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1.
The interrelationships within the clade comprised of turtles, pareiasaurs, and procolophonid-like taxa are investigated via a cladistic analysis incorporating 56 characters. A single most parsimonious tree was found (80 steps, c. i. = 0·8) in which the successive outgroups to turtles are: pareiasaurs, Sclerosaurus, lanthanosuchids, procolophonoids (=Owenetta, Barasaurus and procolophonids), and nyctiphruretians (= nycteroleterids). Thus, as suggested recently by other workers (Reisz, in Fischman, 1993) turtles are the highly modified survivors of a radiation of poorly-known reptiles commonly called ‘parareptiles’. Pareiasaurs are united with turtles on the basis of twenty unambiguous derived features which are absent in other basal amniotes (=‘primitive reptiles’) and reptiliomorph amphibians: for example, the medially located choana, enlarged foramina palatinum posterius, blunt cultriform process, fully ossified medial wall of the prootic, opisthotic-squamosal suture, lateral flange of exoccipital, loss of ventral cranial fissure, thickened braincase floor, ‘pleurosphenoid’ ossification, reduced presacral count, acromion process, trochanter major, reduced fifth pedal digit, and presence of transverse processes on most caudals. Recent phylogenetic proposals linking turtles with captorhinids, with dicynodonts, and with procolophonoids are evaluated. None of the proposed traits supporting the first two hypotheses is compelling. The procolophonoid hypotheses is supported by only one synapomorphy (the slender stapes). All other synapomorphies proposed in favour of the above groupings either occur in many other primitive amniotes, or are not primitive for turtles, or are not primitive for the proposed chelonian sister-group. Nyctiphruretus and Lanthanosuchids and nycteroleterids, often considered to be seymouriamorph amphibians, are demonstrated unequivocally to be amniotes. The ‘rhipaeosaurs’, currently considered to be pareiasaur relatives, are shown to be a heterogenous assemblage of seymouriamorphs, therapsids and nycteroleterids. The phylogeny proposed here indicates that many of the traits of the earliest known turtle, Proganochelys, previously interpreted as unique specialisations, also occur in pareiasaurs and other near outgroups of turtles, and must instead represent the primitive chelonian condition: for example, the wide parietals and the short quadrate flange of the pterygoid. The sequence of acquisition of chelonian traits is discussed: many features once thought to be diagnostic of turtles actually characterize larger groupings of procolophonomorphs, and must have evolved long before the chelonian shell appeared. These traits include most of the chelonian-pareiasaur synapomorphies listed above, and many others which characterize more inclusive groupings found in this analysis. In putting Proganochelys much closer to the main line of chelonian evolution, in elucidating the sequence of acquisition of chelonian traits, and in reducing greatly the number of differences between turtles and their nearest relatives, this study helps bridge one of the major gaps in the fossil record. The failure of previous cladistic analyses to identify correctly the nearest relatives of turtles is attributed to biased character selection, caused by an over-reliance on cranial characters deemed ‘important’ by earlier workers, and by a tendency to shoehorn ‘parareptile’ taxa into phylogenies derived from analyses restricted to ‘mainstream’ groups such as synapsids, diapsids, turtles, and ‘captorhinomorphs’. Many of the synapomorphies that resolve turtle origins are postcranial, and the three nearest outgroups to turtles are all highly bizarre groups which were dismissed as ‘too specialized’ by early workers and continued to be inadequately assessed even by workers using a cladistic framework.  相似文献   

2.
The homology of the fifth metatarsal in turtles and in diapsid reptiles is reassessed in the light of new phylogenetic studies. The two nearest outgroups to turtles — pareiasaurs and procolophonoids — both lack a fifth distal tarsal but retain a normal fifth metatarsal. The fifth distal tarsal was therefore lost at the base of the clade that contains turtles. Thus, the hooked fifth metatarsal in turtles must consist entirely of a modified fifth metatarsal: it does not include contributions from the fifth distal tarsal, as commonly supposed. In diapsids, loss of the fifth distal tarsal appears to have occurred at the base of crown-clade diapsids, hooking of the fifth metatarsal subsequently occurring within lepidosauromorphs, and in archosauromorphs. If so, the hooked fifth metatarsal in archosauromorphs, and some lepidosauromorphs, consists entirely of a modified fifth metatarsal. In both turtles and diapsids, integration of the elements distal to the mesotarsal joint precedes evolution of the hooked fifth metatarsal, supporting the view that the latter element evolved to perform a lever function (analogous to the “heel bone” of mammals).  相似文献   

3.
The neurocranial osteology of the giant monitor lizard Varanus (Megalania) prisca Owen, 1859 is described in detail for the first time. Optimization of neurocranial characters onto phylogenetic topologies for varanoids, including Lanthanotus, Heloderma, and Varanus species nests V. prisca within an Indo‐Australian clade of Varanus on the basis of characters of the otic capsule. A sister‐taxon relationship between V. prisca and Varanus komodoensis Ouwens, 1912 is proposed based on apomorphies of the crista prootica, fenestra vestibuli, occipital recess, and supraoccipital. These results support a monophyletic clade of giant monitors among Indo‐Australian species, and unambiguously synonymize Megalania with Varanus at both generic and subgeneric levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 445–457.  相似文献   

4.
Despite the attention squamate lizards have received in the study of digit and limb loss, little is known about limb morphogenesis in pentadactyl lizards. Recent developmental studies have provided a basis for understanding lizard autopodial element homology based on developmental and comparative anatomy. In addition, the composition and identity of some carpal and tarsal elements of lizard limbs, and reptiles in general, have been the theme of discussions about their homology compared to non‐squamate Lepidosauromorpha and basal Amniota. The study of additional embryonic material from different lizard families may improve our understanding of squamate limb evolution. Here, we analyze limb morphogenesis in the gekkonid lizard Gonatodes albogularis describing patterns of chondrogenesis and ossification from early stages of embryonic development to hatchlings. Our results are in general agreement with previous developmental studies, but we also show that limb development in squamates probably involves more chondrogenic elements for carpal and tarsal morphogenesis, as previously recognized on the grounds of comparative anatomy. We provide evidence for the transitory presence of distal carpale 1 and intermedium in the carpus and tibiale, intermedium, distal centralia, and distal tarsale 2 in the tarsus. Hence, we demonstrate that some elements that were believed to be lost in squamate evolution are conserved as transitory elements during limb development. However, these elements do not represent just phylogenetic burden but may be important for the morphogenesis of the lizard autopodium. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.  相似文献   

5.
We developed specific primers for microsatellite DNA regions of Podocnemis unifilis and tested their utility in population genetic and paternity studies on the species and other closely related Amazonian chelonians. Seventeen microsatellite loci were polymorphic in P. unifilis and all, plus two monomorphic microsatellites in P. unifilis, were polymorphic in at least one additional chelonian species, including Peltocephalus dumeriliana.  相似文献   

6.
7.
Vavilovia formosa is a relict, endangered species from the highlands of the Caucasus and the Near East. Described in 1812, it has had an uncertain status and was finally recognized as a separate genus of tribe Fabeae (Fabaceae). Our informal international group was established in 2007 to revive the interest in this species as it had been seriously neglected for decades. Here, we provide an overview of the accumulated knowledge on V. formosa and present the results of the most recent multidisciplinary research. Three expeditions were made to two locations in Armenia in 2009, providing the material for anatomical, morphological, chemical and molecular analysis. Unlike previous attempts, ex situ conservation in Yerevan and in vitro propagation, important for potential interspecific hybridization, were successful. Molecular tools were used to clarify the taxonomic position of V. formosa, often considered the closest to the extinct ancestor of the whole tribe. The analysis of four informative regions of plastid and nuclear DNA showed that V. formosa belongs to the same clade as Lathyrus and Pisum, with a distinct status. Preservation and maintenance of V. formosa remains the only basis for further development of all other scientific aspects, especially breeding and uses in agronomy. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 524–531.  相似文献   

8.
Although physical features of the environment are known to influence shape, to date, the effects of many environmental conditions on asymmetry remain unexamined. The present study aimed to examine the relationship between asymmetry and the fluid media (air versus water) that an organism inhabits and, subsequently, to use these findings to test a novel hypothesis that fluid environments can impose selective forces acting to constrain asymmetry to maintain or enhance biomechanical function. We examined carapace asymmetry in 114 species (69 aquatic and 45 terrestrial) from the chelonian superfamily Testudinoidea. The results obtained indicate that environment is correlated with the degree of asymmetry, but in different directions for the two clades that comprise the Testudinoidea. Within the Testuguria, aquatic turtles have lower levels of asymmetry than terrestrial turtles, which is consistent with our proposed biomechanical hypothesis. This pattern was not observed within the Emydidae, possibly due to the much shorter time that terrestrial taxa in this clade have existed. Nevertheless, the present study provides the first evidence for a relationship between fluid media and asymmetry in any taxonomic group.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 483–489.  相似文献   

9.
Twenty‐two chromosome counts are reported in 16 species, four subspecies and two varieties of the genus Centaurea. These are mostly Turkish local endemics of section Cheirolepis, a complicated group from the Eastern clade of the Jacea group. Twenty‐one reports are new. Prevalence of the basic chromosome number x = 9 among the eastern sections of the Jacea group is confirmed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 280–286.  相似文献   

10.
The cranial osteology of the small theropod dinosaur Proceratosaurus from the Bathonian of Minchinhampton, England, is described in detail, based on new preparation and computed tomography (CT) scan images of the type, and only known, specimen. Proceratosaurus is an unusual theropod with markedly enlarged external nares and a cranial crest starting at the premaxillary–nasal junction. The skull is highly pneumatic, with pneumatized nasals, jugals, and maxillae, as well as a highly pneumatic braincase, featuring basisphenoid, anterior tympanic, basipterygoid, and carotid recesses. The dentition is unusual, with small premaxillary teeth and much larger lateral teeth, with a pronounced size difference of the serrations between the mesial and distal carina. The first dentary tooth is somewhat procumbent and flexed anteriorly. Phylogenetic analysis places Proceratosaurus in the Tyrannosauroidea, in a monophyletic clade Proceratosauridae, together with the Oxfordian Chinese taxon Guanlong. The Bathonian age of Proceratosaurus extends the origin of all clades of basal coelurosaurs back into the Middle Jurassic, and provides evidence for an early, Laurasia‐wide, dispersal of the Tyrannosauroidea during the late Middle to Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009.  相似文献   

11.
We present a mtDNA analysis of Podocnemis expansa (n = 81) and Podocnemis unifilis (n = 228) turtles traded in Peru to evaluate the potential origin of these animals. In particular, we were interested in the relationship between samples reported in the Iquitos markets (IMs) and a Pacaya Samiria Natural Reserve (PSNR) where illegal hunting is presumed. Our mtDNA data showed that, for both species, all haplotypes found within the PSNR were observed in the IM, and that these markets also displayed haplotypes not documented in the reserve. This suggests that the IMs are recipients of Podocnemis turtles from within and outside the PSNR. The fact that most of the haplotype diversity observed in the markets was not found within the PSNR strongly suggests that Podocnemis genetic diversity is exploited in areas where conservation actions are limited. Hence, we recommend expanding Podocnemis conservation efforts outside of protected areas.  相似文献   

12.
The Balkan Peninsula is considered the most important refugium for species during the Pleistocene glaciations and today harbours c. 2000 endemic species, but we know surprisingly little about the evolution of taxa in this region. Veronica saturejoides, V. thessalica and V. erinoides are a group of closely related alpine taxa endemic to the Balkan Peninsula. Here, we analyse four DNA regions [the nuclear chalcone synthase intron (CHSi) and ribosomal internal transcribed spacer (ITS) region and the plastid rpoBtrnC spacer and trnLtrnLtrnF region] and amplified fragment length polymorphism (AFLP) fingerprints to provide a phylogenetic hypothesis for the relationships among these taxa. Additionally, we analyse leaf morphological characters used to distinguish the three subspecies of V. saturejoides. The analyses support the distinction of the three subspecies based on previously intuitively suggested characters. Nuclear chalcone synthase intron data indicate that the southern taxa are genetically much more diverse than the more northern V. saturejoides subsp. saturejoides. Phylogenetic relationships inferred from this region and AFLP fingerprints support the monophyly of V. saturejoides. In contrast, plastid DNA regions suggest a closer relationship of V. saturejoides subsp. saturejoides to V. thessalica. The most likely scenario involves introgression into V. saturejoides subsp. saturejoides from V. thessalica. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 616–636.  相似文献   

13.
Rensch's rule, a macroevolutionary pattern in which sexual size dimorphism (SSD) increases with body size in male‐biased SSD species, or decreases with female‐biased SSD species, has been investigated in many vertebrates because it indicates whether SSD is being driven by sexual selection or a different force (i.e. fecundity or natural selection). Evidence in turtles has shown some conflicting results, which may be explained by the different phylogenies used in the analyses. Because the newly available well‐resolved phylogeny of family Kinosternidae provides evidence for the ancient monophyly of Staurotypidae and Kinosternidae and their recognition as separate families (previously Staurotypidae was considered as a subfamily within Kinosternidae) and introduced the genus Cryptochelys for the monophyletic leucostomum clade, we revisit the pattern of SSD and body size in Kinosternidae. By contrast to what had been proposed, we found that the Kinosternidae as formerly recognized (i.e. including Staurotypus and Claudius) and the restricted Kinosternidae both follow a pattern consistent with Rensch's rule. Our analysis with published body size data did not change our results, confirming the importance of the phylogeny used in macroevolutionary studies. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 806–809.  相似文献   

14.

Candida auris, a multidrug-resistant nosocomial pathogen, has emerged globally with high morbidity and mortality among immunocompromised individuals and COVID19 hospitalized patients. Five major clades of C. auris have been previously described. The fifth clade is exclusively found in Iran where C. auris isolates are genetically distinct from other clades by?>?200,000 single-nucleotide polymorphisms. The origin of C. auris remains unclear, and limited clinical data are available at present regarding clade V infection or colonization. Herein, another case of otomycosis in Iran caused by an isolate of C. auris belonging to the fifth clade is reported. Genotyping revealed that the obtained C. auris isolate from Isfahan clustered with earlier clade V isolates from Babol, cities around 600 km separated, which indicates that C. auris clade V is established in Iran. C. auris is thought to exist more commonly in Iran, given that limited diagnostic capacity in the country has probably curbed the identification of more C. auris cases. Therefore, surveillance of the environment, patients and healthcare facilities in different geographical regions in Iran is urgently required.

  相似文献   

15.
In this contribution we present a new species of horned turtle, G affneylania auricularis gen. et sp. nov. , from the Paleogene of Patagonia. The specimens come from the lower section of the Sarmiento Formation (Middle Eocene) at Cerro Verde (Cañadón Hondo area, Province of Chubut, Argentina). The level containing turtles and crocodyliforrmes is located at the base of the section and it consists of laminated, fine tuffs interpreted as shallow pond sediments. It underlies another fossiliferous level comprising lenticular, massive sandstones bearing skeletal remains of mammals, referred by previous authors to the Casamayoran SALMA. Gaffneylania auricularis represents one of the most complete meiolaniids from South America found to date and it is distinguished from other meiolaniids by the presence of a peculiar half‐moon‐shaped, thick rim surrounding the cavum tympani, the presence of three cranial scutes K and an unenclosed canalis chorda tympani mandibularis, among others. This new species sheds new light on the evolution and palaeobiogeographical history of the clade Meiolaniidae in Australasia and South America during the Cainozoic. The break up of southern Gondwana provoked major global climatic changes during the Cainozoic that probably influenced the evolution of meiolaniid turtles. The co‐evolution of meiolaniids with other amniotes (e.g. chelid turtles, mammals) suggests a common palaeobiogeographical history of those clades in southern Gondwana. © 2015 The Linnean Society of London  相似文献   

16.
Carpal and tarsal anatomy was documented based on the observation of dry skeletons of adult specimens representing 25 species in 15 genera and on data taken from the literature. In addition, histological sections and cleared and double‐stained autopodia of recently hatched and juvenile specimens representing seven chelid and pelomedusoid species were studied. There is much more morphological diversity in the manus than in the pes. Variation in autopodial skeletons includes: the astragalus and calcaneum are either separated or fused; fusion of distal carpals 3–4−5 or just 4–5; number of centralia in the carpus; and presence/absence of a pisiform and of an accessory radial element. The widespread and probably basal phalangeal formula for Pleurodira is 2.3.3.3.3. Deviations are Pelomedusa subrufa, exhibiting a reduction to 2.2.2.2.2, Pelusios spp. with one phalanx less in digit I and for one species in digit V as well, and Acanthochelys pallidipectoris with an additional phalanx in the fourth finger. Six discrete characters itemizing some of the morphological variation observed were plotted on a composite pleurodire phylogeny, revealing not only homoplastic patterns but also the utility of some characters in supporting the monophyly of several clades. The pisiform is the last carpal element to ossify in Chelus fimbriatus. We hypothesize that the so‐called fifth hooked metatarsal represents the fusion of distal tarsal 5 with metatarsal V. The accessory radial element that was occasionally present in the turtles examined may represent an atavism of the otherwise lost radiale of turtles.  相似文献   

17.
The early Miocene sites of Moruorot and Kalodirr (Kenya, 17.5 Myr) have yielded a rich collection of mammals. New listriodont material from these localities, including a complete skull and a partial mandible, provide long awaited information on cranial features of early bunodont Listriodontinae. The evolution and systematics of the group are highly debated, especially regarding its first representatives. The new material described here sheds light on the differentiation of bunodont Listriodontinae in Africa and clarifies the systematics of the group. The first phylogenetic analysis of the Listriodontinae is here performed and supports close relationships between Kubanochoerus and a clade (Eurolistriodon, Listriodon). Lopholistriodon is the most basal representative of the listriodontine clade. These first results stress the role of the African continent in the biogeographical history of the Listriodontinae. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 653–678.  相似文献   

18.
Miniaturization, or the evolution of a dramatically reduced body size compared to related lineages, is an extraordinarily widespread phenomenon among metazoans. Evolutionary biologists have been fascinated by miniaturization because this transition has occurred numerous times, often among close relatives, providing a model system for studying convergent evolution and its underlying mechanisms. Much of the developmental work describing the ontogeny of miniature species suggests that paedomorphosis is the predominant avenue of miniaturization. Nevertheless, specific alterations to ontogeny appear highly variable, so that even related lineages with similar miniaturized traits produce those similarities via distinct ontogenetic paths. One major vertebrate group that has been overlooked in research on miniaturization is turtles. In the present study, we examined patterns of shape change in the plastron (the ventral part of the shell) over the course of ontogeny in a small clade of turtles (Emydinae) aiming to investigate whether two independently evolved diminutive members of the clade (Glyptemys muhlenbergii and Clemmys guttata) should be considered as miniaturized. We employ geometric morphometric methods to quantify the patterns of shape change these potentially miniaturized species and their relatives undergo during ontogeny, and use molecular phylogenetic trees to reconstruct ancestral conditions and provide information on the polarity of shape changes. We find that differing changes in ontogenetic parameters relative to ancestral conditions accompany the evolution of small size in emydines: G. muhlenbergii changes the duration of ontogeny and rate of shape change, whereas C. guttata changes growth rate. The observed ontogenetic repatterning of these species is reminiscent of changes in ontogeny and life history often found in miniaturized taxa. However, we conclude that C. guttata and G. muhlenbergii are not truly miniaturized because they still produce typical adult shell morphologies, and larger emydines display comparable ontogenetic flexibility. Because no emydines carry juvenile shell features forward into adulthood, we speculate that few, if any turtles, will show paedomorphic shell traits without corresponding changes in defensive strategy because such shells may offer insufficient protection. © 2013 The Linnean Society of London  相似文献   

19.
Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.  相似文献   

20.
Fishes of the order Alepocephaliformes, slickheads and tubeshoulders, constitute a group of deep‐sea fishes poorly known in respect to most areas of their biology and systematics. Morphological studies have found alepocephaliform fishes to display a mosaic of synapomorphic and symplesiomorphic characters, resulting in great difficulties when attempting to resolve intra‐ and interrelationships. Molecular data recently added to the confusion by removing Alepocephaliformes from the Euteleostei and placed them as incertae sedis within the Otocephala. In the present study we attempt to further clarify relationships of Alepocephaliformes by adding newly determined whole mitogenome sequences from 19 alepocephaliforms in order to address 1) phylogenetic position of Alepocephaliformes within the Otocephala; and 2) intrarelationships of Alepocephaliformes. The present study includes 96 taxa of which 30 are alepocephaliforms and unambiguously aligned sequences were subjected to partitioned maximum likelihood and Bayesian analyses. Results from the present study support Alepocephaliformes as a genetically distinct otocephalan order as sister clade to Ostariophysi (mostly freshwater fishes comprising Gonorynchiformes, Cypriniformes, Characiformes, Siluriformes and Gymnotiformes). The disputed family Bathylaconidae was found to be an artificial assemblage of the two genera Bathylaco and Herwigia, with the former as the sister group of the family Alepocephalidae and the latter nested within Alepocephalidae. Platytroctidae was found to be monophyletic as sister clade to the rest of Alepocephaliformes. Previously unrecognized clades within the family Alepocephalidae are presented and a clade comprising Alepocephalus, Conocara and Leptoderma was recovered as the most derived. As long as the current classification is being followed, the genera Alepocephalus, Bathytroctes, Conocara and Narcetes were all found non‐monophyletic. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 923–936.  相似文献   

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