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     

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.     

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.     

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.     

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.     

New insights into sex chromosome evolution in anole lizards (Reptilia,Dactyloidae)

Anoles are a clade of iguanian lizards that underwent an extensive radiation between 125 and 65 million years ago. Their karyotypes show wide variation in diploid number spanning from 26 (Anolis evermanni) to 44 (A. insolitus). This chromosomal variation involves their sex chromosomes, ranging from simple systems (XX/XY), with heterochromosomes represented by either micro- or macrochromosomes, to multiple systems (X₁X₁X₂X₂/X₁X₂Y). Here, for the first time, the homology relationships of sex chromosomes have been investigated in nine anole lizards at the whole chromosome level. Cross-species chromosome painting using sex chromosome paints from A. carolinensis, Ctenonotus pogus and Norops sagrei and gene mapping of X-linked genes demonstrated that the anole ancestral sex chromosome system constituted by microchromosomes is retained in all the species with the ancestral karyotype (2n?=?36, 12 macro- and 24 microchromosomes). On the contrary, species with a derived karyotype, namely those belonging to genera Ctenonotus and Norops, show a series of rearrangements (fusions/fissions) involving autosomes/microchromosomes that led to the formation of their current sex chromosome systems. These results demonstrate that different autosomes were involved in translocations with sex chromosomes in closely related lineages of anole lizards and that several sequential microautosome/sex chromosome fusions lead to a remarkable increase in size of Norops sagrei sex chromosomes.     

Daily patterns of metabolic rate among New Zealand lizards (Reptilia: Lacertilia: Diplodactylidae and Scincidae)

In addition to the effects of temperature fluctuations on metabolic rate, entrained endogenous rhythms in metabolism, which are independent of temperature fluctuations, may be important in overall energy metabolism in ectotherms. Daily entrained endogenous rhythms may serve as energy-conserving mechanisms during an animal's active or inactive phase. However, because nocturnal lizards often take advantage of thermal opportunities during the photophase (light), their daily metabolic rhythms may be less pronounced than those of diurnal species. We measured the rate of oxygen consumption (VO(2)) as an index of metabolic rate of eight temperate lizard species (four nocturnal, three diurnal, and one crepuscular/diurnal; n = 7-14) over 24 h at 13 degrees C and in constant darkness to test whether daily patterns (including amplitude, magnitude, and time of peak VO(2)) of metabolic rate in lizards differ with activity period. We also tested for phylogenetic differences in metabolic rate between skinks and geckos. Three daily patterns were evident: 24-h cycle, 12-h cycle, or no daily cycle. The skink Cyclodina aenea has a 12-h crepuscular pattern of oxygen consumption. In four other species, VO(2) increased with, or in anticipation of, the active part of the day, but three species had rhythms offset from their active phase. Although not correlated with activity period or phylogeny, amplitude of VO(2) may be correlated with whether a species is temperate or tropical. In conclusion, the metabolic rate of many species does not always correlate with the recorded activity period. The dichotomy of ecology and physiology may be clarified by more in-depth studies of species behaviors and activity periods.     

Body elongation and decreased reproductive output within a restricted clade of lizards (Reptilia: Scincidae)

Relationships between body shape and relative abdominal size were compared among differentially elongate species within the scincid lizard genus Brachymeles , to investigate how morphological evolution affects the proportion of body volume available to hold eggs and offspring. Relative abdominal size is inversely related to elongation, suggesting that relative clutch mass decreases with addition of abdominal body segments. Shape-volume relationships contradict trends seen in comparisons among distantly related limbed and limbless squamates (lizards and snakes), in which snakes have relatively more abdominal volume. Comparison within a phylogenetically restricted group allows the identification of functional and ontogenetic factors potentially limiting reproductive output. In Bruchymeles , constraining factors include retention of anterior body segments bearing parasternal ribs, which prevents extension of the clutch anteriorly within the body, and reduction of allometry of abdominal segments, which provides extended series of uniformly-sized vertebrae for limbless locomotion, but reduces the relative size of the abdomen. The latter trait is associated with overall size reduction, which affects relative egg-size and packing. Factors constraining abdominal volume in this genus are probably common to other elongate lizards, a morphological group that has been rarely represented in comparative studies of life history.     

Hemiclitoris discovered: a fully differentiated erectile structure in female monitor lizards (Varanus spp.) (Reptilia: Varanidae)

In female monitor lizards (Varanus spp.), the ubiquitous presence of paired evertible and erectile structures in the ventral portion of the tail root, which are miniaturized mirror images of the hemipenes of the males, was discovered. These organs contain all the structural elements (including a forked retractor muscle, supporting ossifications, epidermal flounces etc.) that are also characteristic for the hemipenes of the respective conspecific males. Consequently, it is proposed that the term hemiclitoris be used for this organ. A preliminary survey showed this organ to be present in several other squamate families, suggesting that it is ubiquitous in all female Squamata.     

Hemiclitoris discovered: a fully differentiated erectile structure in female monitor lizards (Varanus spp.) (Reptilia: Varanidae)

In female monitor lizards ( Varanus spp.), the ubiquitous presence of paired evertible and erectile structures in the ventral portion of the tail root, which are miniaturized mirror images of the hemipenes of the males, was discovered. These organs contain all the structural elements (including a forked retractor muscle, supporting ossifications, epidermal flounces etc.) that are also characteristic for the hemipenes of the respective conspecific males. Consequently, it is proposed that the term hemiclitoris be used for this organ. A preliminary survey showed this organ to be present in several other squamate families, suggesting that it is ubiquitous in all female Squamata.

Zusammenfassung

Die Entdeckung der Hemiclitoris, eines differenzierten erektilen und ausstülpbaren Organs bei Waranweibchen (Reptilia: Varanidae)
In der Schwanzwurzel weiblicher Warane ( Varanus spp.) wurde das obligate Vorhandensein paariger, ausstülpbarer und erektiler Strukturen entdeckt, welche verkleinerte Spiegelbilder der Hemipenes der Männchen darstellen. Diese Organe enthalten sämtliche Strukturelemente (z.B. den gegabelten Rückziehmuskel, Stützknochen, epidermale Paryphasmen), die auch für die Hemipenes der betreffenden artgleichen Männchen charakteristisch sind. Ein erster Erfassungsansatz (mit ausführlicher Artenliste in einer demnächst zu publizierenden, derzeit erstellten Diplomarbeit) zeigt, daß dieses Organ auch bei diversen weiteren Squamatenfamilien nachweisbar ist. Dies legt nahe, daß es tatsächlich obligat bei alien weiblichen Squamata ausgebildet ist.     

Parapatric distribution of the lizards Plestiodon (formerly Eumeces) latiscutatus and P. japonicus (Reptilia: Scincidae) around the Izu Peninsula, Central Japan, and its biogeographic implications

The scincid lizard Plestiodon latiscutatus is found in the Izu Islands and Izu Peninsula of central Japan, whereas P. japonicus, a close relative, is found over the entire main island group of Japan, except the Izu Peninsula. The precise area of occupancy of these species was surveyed around the Izu Peninsula. Species identification was made through comparison of mitochondrial DNA partial sequences of specimens from the Izu Peninsula with those from the other regions, since morphological differences between these species have not yet been characterized. This study determined that these species are deeply diverged from each other in mitochondrial DNA sequence, and that the ranges of these species overlap only in a narrow zone. The results imply that gene flow between these species, if any, is restricted to a low level, without physical barriers. The boundary between the geographic ranges of these species was established as occurring along the lower Fuji River, Mt. Fuji, and the Sakawa River. This region is concordant with that of the old sea that is assumed to have separated the Izu Peninsula from other parts of the Japanese main island group until the middle Pleistocene. This pattern suggests that P. latiscutatus and P. japonicus were differentiated allopatrically before the connection of land areas of the Izu Peninsula and Honshu, the main island of Japan, and come into secondary contact through this connection. Thus, the species boundary is likely to have been maintained in situ, without physical barriers, since the secondary contact in the middle Pleistocene.