Live birth in a 47-million-year-old snake.

Viviparity is a widespread reproductive trait in snakes, although fossil evidence bearing on its evolution is extremely sparse. Here, we report an exceptional specimen of the minute booid snake Messelophis variatus recovered in the paleolake of the Messel Formation (early-middle Eocene, Germany). This gravid female contains at least two embryos located in the posterior third of the trunk region. The morphology, size, and degree of ossification of the cranial and postcranial remains indicate they correspond with late embryos. This specimen documents the first occurrence of viviparity in a fossil snake and extends the temporal distribution of this reproductive strategy in booid snakes by over 47 Ma. The evolution of viviparity in squamates has traditionally been associated with cold climates, but its presence at the dawn of the evolution of booids during early Palaeogene thermal peaks indicates that viviparity may have evolved under different selective pressures in this clade.

A giant on the ground: another large-bodied Atractus (Serpentes: Dipsadinae) from Ecuadorian Andes, with comments on the dietary specializations of the goo-eaters snakes.

Body-size is significantly correlated with the number of vertebrae (pleomerism) in multiple vertebrate lineages, indicating that somitogenesis process is an important factor dictating evolutionary change associated to phyletic allometry and, consequently, species fitness and diversification. However, the role of the evolution of extreme body sizes (dwarfism and gigantism) remains elusive in snakes, mainly with respect to postnatal ontogeny in dietary preferences associated with evolution of gigantism in many lineages. We described herein a new species in the highly diversified and species-rich genus Atractus on the basis of four specimens from the southeastern slopes of the Ecuadorian Andes. The new species is morphologically similar and apparently closely related to two other allopatric giant congeners (A. gigas and A. touzeti), from which it can be distinguished by their distinct dorsal and ventral coloration, the number of supralabial and infralabial scales, the number of maxillary teeth, and relative width of the head. In addition, we discuss on the ontogenetic trajectories hypotheses and dietary specializations related to evolution of gigantism in the goo-eaters genus Atractus.

Ancient DNA from the extinct South American giant glyptodont Doedicurus sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos.

Glyptodonts were giant (some of them up to ~2400 kg), heavily armoured relatives of living armadillos, which became extinct during the Late Pleistocene/early Holocene alongside much of the South American megafauna. Although glyptodonts were an important component of Cenozoic South American faunas, their early evolution and phylogenetic affinities within the order Cingulata (armoured New World placental mammals) remain controversial. In this study, we used hybridization enrichment and high-throughput sequencing to obtain a partial mitochondrial genome from Doedicurus sp., the largest (1.5 m tall, and 4 m long) and one of the last surviving glyptodonts. Our molecular phylogenetic analyses revealed that glyptodonts fall within the diversity of living armadillos. Reanalysis of morphological data using a molecular 'backbone constraint' revealed several morphological characters that supported a close relationship between glyptodonts and the tiny extant fairy armadillos (Chlamyphorinae). This is surprising as these taxa are among the most derived cingulates: glyptodonts were generally large-bodied and heavily armoured, while the fairy armadillos are tiny (~9-17 cm) and adapted for burrowing. Calibration of our phylogeny with the first appearance of glyptodonts in the Eocene resulted in a more precise timeline for xenarthran evolution. The osteological novelties of glyptodonts and their specialization for grazing appear to have evolved rapidly during the Late Eocene to Early Miocene, coincident with global temperature decreases and a shift from wet closed forest towards drier open woodland and grassland across much of South America. This environmental change may have driven the evolution of glyptodonts, culminating in the bizarre giant forms of the Pleistocene.

Paedomorphosis and retention of juvenile diet lead speciation in a group of Neotropical snakes (Colubroides-Philodryadini).

Dipsadidae is one of the largest clades of extant reptiles, showing an impressive morphological and ecological diversity. Despite this fact, the developmental processes behind its diversity are still largely unknown. In this study, we used 3D reconstructions based on micro-CT data and geometric morphometrics to evaluate the skull morphology of Philodryas agassizii, a small, surface-dwelling dipsadid that consume spiders. Adult individuals of P. agassizii exhibit a cranial morphology frequently observed in juveniles of other surface-dwelling colubroideans, represented in our analysis by its close relative Philodryas patagoniensis. Large orbits, gibbous neurocranial roof and a relatively short jaw complex are features present in juveniles of the latter species. Furthermore, we performed an extensive survey about diet of P. patagoniensis in which we detected an ontogenetic dietary shift, indicating that arthropods are more frequently consumed by juveniles of this dietary generalist. Thus, we infer that P. agassizzii retained not only the ancestral juvenile skull morphology but also dietary preferences. This study reveals that morphological changes driven by heterochronic changes, specifically paedomorphosis, influenced the retention of ancestral life history traits in P. agassizii, and therefore promoted cladogenesis. In this way, we obtained first evidence that heterochronic processes lead speciation in the snake megadiverse clade Dipsadidae.

Postnatal ontogeny and the evolution of macrostomy in snakes.

Macrostomy is the anatomical feature present in macrostomatan snakes that permits the ingestion of entire prey with high cross-sectional area. It depends on several anatomical traits in the skeleton and soft tissues, of which the elongation of gnathic complex and backward rotation of the quadrate represent crucial skeletal requirements. Here, the relevance of postnatal development of these skull structures and their relationship with macrohabitat and diet are explored. Contrary to the condition present in lizards and basal snakes that occupy underground macrohabitats, elements of the gnathic complex of most macrostomatan snakes that exploit surface macrohabitats display conspicuous elongation during postnatal growth, relative to the rest of the skull, as well as further backward rotation of the quadrate bone. Remarkably, several clades of small cryptozoic macrostomatans reverse these postnatal transformations and return to a diet based on prey with low cross-sectional area such as annelids, insects or elongated vertebrates, thus resembling the condition present in underground basal snakes. Dietary ontogenetic shift observed in most macrostomatan snakes is directly linked with this ontogenetic trajectory, indicating that this shift is acquired progressively as the gnathic complex elongates and the quadrate rotates backward during postnatal ontogeny. The numerous independent events of reversion in the gnathic complex and prey type choice observed in underground macrostomatans and the presence of skeletal requirements for macrostomy in extinct non-macrostomatan species reinforce the possibility that basal snakes represent underground survivors of clades that had the skeletal requirements for macrostomy. Taken together, the data presented here suggest that macrostomy has been shaped during multiple episodes of occupation of underground and surface macrohabitats throughout the evolution of snakes.

Postnatal development of the skull of Dinilysia patagonica (Squamata-stem Serpentes).

The snake skull represents a profound transformation of the ancestral squamate cranium in which dermal skull roof bones were integrated with the braincase, in a manner convergent with that which occurred during the origin of mammals. However, the ontogeny of snake characters at the origin of the clade has until now been inaccessible. Here we describe a postnatal ontogenetic series of the Late Cretaceous stem snake Dinilysia patagonica and compare it to that of extant lizards and snakes. Comparative analysis indicates notable ontogenetic changes, including advanced state of ossification, isometric growth of the otic capsule, fusion of the stylohyal to the quadrate, and great posterior elongation of the supratemporal. Of these transformations, the unfused condition of braincase bones and the retention of a large otic capsule in adults are examples of paedomorphic and peramorphic processes, respectively. Some ontogenetic transformations detected, in particular those present in middle ear, skull roof and suspensorium, are strikingly similar to those present in extant snakes. Nevertheless, Dinilysia retains a lizard-like paroccipital process without an epiphyseal extremity, and a calcified epiphysis that caps the sphenoccipital tubercle. Finally, the integration of the dermal skull roof with the braincase is similar to that seen in mammals with regard to the overall closure of the braincase, but the two evolutionary and developmental modules appear less integrated in snakes in that the parietal bone of the dermal skull roof progressively overlaps the supraoccipital of the chondrocranial braincase.

A new snake skull from the Paleocene of Bolivia sheds light on the evolution of macrostomatans.

Macrostomatan snakes, one of the most diverse extant clades of squamates, display an impressive arsenal of cranial features to consume a vast array of preys. In the absence of indisputable fossil representatives of this clade with well-preserved skulls, the mode and timing of these extraordinary morphological novelties remain obscure. Here, we report the discovery of Kataria anisodonta n. gen. n. sp., a macrostomatan snake recovered in the Early Palaeocene locality of Tiupampa, Bolivia. The holotype consists of a partial, minute skull that exhibits a combination of booid and caenophidian characters, being the presence of an anisodont dentition and diastema in the maxilla the most distinctive trait. Phylogenetic analysis places Kataria basal to the Caenophidia+Tropidophiidae, and represents along with bolyeriids a distinctive clade of derived macrostomatans. The discovery of Kataria highlights the morphological diversity in the maxilla among derived macrostomatans, demonstrating the relevance of maxillary transformations in the evolution of this clade. Kataria represents the oldest macrostomatan skull recovered, revealing that the diversification of macrostomatans was well under way in early Tertiary times. This record also reinforces the importance of Gondwanan territories in the history of snakes, not only in the origin of the entire group but also in the evolution of ingroup clades.