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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' (1), particularly the 'weird wonders' (2) 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 (3), 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 (4,5), polychaetes (4), gastropods (4), conodonts (6), and the stem arthropod Opabinia (4). 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 (3-6). 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 (7-9), a chordate affinity for T. gregarium resolves the nature of a soft-bodied fossil which has been debated for more than 50 years., Since T. gregarium was originally described as a representative of an extinct phylum (3,5), there have been only two attempts using extensive additional specimens to resolve its affinity (4,6). These [...]

We introduce a new non-destructive source of skeletochronological data with applications to species identification, associating disarticulated remains, assessing minimum number of individuals (MNI), and collection management of fossil snakes, but with potential implications for all bony vertebrates, extinct or extant. Study of a diverse sample of Recent henophidian snakes confirms that annual growth cycles (AGCs) visible on the surface of the vertebral zygantrum correspond to lines of arrested growth in osteohistological thin sections and accordingly reflect chronological age. None of the specimens considered here showed signs of remodelling of the zygantrum, suggesting that a complete, unaltered age record is preserved. We tested potential influences on AGCs with a single experimental organism, a male Bogertophis subocularis, that was raised at a controlled temperature and with constant access to mice and water. The conditions in which this individual was maintained, including that it had yet to live through a full reproductive cycle, enabled us to determine that its AGCs reflect only the annual solar cycle, and neither temperature, nor resource availability, nor energy diversion to gametogenesis could explain that it still exhibited lines of arrested growth. Moreover, growth lines in this specimen are deposited toward the end of the growth season in the fall, and not in the winter, during which this individual continued to feed and grow, even though this mid-latitude species would normally be hibernating and not growing. This suggests that growth lines are not caused by hibernation, but reflect the onset of a physiological cycle preparing Bogertophis subocularis for winter rest. That being said, hibernation and reproductive cycle could still influence the amount of time represented by an individual growth line. Growth-line number and AGC spacing-pattern, plus centrum length, are used to estimate MNI of the Early Eocene fossil snake Boavus occidentalis collected from the Willwood Formation over two field seasons during the late 19th century. We identified eight or nine individuals among specimens previously parcelled among two specimen lots collected during those expeditions. [ABSTRACT FROM AUTHOR]