Vertebrae of the sea snake <Emphasis Type="Italic">Palaeophis nessovi</Emphasis> Averianov (Acrochordoidea,Palaeophiidae) from the Eocene of western Kazakhstan and phylogenetic analysis of the superfamily Acrochordoidea

The vertebrae of sea snakes from five Eocene localities in western Kazakhstan are assigned to the species Palaeophis nessovi Averianov, 1997. The anterior trunk vertebrae have subcentral ridges, large posterior hypapophysis, and large synapophyses; the middle trunk vertebrae are slightly laterally compressed and their synapophyses are positioned highly; the posterior trunk vertebrae are strongly laterally compressed and have a well-developed haemal keel. Cladistic analysis has shown that the genus Palaeophis is not monophyletic; the genus Archaeophis is a more advanced palaeophiid than was previously thought.     

Additional vertebral material of Thaumastophis (Serpentes: Caenophidia) from the early Eocene of India provides new insights on the early diversification of colubroidean snakes

The Ypresian Cambay Shale Formation at Vastan, Mangrol, and Tadkeshwar lignite mines in Gujarat, western India, has yielded a rich vertebrate fauna including madtsoiid, palaeophiid, booid, and colubroidean-like snakes. The latter are particularly abundant, but their systematic affinities are difficult to resolve. Here we describe new specimens of the colubroidean-like snake Thaumastophis missiaeni, including anterior, middle, and posterior trunk vertebrae, as well as caudal vertebrae. The combination of primitive and derived caenophidian and colubroidean vertebral characters confirms Thaumastophis as the earliest known stem-colubriform snake while Procerophis, from the same beds, is more derived and considered to represent a crown-Colubriformes. Additionally, Thaumastophis shares with Renenutet enmerwer from the late Eocene of Egypt a unique combination of vertebral characters that suggests an exchange with North Africa was possible along the southern margin of the Neotethys. We erect the new family Thaumastophiidae for Thaumastophis and Renenutet on the basis of their shared derived vertebral morphology.     

A new species of Lapparentophis from the mid-Cretaceous Kem Kem beds,Morocco, with remarks on the distribution of lapparentophiid snakes

Two isolated trunk vertebrae from the ?uppermost Albian–lower Cenomanian Kem Kem beds of Morocco are described and assigned to Lapparentophis, an early snake genus known from coeval deposits in Algeria. The Moroccan specimens represent a new species, Lapparentophis ragei, which can be distinguished from the type and only known species, Lapparentophis defrennei, by its smaller size, its more elongate vertebrae, the presence of parazygosphenal foramina, and paradiapophyses extending anteroventrally closer to the cotyle. The discovery of Lapparentophis in the Kem Kem beds adds to the relatively diverse snake assemblage previously reported from this formation and extends the geographical range of the genus. The distribution of Lapparentophis and lapparentophiid-grade (?lapparentophiid) snakes is discussed. This poorly known family of terrestrial snakes seems to be restricted to the latest Albian–early Cenomanian of North Africa, with the exception of Pouitella from the early–middle Cenomanian of France. As for many other vertebrate taxa of this period, this distribution is consistent with a dispersal event from Africa to the western part of the European archipelago.     

Anatomy and relationships of Pachyrhachis problematicus,a primitive snake with hindlimbs

The anatomy of Pachyrhachis problematicus, an elongate, limb-reduced squamate from the Upper Cretaceous of Israel, is described and evaluated in detail. Previously considered a snake-like ''lizard'' of uncertain affinities, it is here shown to be the most primitive snake, and the sister-group to all other snakes. Pachyrhachis exhibits numerous derived characters uniting it with modern snakes (scolecophidians and alethinophidians): e.g. mobile premaxilla-maxilla articulation, braincase enclosed by frontals and parietals, sagittal parietal crest, absence of tympanic recess, single postdentary bone, over 140 presacral vertebrae, and complete loss of shoulder girdle and forelimb. However, it is more primitive than all modern snakes in retaining some strikingly primitive (lizard-like) features: presence of a jugal, squamosal, normal sacral attachment, and well-developed hindlimb composed of femur, tibia, fibula, and tarsals. Pachyrhachis provides additional support for the hypothesis that snakes are most closely related to Cretaceous marine lizards (mosasauroids). Almost all of the derived characters proposed to unite snakes and mosasauroids are highly developed in Pachyrhachis: the mobile mandibular symphysis, intramandibular joint, long and recurved pterygoid teeth, quadrate suspended by the supratemporal, loosely united pelvic elements (ilium, ischium, and pubis), and separate astragalus and calcaneum.     

Ecological and phylogenetic variability in the spinalis muscle of snakes

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.     

Radiation within the advanced snakes (Caenophidia) with special emphasis on African opistoglyph colubrids, based on mitochondrial sequence data

A parsimony analysis of parts of the mitochondrial genes 12S and 16S (722 base pairs) from 43 species of the advanced snakes (Caenophidia) resulted in two most parsimonious topologies. Based on a strict consensus of these the following objectives were addressed: (1) Which groupings of caenophidian taxa can be recognized? (2) Is Acrochordus sister group to, or included in, Caenophidia? (3) Are boigine snakes (sensu stricto) a monophyletic grouping and where do these taxa belong in a broader caenophidian context? (4) What are the systematic affinities of the African egg-eating genusDasypeltis ? The phylogeny was then used to discuss: (5) the evolution of the posterior maxillary dentition and the composition of the retinal visual cells. The results reveal that, when using Boa constrictor as outgroup,Acrochordus is the sistergroup to the remainder of the ingroup, and a further eight clades are defined. Five genera of elapids did not appear to be monophyletic and a number of colubrids (sensu lato) such as Mehelya,Lycodonomorphus , Lamprophis, Atractaspis, and Buhoma (formerly Geodipsas), which have traditionally been problematic to place systematically, did not group with any of the larger clades. These taxa are together with the elapids representatives of very early radiations in the evolution of the Advanced snakes. The homalopsine Enhydris enhydris appears as a sistergroup to the viperine clade (Clade 1). When plotted onto the topology the posterior maxillary dentition appear to express three, maybe four, independent origins of the Opistoglyph State. The retinal evolution also appeared very complex. The suggested very primitive placing of the Boigine snakes (sensu stricto) due to their lack of double cones in the retina of the eye was not supported here; instead the sequence data suggests this observation to be the result of a secondary loss.     

Ostéologie et affinités de Trematochampsa taqueti (Crocodylia,Mesosuchia) du Sénonien inférieur d'In Beceten (République du Niger)

Among the abundant remains of Mesosuchia that have been yielded by the early Senonian locality of In Beceten (Niger), the most numerous belong to the species Trematochampsa taquetiBuffetaut, 1974. The osteology of this species is described here. This medium-sized crocodilian, with a moderately elongated skull, is characterized, among other features, by its antorbital fenestra, its surangular-quadratojugal articulation, its slightly displaced post-orbital pillar, its teeth with wrinkled enamel, and its amphicoelous vertebrae. In its general outlook, this animal is rather reminiscent of the Goniopholidae, from the Upper Jurassic and the Cretaceous of Laurasia, but it is more primitive than them, and resemblances are probably due to convergence phenomena. The family Trematochampsidae is probably essentially Gondwanian, and could persist until the Senonian thanks to a certain isolation from Laurasia, where more progressive Crocodylia became predominant much earlier than in Africa and South America.     

From Lizard to Snake; Behind the Evolution of an Extreme Body Plan

The elongated, snake-like skeleton, as it has convergently evolved in numerous reptilian and amphibian lineages, is from a developmental biologist’s point of view amongst the most fascinating anatomical peculiarities in the animal kingdom. This type of body plan is characterized by a greatly increased number of vertebrae, a reduction of skeletal regionalization along the primary body axis and loss of the limbs. Recent studies conducted on both mouse and snakes now hint at how changes inside the gene regulatory circuitries of the Hox genes and the somitogenesis clock likely underlie these striking departures from standard tetrapod morphology, suggesting scenarios by which snakes and other elongated species may have evolved from more ordinarily bodied ancestors.     

New Data on Dinosaurs of the Crimean Peninsula

Reexamination of the holotype of Riabininohadros weberae from the Upper Cretaceous (upper Maastrichtian) of the Crimean Peninsula (Besh-Kosh) allowed determination of previously unknown elements of the femur, astragalus, and calcaneus. This taxon shows a set of primitive characters observed in iguanodontids and basal ornithischians and is referred to as Styracosterna indet. The second dinosaur specimen from Crimea (Aleshino) is a fragmentary skeleton, including cervical and dorsal vertebrae. It possibly belongs to advanced iguanodontids or primitive hadrosauroids. Thus, in the Maastrichtian of the Crimean Peninsula, at least two dinosaur species coexisted.     

A new species of lesser panda Parailurus (Mammalia,Carnivora) from the Pliocene of Transbaikalia (Russia) and some aspects of ailurine phylogeny

A new species of the lesser panda, Parailurus baikalicus sp. nov., from the Pliocene of Transbaikalia is described. In contrast to the European taxa P. anglicus and P. hungaricus, it retains a primitive occlusal pattern of M¹-M², with a concave buccal outline, small mesostyle on M¹, and undeveloped styles on M². At the same time, the Transbaikalian panda is more advanced than other representatives of Parailurus in the upper molars with a reduced lingual cingulum and an enlarged paraconule, which is partially (M¹) or completely (M²) separated from the protocone. This combination of primitive and advanced characters points to the separation of the Asian branch at the earliest stages of the genus development. The Transbaikalian lesser panda may represent a terminal form of this phylogenetic lineage.     

Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya

This paper describes the morphology of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene primate species excavated from Nachola, Kenya in 1999-2002. The cervical vertebrae in Nacholapithecus are larger than those of Papio cynocephalus. They are more robust relative to more caudal vertebral bones. Since Nacholapithecus had large forelimbs, it is assumed that strong cervical vertebrae would have been required to resist muscle reaction forces during locomotion. On the other hand, the vertebral foramen of the lower cervical vertebrae in Nacholapithecus is almost the same size as or smaller than that of P. cynocephalus. Atlas specimens of Nacholapithecus resemble those of extant great apes with regard to the superior articular facet, and they have an anterior tubercle trait intermediate between that of extant apes and other primate species. Nacholapithecus has a relatively short and thick dens on the axis, similar to those of extant great apes and the axis body shape is intermediate between that of extant apes and other primates. Moreover, an intermediate trait between extant great apes and other primate species has been indicated with regard to the angle between the prezygapophyseal articular facets of the axis in Nacholapithecus. Although the atlas of Nacholapithecus is inferred as having a primitive morphology (i.e., possessing a lateral bridge), the shape of the atlas and axis leads to speculation that locomotion or posture in Nacholapithecus involved more orthograde behavior similar to that of extant apes, and, in so far as cervical vertebral morphology is concerned, it is thought that Nacholapithecus was incipiently specialized toward the characteristics of extant hominoids.     

Comparison of Phylogeny,Venom Composition and Neutralization by Antivenom in Diverse Species of Bothrops Complex

In Latin America, Bothrops snakes account for most snake bites in humans, and the recommended treatment is administration of multispecific Bothrops antivenom (SAB – soro antibotrópico). However, Bothrops snakes are very diverse with regard to their venom composition, which raises the issue of which venoms should be used as immunizing antigens for the production of pan-specific Bothrops antivenoms. In this study, we simultaneously compared the composition and reactivity with SAB of venoms collected from six species of snakes, distributed in pairs from three distinct phylogenetic clades: Bothrops, Bothropoides and Rhinocerophis. We also evaluated the neutralization of Bothrops atrox venom, which is the species responsible for most snake bites in the Amazon region, but not included in the immunization antigen mixture used to produce SAB. Using mass spectrometric and chromatographic approaches, we observed a lack of similarity in protein composition between the venoms from closely related snakes and a high similarity between the venoms of phylogenetically more distant snakes, suggesting little connection between taxonomic position and venom composition. P-III snake venom metalloproteinases (SVMPs) are the most antigenic toxins in the venoms of snakes from the Bothrops complex, whereas class P-I SVMPs, snake venom serine proteinases and phospholipases A₂ reacted with antibodies in lower levels. Low molecular size toxins, such as disintegrins and bradykinin-potentiating peptides, were poorly antigenic. Toxins from the same protein family showed antigenic cross-reactivity among venoms from different species; SAB was efficient in neutralizing the B. atrox venom major toxins. Thus, we suggest that it is possible to obtain pan-specific effective antivenoms for Bothrops envenomations through immunization with venoms from only a few species of snakes, if these venoms contain protein classes that are representative of all species to which the antivenom is targeted.     

The phylogeny of varanoid lizards and the affinities of snakes

Evidence that platynotan squamates (living varanoid lizards, snakes and their fossil relatives) are monophyletic is presented. Evolutionary relationships within this group are then ascertained through a cladistic analysis of 144 osteological characters. Mosasauroids (aigialosaurs and mosasaurs), a group of large marine lizards, are identified as the nearest relatives of snakes, thus resolving the long-standing problem of snake affinities. The mosasauroid–snake clade (Pythonomorpha) is corroborated by 40 derived characters, including recumbent replacement teeth, thecodonty, four or fewer premaxillary teeth, supratemporal–prootic contact, free mandibular tips, crista circumfenestralis, straight vertical splenio-angular joint, loss of posterior ramus of the coronoid, reduced basipterygoid processes, reduced interpterygoid vacuity, zygosphene–zygantral articulations, and absence of epiphyses on the axial skeleton and skull. After mosasauroids, the next closest relatives of snakes are varanids (Varanus, Saniwa and Saniwides) and lanthanotids (Lanthanotus and Cherminotus). Derived features uniting varanids and lanthanotids include nine cervical vertebrae and three or fewer pairs of sternal ribs. The varanid–lanthanotid–pythonomorph clade, here termed Thecoglossa, is supported by features such as the anteriorly positioned basal tubera, and the loss of the second epibranchial. Successive outgroups to thecoglossans are Telmasaurus, an unresolved polytomy (Estesia, Gobidermatidae and Helodermatidae), Paravaranus and Proplatynota. The ''necrosaurs'' are demonstrated to be an artificial (polyphyletic) assemblage of primitive platynotans that are not particularly closely related to each other.Snakes are presumed to have evolved from small, limbless, burrowing lizards and the inability of previous analyses to resolve the affinities of snakes has been attributed to extensive convergence among the numerous lineages of such lizards. The present study contradicts this claim, demonstrating that the problem is due instead to omission of critical fossil taxa. No modern phylogenetic analysis of squamate relationships has simultaneously included both mosasauroids and snakes: previous studies have therefore failed to identify the mosasauroid–snake association and the suite of derived characters supporting it. Mosasauroids are large aquatic animals with well-developed appendages, and none of the derived characters uniting mosasauroids and snakes is obviously correlated with miniaturization, limb reduction or fossoriality. Recognition that mosasauroids, followed by varanids and lanthanotids, are the nearest relatives of snakes will also facilitate studies of relationships within snakes, which until now have been hampered by uncertainty over the most appropriate (closely-related) lizard outgroups.     

Diversity of haemogregarine parasites infecting Brazilian snakes from the Midwest and Southeast regions with a description of two new species of Hepatozoon (Apicomplexa: Adeleorina: Hepatozoidae)

Although Brazil is a hotspot for snake species, there is a lack of information on the biodiversity of haemoparasites infecting these hosts. Thus, the present study aimed to bring new insights on the diversity of species of Hepatozoon (Apicomplexa: Adeleorina: Hepatozoidae) infecting Brazilian snakes from the Midwest and Southeast regions. The snakes were captured from 2018 to 2020 from the states of Mato Grosso, Mato Grosso do Sul, Goiás, and São Paulo. Three to five blood smears were made and the remaining blood sample was stored for further molecular analysis. Moreover, histopathological slides of the organs were stained with haematoxylin-eosin. Regarding molecular diagnosis, PCR was performed targeting different regions of the 18S rRNA gene of apicomplexan parasites. From the 13 free-living snakes screened, ten (76.92%) were found infected with Hepatozoon spp. Based on morphological and morphometric tools, five different morphotypes of species of Hepatozoon gamonts were detected. Molecular data and phylogenetic analysis support the morphological data, identifying five species of Hepatozoon from snakes, of which three species belong to previously described species, Hepatozoon cevapii, Hepatozoon cuestensis, and Hepatozoon quagliattus, with a genetic similarity of 100% (based on the 18S rRNA genetic marker). The present study identifies and describes two new species of Hepatozoon, Hepatozoon annulatum sp. nov. infecting the snake Leptodeira annulata and Hepatozoon trigeminum sp. nov. infecting the snake Oxyrhopus trigeminus. Thus, based on morphological and molecular data the present study provides new insights on haemogregarine diversity infecting Brazilian snakes from the Midwest and Southeast regions.     

An analysis of two Myosorex species (Soricidae) from the Early Pliocene site of Langebaanweg (West coast, South Africa) using geometric morphometrics, linear measurements, and non-metric characters

This represents the first study of the soricid community from the Early Pliocene site of Langebaanweg (LBW) which is represented by four species, two of which belong to the soricid genus Myosorex (Forest shrew). Using geometric morphometrics the two Myosorex fossil species are compared with each other, and the extant western and eastern Cape species: Myosorex varius, M. cafer and M. longicaudatus. In addition, the results from an analysis of a number of non-metric characters and linear measurements on the fossil species are presented. The mandibles of the fossil species are characterized by extremely broad teeth, ascending rami which are relatively gracile and differ in shape and size from modern species, and the existence of a number of primitive features of the mandibles and teeth. This evidence is interpreted as indicating that the two LBW Myosorex species represent an archaic lineage now extinct in southern Africa.     

Phylogenetic Relationships among DROSOPHILA LONGICORNIS, DROSOPHILA PROPACHUCA and DROSOPHILA PACHUCA, a Triad of Sibling Species

Drosophila longicornis, D. propachuca and D. pachuca comprise a triad of sibling species. They are morphologically indistinguishable, sympatric forms that, under laboratory conditions, are capable of exchanging genes through the production of fertile F₁ females. However, we have no evidence for introgressive hybridization in nature. The chromosomal constitution of our strains indicates that the ancestral species had the Primitive E gene sequence, and therefore differed from the standard repleta sequence by being Xabc; 2abcg; 3abc. This Primitive E sequence is found in both D. propachuca and D. longicornis. Each of these two species has its own unique rearrangements. D. pachuca is a derived species, which evolved from propachuca. It is cytologically more advanced and has, as its most primitive gene arrangement, one of the more advanced arrangements found in propachuca.     

Study of the Vallesian hipparions of the lower Axios valley(Macedonia,Greece)

The hipparions of the Vallesian locality «Ravin dela Pluie in the lower Axios valley are studied. Two different species were found, a large one, H. primigenium and another small one, H. macedonicum. Their characteristics and dimensions as well as their comparison to the eurasiatic Vallesian samples allow us to find that they belong to Upper Vallesian. H. primigenium of Ravin de la Pluie is more evolved than the typical one of Eppelsheim, while H. macedonicum represents to oldest small hipparion which probably comes from a large form of Lower Vallesian. From the former took rise the later small hipparions. Some primitive features of hipparions are also discussed.     

New macrostomatan snake from the Paleogene of northwestern Argentina

The lower Eocene Lumbrera Formation in Salta province, northwestern Argentina, outstands for providing snake remains from a non-Patagonian Paleogene site. The material consists of articulated precloacal vertebrae that represent a new medium-sized macrostomatan snake, namely Amaru scagliai nov. gen., nov. sp. The vertebral characters of Amaru scagliai nov. gen., nov. sp., suggest affinities with advanced clades, which is consistent with the recognition of derived macrostomatans in the early Paleocene of Bolivia and early Eocene of Brazil. The new snake confirms the presence of macrostomatan snakes in South America as early as the Eocene and suggests that the southern continents may have played an unsuspected role in the origin and evolution of advanced macrostomatans during the earliest Cenozoic.     

Venom glands and some associated muscles in sea snakes

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.