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Problematic fossils, extinct taxa of enigmatic morphology that cannot be assigned to a known major group, were once a major issue in palaeontology. A long-favoured solution to the 'problem of the problematica', particularly the 'weird wonders' of the Cambrian Burgess Shale, was to consider them representatives of extinct phyla. A combination of new evidence and modern approaches to phylogenetic analysis has now resolved the affinities of most of these forms. Perhaps the most notable exception is Tullimonstrum gregarium, popularly known as the Tully monster, a large soft-bodied organism from the late Carboniferous Mazon Creek biota (approximately 309-307 million years ago) of Illinois, USA, which was designated the official state fossil of Illinois in 1989. Its phylogenetic position has remained uncertain and it has been compared with nemerteans, polychaetes, gastropods, conodonts, and the stem arthropod Opabinia. Here we review the morphology of Tullimonstrum based on an analysis of more than 1,200 specimens. We find that the anterior proboscis ends in a buccal apparatus containing teeth, the eyes project laterally on a long rigid bar, and the elongate segmented body bears a caudal fin with dorsal and ventral lobes. We describe new evidence for a notochord, cartilaginous arcualia, gill pouches, articulations within the proboscis, and multiple tooth rows adjacent to the mouth. This combination of characters, supported by phylogenetic analysis, identifies Tullimonstrum as a vertebrate, and places it on the stem lineage to lampreys (Petromyzontida). In addition to increasing the known morphological disparity of extinct lampreys, a chordate affinity for T. gregarium resolves the nature of a soft-bodied fossil which has been debated for more than 50 years.

The evolutionary trajectory of early complex life on Earth is interpreted largely from the fossils of the Precambrian soft-bodied Ediacara Biota, which appeared and evolved during a time of dynamic biogeochemical and environmental fluctuation in the global ocean. The Ediacara Biota is historically divided into three successive Assemblages—the Avalon, the White Sea, and the Nama—which are marked by the appearance of novel biological traits and ecological strategies. In particular, the younger White Sea and Nama Assemblages record a “second wave” of ecological innovations, which included not only the development of uniquely Ediacaran body plans and ecologies, such as matground adaptations, but also the dual emergence of bilaterian-grade animals and Phanerozoic-style ecological innovations, including spatial heterogeneity, complex reproductive strategies, ecospace utilization, motility, and substrate competition. The late Ediacaran was an evolutionarily dynamic time characterized by strong environmental control over the distribution of taxa in time and space.

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