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Abstract Recent phylogenetic analyses differ in their interpretations of the origin and interrelationships of snakes, resulting in polarized views of snake ecology, habit and acquisition of features associated with wide-gaped feeding (macrostomy). Here, we report a new specimen of the Late Cretaceous nest predator Sanajeh indicus that helps to resolve the origin of macrostomy. The new specimen preserves an ossified upper temporal bar and a posteriorly expanded otooccipital region that lacks a free-ending supratemporal bone and retains a lizard-like palatomaxillary arch that allows limited movements during swallowing. Phylogenetic analyses of a large-scale total evidence dataset resolve Sanajeh near the base of Pan-Serpentes, as the sister group of Najash, Dinilysia and crown-group Serpentes. The Cretaceous Tetrapodophis and Coniophis represent the earliest-diverging members of Pan-Serpentes. The Cretaceous hindlimbed pachyophiids and Cenozoic Australian ‘madtsoiids’ are inside crown Alethinophidia, whereas mosasaurs are recovered invariably within anguimorphs. Our results suggest that the wide-gape condition in mosasaurs and snakes might have evolved independently, as functionally distinct mechanisms of prey ingestion. The intermediate morphology preserved in Sanajeh indicates that ingestion of large prey items (macrophagy) preceded wide-gaped, unilateral feeding (macrostomy), which appeared 35 Myr later, in the common ancestor of pachyophiids, Cenozoic Australian ‘madtsoiids’ and alethinophidians. 

Cretaceous dinosaurs were first reported from the Indian subcontinent in the late 1800s, and titanosaur sauropod and abelisauroid theropod remains are now known from central, western, and southern parts of India and from central western Pakistan. Although dinosaur remains are abundant, associated or articulated specimens are extremely rare, and so are complex skeletal elements such as cranial bones and presacral vertebrae. The historical pattern of sampling and collecting has limited the inferences about patterns of diversity, phylogenetic affinity, and paleobiogeographic relationships of Indian dinosaurs. Here we report on three titanosaur vertebrae representing regions of the skeleton that are complex and otherwise poorly represented in the Indian record, including two anterior dorsal vertebrae pertaining to a single individual from Rahioli, in Gujarat State (western India), and an anterior caudal neural arch from Bara Simla, in Madhya Pradesh State (central India). Phylogenetic analysis places the two individuals within Titanosauria, but further resolution of their affinities is precluded by their incompleteness and that of titanosaur vertebral columns in general, lack of coding of character data for titanosaur presacral and anterior caudal vertebrae, and relatively coarse understanding of the evolutionary relationships of titanosaurs. Comparisons with contemporaneous and spatially proximal titanosaurs from Indo-Pakistan, Madagascar, and South America provide insights into their affinities. The dorsal vertebrae share close affinity with Isisaurus from India and Mendozasaurus from Argentina. Few local comparisons are available for the anterior caudal vertebra, which shares characteristics with Tengrisaurus from the Early Cretaceous of Russia.