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Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes).

ABSTRACT: Caenophidian snakes include the file snake genus Acrochordus and advanced colubroidean snakes that radiated mainly during the Neogene. Although caenophidian snakes are a well-supported clade, their inferred affinities, based either on molecular or morphological data, remain poorly known or controversial. Here, we provide an expanded molecular phylogenetic analysis of Caenophidia and use three non-parametric measures of support-Shimodaira-Hasegawa-Like test (SHL), Felsentein (FBP) and transfer (TBE) bootstrap measures-to evaluate the robustness of each clade in the molecular tree. That very different alternative support values are common suggests that results based on only one support value should be viewed with caution. Using a scheme to combine support values, we find 20.9% of the 1265 clades comprising the inferred caenophidian tree are unambiguously supported by both SHL and FBP values, while almost 37% are unsupported or ambiguously supported, revealing the substantial extent of phylogenetic problems within Caenophidia. Combined FBP/TBE support values show similar results, while SHL/TBE result in slightly higher combined values. We consider key morphological attributes of colubroidean cranial, vertebral and hemipenial anatomy and provide additional morphological evidence supporting the clades Colubroides, Colubriformes, and Endoglyptodonta. We review and revise the relevant caenophidian fossil record and provide a time-calibrated tree derived from our molecular data to discuss the main cladogenetic events that resulted in present-day patterns of caenophidian diversification. Our results suggest that all extant families of Colubroidea and Elapoidea composing the present-day endoglyptodont fauna originated rapidly within the early Oligocene-between approximately 33 and 28 Mya-following the major terrestrial faunal turnover known as the "Grande Coupure" and associated with the overall climate shift at the Eocene-Oligocene boundary. Our results further suggest that the caenophidian radiation originated within the Caenozoic, with the divergence between Colubroides and Acrochordidae occurring in the early Eocene, at ~ 56 Mya.

SUBMITTER: Zaher H

PROVIDER: S-EPMC6512042 | biostudies-literature |

REPOSITORIES: biostudies-literature

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Project description:BackgroundThe highly derived morphology and astounding diversity of snakes has long inspired debate regarding the ecological and evolutionary origin of both the snake total-group (Pan-Serpentes) and crown snakes (Serpentes). Although speculation abounds on the ecology, behavior, and provenance of the earliest snakes, a rigorous, clade-wide analysis of snake origins has yet to be attempted, in part due to a dearth of adequate paleontological data on early stem snakes. Here, we present the first comprehensive analytical reconstruction of the ancestor of crown snakes and the ancestor of the snake total-group, as inferred using multiple methods of ancestral state reconstruction. We use a combined-data approach that includes new information from the fossil record on extinct crown snakes, new data on the anatomy of the stem snakes Najash rionegrina, Dinilysia patagonica, and Coniophis precedens, and a deeper understanding of the distribution of phenotypic apomorphies among the major clades of fossil and Recent snakes. Additionally, we infer time-calibrated phylogenies using both new 'tip-dating' and traditional node-based approaches, providing new insights on temporal patterns in the early evolutionary history of snakes.ResultsComprehensive ancestral state reconstructions reveal that both the ancestor of crown snakes and the ancestor of total-group snakes were nocturnal, widely foraging, non-constricting stealth hunters. They likely consumed soft-bodied vertebrate and invertebrate prey that was subequal to head size, and occupied terrestrial settings in warm, well-watered, and well-vegetated environments. The snake total-group - approximated by the Coniophis node - is inferred to have originated on land during the middle Early Cretaceous (~128.5 Ma), with the crown-group following about 20 million years later, during the Albian stage. Our inferred divergence dates provide strong evidence for a major radiation of henophidian snake diversity in the wake of the Cretaceous-Paleogene (K-Pg) mass extinction, clarifying the pattern and timing of the extant snake radiation. Although the snake crown-group most likely arose on the supercontinent of Gondwana, our results suggest the possibility that the snake total-group originated on Laurasia.ConclusionsOur study provides new insights into when, where, and how snakes originated, and presents the most complete picture of the early evolution of snakes to date. More broadly, we demonstrate the striking influence of including fossils and phenotypic data in combined analyses aimed at both phylogenetic topology inference and ancestral state reconstruction.

Project description:Ghost shrimps of Callianassidae and Ctenochelidae are soft-bodied, usually heterochelous decapods representing major bioturbators of muddy and sandy (sub)marine substrates. Ghost shrimps have a robust fossil record spanning from the Early Cretaceous (~ 133 Ma) to the Holocene and their remains are present in most assemblages of Cenozoic decapod crustaceans. Their taxonomic interpretation is in flux, mainly because the generic assignment is hindered by their insufficient preservation and disagreement in the biological classification. Furthermore, numerous taxa are incorrectly classified within the catch-all taxon Callianassa. To show the historical patterns in describing fossil ghost shrimps and to evaluate taphonomic aspects influencing the attribution of ghost shrimp remains to higher level taxa, a database of all fossil species treated at some time as belonging to the group has been compiled: 250 / 274 species are considered valid ghost shrimp taxa herein. More than half of these taxa (160 species, 58.4%) are known only from distal cheliped elements, i.e., dactylus and / or propodus, due to the more calcified cuticle locally. Rarely, ghost shrimps are preserved in situ in burrows or in direct association with them, and several previously unpublished occurrences are reported herein. For generic assignment, fossil material should be compared to living species because many of them have modern relatives. Heterochely, intraspecific variation, ontogenetic changes and sexual dimorphism are all factors that have to be taken into account when working with fossil ghost shrimps. Distal elements are usually more variable than proximal ones. Preliminary results suggest that the ghost shrimp clade emerged not before the Hauterivian (~ 133 Ma). The divergence of Ctenochelidae and Paracalliacinae is estimated to occur within the interval of Hauterivian to Albian (133-100 Ma). Callichirinae and Eucalliacinae likely diverged later during the Late Cretaceous (100-66 Ma), whereas Callianassinae did not appear before the Eocene (56 Ma).

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Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes).

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