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1.
We report on the first detailed study of the atlas–axis complex in the lizard clade Dibamidae, a family of poorly known fossorial squamates distributed in tropical or subtropical climates. This skeletal bridge is characterized by several features, such as the complete absence of the first intercentrum or the appearance of the first free cervical rib on the axis (usually less developed in Dibamus relative to that in Anelytropsis). Our study shows morphological differences of the atlas–axis complex in the Mexican blind lizard Anelytropsis relative to those of Asian Dibamus, the only two known extant genera of this clade. With regard to taxonomy and phylogenetic topology of the Dibamidae within Squamata, a huge conflict exists between morphology versus molecules. The morphology of the atlas–axis complex is therefore compared with several potential sister clades + Sphenodon. Dibamids share several features with limbless Gekkota, Scincoidea, and Amphisbaenia. The complete absence of the first intercentrum is observed in Rhineura floridana and in Ateuchosaurus chinensis as well, and the free rib associated with the synapophyses of the axis is also present in Acontias meleagris. However, some of these features may result from a limbless, burrowing ecology and thus could represent homoplastic characters. In any case, the morphology of the atlas–axis shows that dibamids share most character states with skinks. Although the atlas–axis complex forms only an additional source of information, this conclusion is consistent with most morphological rather than molecular tree topologies.  相似文献   

2.
I staged replicate encounters between unrestrained lizards andsnakes in outdoor enclosures to examine size-dependent predationwithin the common garden skink (Lampropholis guichenoti). Yellow-facedwhip snakes (Demansia psammophis) forage widely for activeprey and most often consumed large skinks, whereas death adders(Acanthophis antarcticus) ambush active prey and most oftenconsumed small skinks. Small-eyed snakes (Rhinoplocephalusnigrescens) forage widely for inactive prey and consumed bothsmall and large skinks equally often. Differential predationmay reflect active choice by the predator, differential preyvulnerability, or both. To test for active choice, I presentedforaging snakes with an inert small lizard versus an inertlarge lizard. They did not actively select lizards of a particularbody size. To test for differential prey vulnerability, I quantifiedvariation between small and large lizards in behavior thatis important for determining the outcome of predator—prey interactions. Snakes did not differentiate between integumentarychemicals from small and large lizards. Large lizards tendto flee from approaching predators, thereby eliciting attackby the visually oriented whip snakes. Small lizards were moremobile than large lizards and therefore more likely to passby sedentary death adders. Additionally, small skinks were more effectively lured by this sit-and-wait species and less likelyto avoid its first capture attempt. In contrast, overnightretreat site selection (not body size) determined a lizard'schances of being detected by small-eyed snakes. Patterns ofsize-dependent predation by elapid snakes may arise not becauseof active choice but as a function of species-specific predatortactics and prey behavior.  相似文献   

3.
Summary The structure of the sound-transmitting apparatus in primitive snakes (Scolecophidia, Henophidia) is reviewed and compared with that of advanced snakes (Caenophidia) and of some fossorial lizards. Assuming a constant course of the chorda tympani, the ophidian stylohyal can be homologized with the intercalary cartilage of lizards while the cartilaginous distal portion of the ophidian stapes represents the internal process. The cladistic significance of the stapes-quadrate-articulation in the Henophidia and Caenophidia is discussed. The Booidea and the Caenophidia show a shift of the stapes-quadrate-articulation which is correlated with changes in the suspensorium as an adaptation to relatively larger prey. However, convergence cannot be ruled out. Dibamus is shown to be the only lizard known so far which approaches the ophidian middle ear structure. Convergence has to be assumed since there is no sign of a stylohyal in Dibamus and since the course of the ramus communicans externus n. facialis cum glossopharyngeo supports the hypothesis that snakes are to be derived from a pre-lacertilian stage of lepidosaurian evolution.
Zusammenfassung Die Struktur des schalleitenden Apparates im Mittelohr primitiver Schlangen (Scolecophidia, Henophidia) wird beschrieben und mit höheren Schlangen (Caenophidia) sowie mit einigen grabenden Lacertiliern verglichen. Unter der Annahme eines konstanten Verlaufes der Chorda tympani lässt sich das Stylohyale der Schlangen mit dem Processus dorsalis homologisieren, während das knorpelige distale Ende des Stapes der Schlangen dem Processus internus entspricht. Die kladistischen Implikationen der Struktur des Mittelohres werden diskutiert. Die Booidea und die Caenophidia zeigen eine Verschiebung das Stapes-Quadratum-Gelenkes, welche möglicherweise mit Änderungen der Proportionen des Suspensoriums als Anpassung an relativ grössere Beute zusammenhängt. Konvergenz kann hierbei nicht ausgeschlossen werden. Dibamus ist die einzige bislang bekannte Echse welche im Bau des Mittelohres den Schlangen nahe kommt. Allerdings ist Konvergenz anzunehmen, da Dibamus keinerlei Hinweis auf das Vorhandensein eines Stylohyale liefert, und da der Verlauf des Ramus communicans externus n. facialis cum glossopharyngeo die Hypothese stützt, dass Schlangen nicht von rezenten Unterordnungen der Lacertilier abgeleitet werden können.

Abbreviations (used in the figures) chr chorda tympani - ci carotis interior - crst cartilaginous distal extension of the stapedial shaft - dper ductus perilymphaticus - fc carotis facialis - par articulatory process of quadrate - pt pterygoid - q quadrate - rce ramus communicans externus - rci ramus communicans internus - st supratemporal - stp stapes - sty stylohyal - vcl vena capitis lateralis - lX n. glossopharyngeus - Vllhy ramus hyomandibularis of facial nerve  相似文献   

4.
The affinities of three problematic groups of elongate, burrowing reptiles (amphisbaenians, dibamids and snakes) are reassessed through a phylogenetic analysis of all the major groups of squamates, including the important fossil taxa Sineoamphisbaena, mosasauroids and Pachyrhachis; 230 phylogenetically informative osteological characters were evaluated in 22 taxa. Snakes (including Pachyrhachis) are anguimorphs, being related firstly to large marine mosasauroids, and secondly to monitor lizards (varanids). Scincids and cordylids are not related to lacertiforms as previously thought, but to anguimorphs. Amphisbaenians and dibamids are closely related, and Sineoamphisbaena is the sister group to this clade. The amphisbaenian-dibamid-Sineoamphisbaena clade, in turn, is related to gekkotans and xantusiids. When the fossil taxa are ignored, snakes, amphisbaenians and dibamids form an apparently well-corroborated clade nested within anguimorphs. However, nearly all of the characters supporting this arrangement are correlated with head-first burrowing (miniaturization, cranial consolidation, body elongation, limb reduction), and invariably co-occur in other tetrapods with similar habits. These characters are potentially very misleading because of their sheer number and because they largely represent reductions or losses. It takes very drastic downweighting of these linked characters to alter tree topology: if fossils are excluded from the analysis, a (probably spurious) clade consisting of elongate, fossorial taxa almost always results. These results underscore the importance of including all relevant taxa in phylogenetic analyses. Inferring squamate phylogeny depends critically on the inclusion of certain (fossil) taxa with combinations of character states that demonstrate convergent evolution of the elongate, fossorial ecomorph in amphisbaenians and dibamids, and in snakes. In the all-taxon analysis, the position of snakes within anguimorphs is more strongly-corroborated than the association of amphisbaenians and dibamids with gekkotans. When the critical fossil taxa are deleted, snakes ‘attract’ the amphisbaenian-dibamid clade on the basis of a suite of correlated characters. While snakes remain anchored in anguimorphs, the amphisbaenian-dibamid clade moves away from gekkotans to join them. Regardless of the varying positions of the three elongate burrowing taxa, the interrelationships between the remaining limbed squamates (‘lizards’) are constant; thus, the heterodox affinities of scincids, cordylids, and xantusiids identified in this analysis appear to be robust. Finally, the position of Pachyrhachis as a basal snake rather than (as recently suggested) a derived snake is supported on both phylogenetic and evolutionary grounds.  相似文献   

5.
Abstract Squamate autapomorphies seen in sperm of the Scincidae (e.g. Ctenotus robustus, Carlia pectoralis, Cryptoblepharus virgatus, and Lampropholis delicata) are penetration of the fibrous sheath of the axoneme into the midpiece, and the paracrystalline subacrosomal cone. Sphenomorphus group spermatozoa (e.g. Ctenotus) and the Egernia group (Tiliqua) differ from the more derived Eugongylus group (C. virgatus, L. delicata and C. pectoralis) in that the acrosome is elongate and apically depressed; the perforatorium is strongly oblique; the midpiece is relatively short, with four dense ring structures in longitudinal succession; mitochondria are columnar; and enlarged peripheral fibres 3 and 8 do not show the gross anterior enlargement seen in Carlia and Lampropholis. Heteronotia binoei (Gekkonidae) sperm have no epinuclear electron-lucent region; nuclear shoulders are smooth, as in sphenomorph but not Eugongylus group skinks; mitochondria are columnar; unlike skinks, the median surfaces of the mitochondria are indented by triangular, sometimes longitudinally, interconnected dense bodies. In Lialis burtonis (Pygopodidae) sperm, the perforatorium extends virtually to the tip of the fore-shortened apically domed acrosome; nuclear shoulders are absent; the mitochondria alternate singly or in groups with one or more dense bodies which also form an interrupted collar around the distal centriole. Spermatozoal ultrastructure suggests that a common ancestry of snakes and pygopods deserves consideration.  相似文献   

6.
7.
Information from lizard lineages that have evolved a highly elongate (snake‐like) body form may clarify the selective forces important in the early evolution of snakes. Lizards have evolved bodily elongation via two distinct routes: as an adaptation to burrowing underground or to rapid locomotion above ground. These two routes involve diametrically opposite modifications to the body plan. Burrowing lizards have elongate trunks, small heads, short tails, and relatively constant body widths, whereas surface‐active taxa typically have shorter trunks, wider heads, longer tails, and more variable body widths. Snakes resemble burrowing rather than surface‐active (or aquatic) lizards in these respects, suggesting that snakes evolved from burrowing lizards. The trunk elongation of burrowing lizards increases the volume of the alimentary tract, so that an ability to ingest large meals (albeit consisting of small individual prey items) was present in the earliest snakes. Subsequent shifts to ingestion of wide‐bodied prey came later, after selection dismantled other gape‐constraining morphological attributes, some of which may also have arisen as adaptations to burrowing through hard soil (e.g. relatively small heads, rigid skulls). Adaptations of snake skulls to facilitate ingestion of large prey have evolved to compensate for the reduction of relative head size accompanying bodily elongation; relative to predator body mass, maximum sizes of prey taken by snakes may not be much larger than those of many lizards. This adaptive scenario suggests novel functional links between traits, and a series of testable predictions about the relationships between squamate morphology, habitat, and trophic ecology. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 293–304.  相似文献   

8.
The concept of ecomorphs, whereby species with similar ecologies have similar phenotypes regardless of their phylogenetic relatedness, is often central to discussions regarding the relationship between ecology and phenotype. However, some aspects of the concept have been questioned, and sometimes species have been grouped as ecomorphs based on phenotypic similarity without demonstrating ecological similarity. Within snakes, similar head shapes have convergently evolved in species living in comparable environments and/or with similar diets. Therefore, ecomorphs could exist in some snake lineages, but this assertion has rarely been tested for a wide-ranging group within a single framework. Natricine snakes (Natricinae) are ecomorphologically diverse and currently distributed in Asia, Africa, Europe and north-central America. They are primarily semiaquatic or ground-dwelling terrestrial snakes, but some are aquatic, burrowing or aquatic and burrowing in habit and may be generalist or specialist in diet. Thus, natricines present an interesting system to test whether snakes from different major habit categories represent ecomorphs. We quantify morphological similarity and disparity in head shape among 191 of the ca. 250 currently recognized natricine species and apply phylogenetic comparative methods to test for convergence. Natricine head shape is largely correlated with habit, but in some burrowers is better explained by dietary specialism. Convergence in head shape is especially strong for aquatic burrowing, semiaquatic and terrestrial ecomorphs and less strong for aquatic and burrowing ecomorphs. The ecomorph concept is useful for understanding natricine diversity and evolution, though would benefit from further refinement, especially for aquatic and burrowing taxa.  相似文献   

9.
《Journal of morphology》2017,278(12):1706-1715
Of the few elongate, fossorial vertebrates that have been examined for their burrowing mechanics, all were found to use an akinetic, reinforced skull to push into the soil, powered mostly by trunk muscles. Reinforced skulls were considered essential for head‐first burrowing. In contrast, I found that the skull of the fossorial shield‐nosed cobra (Aspidelaps scutatus ) is not reinforced and retains the kinetic potential typical of many non‐fossorial snakes. Aspidelaps scutatus burrows using a greatly enlarged rostral scale that is attached to a kinetic snout that is independently mobile with respect to the rest of the skull. Two mechanisms of burrowing are used: (1) anteriorly directed head thrusts from a loosely bent body that is anchored against the walls of the tunnel by friction, and (2) side‐to‐side shovelling using the head and rostral scale. The premaxilla, to which the rostral scale is attached, lacks any direct muscle attachments. Rostral scale movements are powered by, first, retractions of the palato‐pterygoid bar, mediated by a ligament that connects the anterior end of the palatine to the transverse process of the premaxilla and, second, by contraction of a previously undescribed muscle slip of the m. retractor pterygoidei that inserts on the skin at the edge of the rostral scale. In derived snakes, palatomaxillary movements are highly conserved and power prey capture and transport behaviors. Aspidelaps scutatus has co‐opted those mechanisms for the unrelated function of burrowing without compromising the original feeding functions, showing the potential for evolution of functional innovations in highly conserved systems.  相似文献   

10.
Ecosystem engineers such as burrowing seabirds can increase habitat availability for sympatric taxa – but only if the burrow's owner allows other species to use the newly created shelter site. Our studies on a small Pacific island suggest that an avian burrower (the wedge‐tailed shearwater Puffinus pacificus) is both a facilitator and a competitor for amphibious seasnakes. Video camera inspection of 102 burrows revealed frequent usage of these burrows as retreat sites by the snakes, with Laticauda laticaudata restricted to burrows <4 m from the water's edge, whereas Laticauda saintgironsi often used burrows further inland. Snakes never occupied burrows that contained adult shearwaters, suggesting active burrow defence by the birds. Model snakes that we inserted into burrows were attacked, especially on the head and upper body, and we found one snake pecked to death outside a burrow. Wedge‐tailed shearwaters act as facilitators, creating a thermally favourable microhabitat and substantially enhancing habitat suitability for snakes; but they are also competitors, aggressively competing with snakes for occupancy of the resource that has been created.  相似文献   

11.
12.
Differentiated sex chromosomes are believed to be evolutionarily stable, while poorly differentiated sex chromosomes are considered to be prone to turnovers. With around 1700 currently known species forming ca 15% of reptile species diversity, skinks (family Scincidae) are a very diverse group of squamates known for their large ecological and morphological variability. Skinks generally have poorly differentiated and cytogenetically indistinguishable sex chromosomes, and their sex determination was suggested to be highly variable. Here, we determined X-linked genes in the common sandfish (Scincus scincus) and demonstrate that skinks have shared the same homologous XX/XY sex chromosomes across their wide phylogenetic spectrum for at least 85 million years, approaching the age of the highly differentiated ZZ/ZW sex chromosomes of birds and advanced snakes. Skinks thus demonstrate that even poorly differentiated sex chromosomes can be evolutionarily stable. The conservation of sex chromosomes across skinks allows us to introduce the first molecular sexing method widely applicable in this group.  相似文献   

13.
Death adders (genus Acanthophis) differ from most other elapid snakes, and resemble many viperid snakes, in their thickset morphology and ambush foraging mode. Although these snakes are widely distributed through Australia and Papua New Guinea, their basic biology remains poorly known. We report morphological and ecological data based upon dissection of >750 museum specimens drawn from most of the range of the genus. Female death adders grow larger than conspecific males, to about the same extent in all taxa (20% in mean adult snout-vent length,  =  SVL). Most museum specimens were adult rather than juvenile animals, and adult males outnumbered females in all taxa except A. pyrrhus. Females have shorter tails (relative to SVL) than males, and longer narrower heads (relative to head length) in some but not all species. The southern A. antarcticus is wider-bodied (relative to SVL) than the other Australian species. Fecundity of these viviparous snakes was similar among taxa (mean litter sizes 8 to 14). Death adders encompass a broad range of ecological attributes, taking a wide variety of vertebrate prey, mostly lizards (55%), frogs and mammals (each 21%; based on 217 records). Dietary composition differed among species (e.g. frogs were more common in tropical than temperate-zone species), and shifted with snake body size (endotherms were taken by larger snakes) and sex (male death adders took more lizards than did females). Overall, death adders take a broader array of prey types, including active fast-moving taxa such as endotherms and large diurnal skinks, than do most other Australian elapids of similar body sizes. Ambush foraging is the key to capturing such elusive prey.  相似文献   

14.
15.
The venom glands and related muscles of sea snakes conform in their general structure to those of the terrestrial elapids. The venom gland, however, is smaller in size and the accessory gland is considerably reduced. A similar pattern is found in the Australian elapid Notechis. The musculus compressor glandulae is well developed in the sea snakes and in some species its posterior-medial portion runs uninterruptedly from the origin to the insertion of the muscle. This might be considered as a primitive condition suggesting an early divergence of the sea snakes from an ancestral elapid stock. Three species of sea snakes, Aipysurus eydouxi, Emydocephalus annulatus, and E. ijimae, feed on fish eggs and have very small, but still functioning, venom glands. The reduced accessory gland of the sea snakes is apparently connected with their aquatic environment, as a similar condition is found also in the elapine Boulengerina annulata which lives in large lakes of Central Africa. The similarity in structure of the venom gland between sea snakes and Notechis scutatus may point to a possible phylogenetic relationship between this group of Australian elapids and hydrophiine snakes.  相似文献   

16.
The present distribution of lizards is usually explained as a result of relatively recent global events, i.e. faunal turnovers or exchanges within and between particular continents mostly connected with glaciations and land‐bridges. However, today's disjunct distribution of the North American Xantusiidae and African Cordyliformes (close relatives of skinks) does not fit generally accepted biogeographical patterns. A new, exquisitely preserved specimen of the Late Jurassic lizard Ardeosaurus brevipes from the Solnhofen area, Germany sheds some light on the problem. A posterior projection of its parietal is known only in the Dibamidae, Gekkota and Scincoidea, taxa representing first branches on the molecular tree of lizards. The projection of the parietal is proposed to be an apomorphy of the Squamata that was lost in the common ancestor of Lacertoidea and Toxicofera (snakes, anguimorphs and iguanians). This implies a basalmost position of A. brevipes on the Squamata tree. The location of the supratemporal such as in A. brevipes characterizes all the Scincoidea and some geckos, but most geckos lack osteoderms, which are common in Scincoidea. This makes A. brevipes the oldest known crown‐member of this lineage. Moreover, this indicates that the main groups of modern lizards were already present in the Jurassic. Ancestors of Cordyliformes migrated to Gondwana in the Late Jurassic together with dinosaurs, prior to the Cretaceous separation of this ancient continent.  相似文献   

17.
Understanding the origin and maintenance of functionally important subordinate traits is a major goal of evolutionary physiologists and ecomorphologists. Within the confines of a limbless body plan, snakes are diverse in terms of body size and ecology, but we know little about the functional traits that underlie this diversity. We used a phylogenetically diverse group of 131 snake species to examine associations between habitat use, sidewinding locomotion and constriction behaviour with the number of body vertebrae spanned by a single segment of the spinalis muscle, with total numbers of body vertebrae used as a covariate in statistical analyses. We compared models with combinations of these predictors to determine which best fit the data among all species and for the advanced snakes only (N = 114). We used both ordinary least‐squares models and phylogenetic models in which the residuals were modelled as evolving by the Ornstein–Uhlenbeck process. Snakes with greater numbers of vertebrae tended to have spinalis muscles that spanned more vertebrae. Habitat effects dominated models for analyses of all species and advanced snakes only, with the spinalis length spanning more vertebrae in arboreal species and fewer vertebrae in aquatic and burrowing species. Sidewinding specialists had shorter muscle lengths than nonspecialists. The relationship between prey constriction and spinalis length was less clear. Differences among clades were also strong when considering all species, but not for advanced snakes alone. Overall, these results suggest that muscle morphology may have played a key role in the adaptive radiation of snakes.  相似文献   

18.
Most lizards display relatively simple social systems, but more complex and stable social aggregations appear to be common in one lineage of Australian skinks, the Egernia Group. Previous studies on this lineage have focused on species inhabiting crevices in large and disjunct rocky outcrops. Here, we describe the social organization of White's skink, Egernia whitii, a burrowing species that inhabits rocky habitats in southeastern Australia. We examined social group size, composition and stability over two field seasons using a capture‐mark‐recapture study, behavioral observations and genetic analyses. Twenty‐four social groups, each comprising two to six individuals, were present at our study site, with 75% of lizards belonging to a social grouping. A higher proportion of adults than juveniles were part of a group, while more adult females belonged to a group than adult males. Groups generally comprised a single adult pair or an adult pair with juveniles. However, groups comprising one male with multiple females and multiple individuals of both sexes also were present. Groups were highly stable throughout the study, although individual group members were observed singly on half of all observations. Paternity analysis using four microsatellite loci revealed that juveniles within groups were closely related to adults in the group, with 38% living in groups with both parents and 71% in groups with at least one parent. Our data demonstrate the presence of complex sociality in a burrowing Egernia species and, together with previous studies, suggest that stable social organization is widespread across different habitats and phylogenetic groupings within the Egernia Group.  相似文献   

19.
Relationships between the major lineages of snakes are assessed based on a phylogenetic analysis of the most extensive phenotypic data set to date (212 osteological, 48 soft anatomical, and three ecological characters). The marine, limbed Cretaceous snakes Pachyrhachis and Haasiophis emerge as the most primitive snakes: characters proposed to unite them with advanced snakes (macrostomatans) are based on unlikely interpretations of contentious elements or are highly variable within snakes. Other basal snakes include madtsoiids and Dinilysia--both large, presumably non-burrowing forms. The inferred relationships within extant snakes are broadly similar to currently accepted views, with scolecophidians (blindsnakes) being the most basal living forms, followed by anilioids (pipesnakes), booids and booid-like groups, acrochordids (filesnakes), and finally colubroids. Important new conclusions include strong support for the monophyly of large constricting snakes (erycines, boines. pythonines), and moderate support for the non-monophyly of the trophidophiids' (dwarf boas). These phylogenetic results are obtained whether varanoid lizards, or amphisbaenians and dibamids, are assumed to be the nearest relatives (outgroups) of snakes, and whether multistate characters are treated as ordered or unordered. Identification of large marine forms, and large surface-active terrestrial forms, as the most primitive snakes contradicts with the widespread view that snakes arose via minute, burrowing ancestors. Furthermore, these basal fossil snakes all have long flexible jaw elements adapted for ingesting large prey ('macrostomy'), suggesting that large gape was primitive for snakes and secondarily reduced in the most basal living foms (scolecophidians and anilioids) in connection with burrowing. This challenges the widespread view that snake evolution has involved progressive, directional elaboration of the jaw apparatus to feed on larger prey.  相似文献   

20.
Faecal carriage of Pseudomonas aeruginosa was investigated by selective plating and PCR identification test, among healthy captive snakes from zoological and private collections from France as well as from wild snakes from Guinea. P. aeruginosa faecal carriage among captive snakes was high (72 out of 83 individuals), but low among wild specimen (3 out of 23 individuals). Genetic diversity analyses of the isolates, based on SpeI‐PFGE profiles, evidenced five dominant clones or clonal complexes spreading among snakes within a site and between sites and persisting over time. Similar clones or clonal complexes were detected from mouth swabs of the owners and from water and preys used to feed the snakes, evidencing various sources of snake colonization and the first cases of P. aeruginosa cross‐contamination between snakes and owners. These observations led to the conclusion that P. aeruginosa behaves as an opportunistic species within snakes in captivity and that colonization and dissemination occurs consecutively to processes similar to those identified within the hospital. Antibiotic susceptibility testing showed that most isolates had a wild‐type resistance profile except for one persistent clone isolated from both snakes and preys that harboured multiple antimicrobial resistance genes mediated by an integron carrying the qacH, aadB, aadA2 and cmlA10 cassettes, and a tetA(C)‐carrying transposon. Biocides or antibiotics used in the zoological garden could have led to the acquisition of this integron.  相似文献   

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