The potential utility of postnatalskeletal developmental patterns in squamate phylogenetics

Postnatal patterns of skeletal development, includingthe sequence of appearance of ossification centres and the distributionof sesamoids, appear to be highly conserved species-level phenomenain squamates. As such, they are a potential source of charactersfor phylogenetic inquiry. These patterns, from 21 species representing14 crown squamate clades, form the basis for two analyses. In thefirst, the sequence of postnatal skeletal events is coded as charactersusing the sequence unit approach. This analysis reveals that thesequence of postnatal skeletal events might be useful for determiningrelationships at or above the level of crown clades, but not amongthem. The second analysis utilizes discrete data from postnatalskeletal development, such as the presence/absence of sesamoidsand the number of secondary centres in epiphyseal cartilages. Thesediscrete data appear capable of recovering the deeper divergenceswithin Squamata, but evolve too slowly to be informative at thelevel of crown clades. Thus, patterns of postnatal skeletal developmenthave the potential to help illuminate relationships throughout thesquamate tree. Further progress in this area will require the examinationof additional squamate species, the exploration of alternative codingschemes for developmental sequences, and comparable postnatal datafor Sphenodon .  © 2002 The Linnean Societyof London, Zoological Journal of the Linnean Society, 2002, 136 ,277−313.     

The performance of morphological characters in broad-scale phylogenetic analyses

In view of their propositional content (i.e. they can be right or wrong), character statements (i.e. statements that predicate characters of organisms) are treated as low-level hypotheses. The thesis of the present study is that such character statements, as do more complex scientific theories, come with variable scope. The scope of a hypothesis, or theory, is the domain of discourse over which the hypothesis, or theory, ranges. A character statement is initially introduced within the context of a certain domain of discourse that is defined by the scale of the initial phylogenetic analysis. The doctrine of 'total evidence' requires the inclusion of previously introduced characters in subsequent studies. As a consequence, the initial scope of character statements is widened to the extent that the scale of subsequent analyses is broadened. Scope expansion for character statements may result in incomplete characters, in the subdivision of characters, or in ambiguity of reference (indeterminacy of the extension of anatomical terms). Character statements with a wide scope are desirable because they refer to characters with the potential to resolve deep nodes in phylogenetic analyses. Care must be taken to preserve referential unambiguity of anatomical terms if the originally restricted scope of a character statement is expanded to match a broad-scale phylogenetic analysis.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 297–308.     

Molecular phylogenetic diversity in the widespread lizard Cercosaura ocellata (Reptilia: Gymnophthalmidae) in South America

For a long time, Cercosaura ocellata was considered polytypic, with three subspecies: C. ocellata ocellata, C. ocellata petersi and C. ocellata bassleri. Recently, C. ocellata bassleri was elevated to full species, based on analysis of a few molecular samples from Peru. This species complex is widely distributed in South America, occurring in Amazonia, Cerrado, Atlantic forest and Pampa biomes. The monophyly and species diversity of C. ocellata are still unstudied. Here, we infer phylogenetic relationships and species diversity of this group analysing 2326 base pairs of three mitochondrial (12S, 16S, and ND4) and one nuclear (c-mos) genes. Our taxon sampling of 115 specimens includes 72 samples of C. ocellata and sequences of other Cercosaura species and closely related Cercosaurinae. Maximum likelihood phylogenetic analysis recovered the monophyly of Cercosaura and that of C. ocellata with strong support. Our analyses suggest that C. ocellata is a complex of cryptic species, which possibly started diversifying in Amazonia.     

Amphibians and squamates from the middle Eocene of Namibia,with comments on pre-Miocene anurans from Africa

Three middle Eocene localities (Silica North, Silica South, Black Crow) recently discovered in Namibia have produced terrestrial faunas that rank among the few known from the period of insulation of Africa (Aptian-early Miocene). Collectively, the three localities have yielded anuran amphibians (one pipid frog, the earliest assemblage [three taxa] of ranoid frogs in Africa, one indeterminate family) and squamate reptiles (an amphisbaenian ‘lizard’, a snake that likely represents a colubroid, and two indeterminate ‘lizards’). These Eocene faunas suggest that ranoids, colubroids and African pipids are autochthonous to Africa. However, whereas pipids are vicariants inherited from West Gondwana, ranoids and colubroids (if really autochthonous) originated in Africa from unknown stems. Silica North and Silica South correspond to aquatic environments, permanent fresh water being present in the first locality; the environment of Black Crow was drier.     

Ecomorphological variation and sexual dimorphism in a recent radiation of dwarf chameleons (Bradypodion)

Natural selection tends to favour optimal phenotypes either through directional or stabilizing selection; however, phenotypic variation in natural populations is common and arises from a combination of biotic and abiotic interactions. In these instances, rare phenotypes may possess a fitness advantage over the more common phenotypes in particular environments, which can lead to adaptation and ecological speciation. A recently radiated clade of dwarf chameleons (Bradypodion) restricted to southern KwaZulu‐Natal Province, South Africa, is currently comprised of two species (Bradypodion melanocephalum and Bradypodion thamnobates), yet three other phenotypic forms exist, possibly indicating the clade is far more speciose. Very little genetic differentiation exists between these five phenotypic forms; however, all are allopatric in distribution, occupy different habitats and vary in overall size and coloration, which may indicate that these forms are adapting to their local environments and possibly undergoing ecological speciation. To test this, we collected morphometric and habitat data from each form and examined whether ecological relevant morphological differences exist between them that reflect their differential habitat use. Sexual dimorphism was detected in four of the five forms. Yet, the degree and number of dimorphic characters was different between them, with size‐adjusted male‐biased dimorphism being much more pronounced in B. thamnobates. Habitat differences also existed between sexes, with males occupying higher perches in more closed canopy (forested) habitats than females. Clear morphological distinctions were detected between four of the five forms, with the head explaining the vast majority of the variation. Chameleons occupying forested habitats tended to possess proportionally larger heads and feet but shorter limbs than those in open canopy habitats (i.e. grassland). These results show that this species complex of Bradypodion is morphologically variable for traits that are ecologically relevant for chameleons, and that the variation among the five phenotypic forms is associated with habitat type, suggesting that this species complex is in the early stages of ecological speciation. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 113–130.     

Ecology and Behaviour of Burton's Legless Lizard(Lialis burtonis,Pygopodidae) in Tropical Australia

The elongate, functionally limbless flap-footed lizards(family Pygopodidae) are found throughout Australia, ranging into southern New Guinea. Despite their diversity and abundance in most Australian ecosystems, pygopodids have attracted little scientific study. An intensive ecological study of one pygopodid, Burton's legless lizard(Lialis burtonis Gray 1835), was conducted in Australia's tropical Northern Territory. L. burtonis eats nothing but other lizards, primarily skinks, and appears to feed relatively infrequently(only 20.8% of stomachs contained prey). Ovulation and mating occur chiefly in the late dry-season(beginning around September), and most egg-laying takes place in the early to middle wet-season(November–January). Females can lay multiple clutches per year, some of which may be fertilised with stored sperm. Free-ranging L. burtonis are sedentary ambush foragers, with radio-tracked lizards moving on average 5 m/day. Most foraging is done diurnally, but lizards may be active at any time of day or night. Radiotracked lizards were usually found in leaf-litter microhabitats, a preference that was also evident in habitat-choice experiments using field enclosures. Lizards typically buried themselves in 6–8 cm of litter; at this depth, they detect potential prey items while staying hidden from predators and prey and avoiding lethally high temperatures.     

A New Species of the Genus Protobothrops(Squamata: Viperidae) from Southern Tibet,China and Sikkim,India

A new species of the genus Protobothrops Hoge Romano-Hoge, 1983, was described from Jilong County, southern Tibet, China, and Chungthang, northern Sikkim, India. It differs from congeners by the following characters: 1) relatively large body size(total length up to 1510 mm); 2) dorsal scale rows 25–25–19; 3) except for the smooth outermost row, dorsal scales are weakly keeled; 4) relatively high number of ventral(198–216) and subcaudal(65–76 pairs) scales; 5) 7–8 supralabials; 6) 11 to 13 infralabials; 7) dorsal head uniform dark brown, laterally a reddish-brown obscure postocular streak; 8) dorsum of trunk and tail olive, with distinct black edged red brown transverse bands across the body and tail; and 9) eye from bright brown and reddish brown to mildly brown. The new species was also observed from the Haa Valley in western Bhutan.     

Fossil lizard from central Europe resolves the origin of large body size and herbivory in giant Canary Island lacertids

The endemic Canary Island lizard clade Gallotia, which includes the largest members of Europe's dominant reptile group, Lacertidae, is one of the classic examples of insular gigantism. For the first time we use fossil data to test the evolutionary reasons for the association between gigantism and herbivory. We describe an almost completely preserved skeleton of J anosikia ulmensis comb. nov. from the early Miocene of Ulm, Germany (MN 2a, ～ 22 Mya). We show that this species and Oligocene Pseudeumeces cadurcensis (Filhol, 1877) are in fact crown lacertids, and the first known pre‐Quaternary record of the total clade of Gallotia. Pseudeumeces confirms the early origin of crown Lacertidae in the Palaeogene of Europe. More importantly, these fossil taxa show that large body size was already achieved on the European mainland by the early Miocene. Furthermore, Pseudeumeces and Janosikia were faunivorous, thus demonstrating that insularity, not large body size, was crucial to the evolution of herbivory in this lineage. Body size change in Gallotia was more complex than previously thought, encompassing size increase [e.g. in the extinct Gallotia goliath (Mertens, 1942)], but more commonly involving miniaturization. The physical environment may play a crucial role in modulating the evolution of body size in this natural laboratory.     

Anatomical study of the skull of amphisbaenian Diplometopon zarudnyi (Squamata,Amphisbaenia)

This study investigates the amphisbaenian species skull which includes cranium, lower jaw and hyoid apparatus. The medial dorsal bones comprise the premaxilla, nasal, frontal and parietal. The premaxilla carries a large medial tooth and two lateral ones. The nasals are paired bones and separated by longitudinal suture. Bones of circumorbital series are frontal, orbitosphenoid and maxilla. The occipital ring consists of basioccipital, supraoccipital and exooccipital. Supraoccipital and basioccipital are single bones while the exo-occipitals are paired. The bones of the palate comprise premaxilla, maxilla, septomaxilla, palatine, pterygoid, ectopterygoid, basisphenoid, parasphenoid, orbitosphenoid and laterosphenoid. Prevomer and pterygoid teeth are absent. Palatine represent by two separate bones. The temporal bones are clearly visible. The lower jaw consists of the dentary, articular, coronoid, supra-angular, angular and splenial. The hyoid apparatus is represented by a Y-shaped structure. The mandible is long and is suspended from the braincase via relatively short quadrate. There is an extensive contact between the long angular and the large triangular coronoid. Thus inter-mandibular joint is bridged completely by the angular and consequently, the lower jaws are relatively rigid and kinetic. The maxillae are suspended from the braincase largely by ligaments and muscles rather than through bony articulation. In conclusion, the skull shape affects feeding strategy in Diplometopon zarudnyi. The prey is ingested and transported via a rapid maxillary raking mechanism.