Systematics and Phylogenetic Interrelationships of the Enigmatic Late Jurassic Shark Protospinax annectans Woodward, 1918 with Comments on the Shark-Ray Sister Group Relationship.

The Late Jurassic elasmobranch Protospinax annectans is often regarded as a key species to our understanding of crown group elasmobranch interrelationships and the evolutionary history of this group. However, since its first description more than 100 years ago, its phylogenetic position within the Elasmobranchii (sharks and rays) has proven controversial, and a closer relationship between Protospinax and each of the posited superorders (Batomorphii, Squalomorphii, and Galeomorphii) has been proposed over the time. Here we revise this controversial taxon based on new holomorphic specimens from the Late Jurassic Konservat-Lagerstätte of the Solnhofen Archipelago in Bavaria (Germany) and review its skeletal morphology, systematics, and phylogenetic interrelationships. A data matrix with 224 morphological characters was compiled and analyzed under a molecular backbone constraint. Our results indicate a close relationship between Protospinax, angel sharks (Squatiniformes), and saw sharks (Pristiophoriformes). However, the revision of our morphological data matrix within a molecular framework highlights the lack of morphological characters defining certain groups, especially sharks of the order Squaliformes, hampering the phylogenetic resolution of Protospinax annectans with certainty. Furthermore, the monophyly of modern sharks retrieved by molecular studies is only weakly supported by morphological data, stressing the need for more characters to align morphological and molecular studies in the future.

A well-preserved Archaeopteryx specimen with theropod features.

A nearly complete skeleton of Archaeopteryx with excellent bone preservation shows that the osteology of the urvogel is similar to that of nonavian theropod dinosaurs. The new specimen confirms the presence of a hyperextendible second toe as in dromaeosaurs and troodontids. Archaeopteryx had a plesiomorphic tetraradiate palatine bone and no fully reversed first toe. These observations provide further evidence for the theropod ancestry of birds. In addition, the presence of a hyperextendible second toe blurs the distinction of archaeopterygids from basal deinonychosaurs (troodontids and dromaeosaurs) and challenges the monophyly of Aves.