the location at which the tail is grasped, and upon detachment the tail may undergo violent, rapid, and unpredictable movements that appear to be, to some degree, regulated by contact with the physical environment. Neither the mechanism by which tail breakage at a particular location is determined, nor that by which environmental feedback to the tail is received, are known. It has been suggested that mechanoreceptors (sensilla) are the means of mediation of such activities, and reports indicate that the density of sensilla on the tail is high. To determine the feasibility that mechanoreceptors are involved in such phenomena, we mapped scale form and the size, density, distribution, and spacing of sensilla on the head, body, limbs, and tail of the leopard gecko. This species has a full complement of autotomy planes along the length of the tail, and the postautotomic behavior of its tail has been documented. We found that the density of sensilla is highest on the tail relative to all other body regions examined; a dorsoventral gradient of caudal sensilla density is evident on the tail; sensilla are more closely spaced on the dorsal and lateral regions of the tail than elsewhere and are carried on relatively small scales; and that the whorls of scales on the tail bear a one to one relationship with the autotomy planes. Our results are consistent with the hypotheses of sensilla being involved in determining the site at which autotomy will occur, and with them being involved in the mediation of tail behavior following autotomy. These findings open the way for experimental neurological investigations of how autotomy is induced and how the detached tail responds to external environmental input. J. Morphol. 275:961–979, 2014. © 2014 Wiley Periodicals, Inc.