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The origin of snakes: revealing the ecology, behavior, and evolutionary history of early snakes using genomics, phenomics, and the fossil record

机译:蛇的起源:使用基因组学,表型学和化石记录揭示早期蛇的生态,行为和进化史

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Background The 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. Results Comprehensive 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. Conclusions Our 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.
机译:背景技术蛇的高度衍生的形态和惊人的多样性长期以来一直在引发关于蛇总群(Pan-Serpentes)和冠蛇(Serpentes)的生态和进化起源的争论。尽管对最早的蛇的生态,行为和出处有很多推测,但仍未尝试对整个蛇的起源进行严格的,进化枝范围的分析,部分原因是缺乏有关早期茎蛇的足够的古生物学数据。在这里,我们提出了对冠蛇祖先和蛇总群祖先的第一个综合分析重建,这是使用多种祖先状态重建方法推断得出的。我们使用组合数据的方法,包括来自灭绝的冠蛇化石记录的新信息,有关茎蛇Najash rionegrina,Dinilysia patagonica和Coniophis先例的解剖学的新数据,以及对表型无性型分布的更深入了解。化石和最近的蛇的主要进化枝。此外,我们使用新的“尖端约会”和传统的基于节点的方法来推断时间校准的系统发育,从而为蛇的早期进化史上的时间模式提供了新的见解。结果全面的祖先状态重建表明,冠蛇的祖先和全群蛇的祖先都是夜间活动的,广泛觅食的,非限制性的隐身猎人。他们可能消耗了大小不等的软体动物脊椎动物和无脊椎动物,并在温暖,水源充足和植被茂密的环境中占据了陆地环境。蛇的总群(由Coniophis节点近似)被推断是起源于白垩纪中期(〜128.5?Ma)的陆地,而冠群则是在大约2000万年之后的阿尔比亚时期。我们推测的发散日期为白垩纪-古生代(K-Pg)大规模灭绝后主要的三栖类蛇多样性辐射提供了有力证据,从而阐明了现存蛇辐射的模式和时间。尽管蛇冠群最有可能出现在冈瓦纳的超大陆上,但我们的研究结果表明,蛇的总群起源于劳拉西亚。结论我们的研究提供了有关蛇何时,何地以及如何起源的新见解,并提供了迄今为止蛇的早期进化的最完整图景。更广泛地说,我们证明了结合化石和表型数据的惊人影响,旨在进行系统发育拓扑推断和祖先状态重建。

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