Summary:Odontogriphus omalus was originally described as a problematic non-biomineralized lophophorate organism. Here we reinterpret Odontogriphus based on 189 new specimens including numerous exceptionally well-preserved individuals from the Burgess Shale collections of the Royal Ontario Museum. This additional material provides compelling evidence that the feeding apparatus in Odontogriphus is a radula of molluscan architecture comprising two primary bipartite tooth rows attached to a radular membrane and showing replacement by posterior addition. Further characters supporting molluscan affinity include a broad foot bordered by numerous ctenidia located in a mantle groove and a stiffened cuticular dorsum. Odontogriphus has a radula similar to Wiwaxia corrugata but lacks a scleritome. We interpret these animals to be members of an early stem-group mollusc lineage that likely originated in the Neoproterozoic Ediacaran Period, providing support for the retention of a biomat-based grazing community from the late Precambrian until at least the Middle Cambrian.
A remarkable new fossil horseshoe crab, Lunataspis aurora gen. et sp. nov., from recently discovered Upper Ordovician (c. 445 Ma) shallow marine Konservat-Lagerstät-ten deposits in Manitoba (Canada), is characterized by fusion of opisthosomal tergites into two sclerites. A broad mesosoma of six or seven fused segments, followed by a narrow metasoma of three reduced segments, represents an advanced transitional condition in the development of the xiphosurid thoracetron. Lunataspis further possesses a large crescentic prosomal shield bearing lateral compound eyes on weak ophthalmic ridges that flank a low cardiac lobe, and a keeled lanceolate telson. Lunataspis is much older than the proposed 'synziphosurine' stem lineage of Carboniferous and post-Palaeozoic Xiphosurida, yet is strikingly similar to crown group limuline horseshoe crabs, indicating that major features of the distinctive and highly conserved xiphosurid Bauplan evolved considerably earlier in the Palaeozoic than was previously suspected. In addition to establishing a new temporal benchmark for assessing hypotheses of early chelicerate relationships, the discovery of horseshoe crabs in a Late Ordovician marginal marine setting marks the earliest definitive record of this persistent ecological association.
Examples of the trilobites Toxochasmops extensus (Boeck, 1838), Asaphiscus wheeleri Meek, 1874, Encrinurus mitchelli Foerste, 1888, Ogygopsis klotzi (Rominger, 1887), Paradoxides davidis Salter, 1863 and Oryctocephalus spp. which are interpreted as exuvial configurations, are described. Four specimens of Toxochasmops extensus arc known in which the pygidium rests either directly behind the eephalon, or with only three intervening thoracic segments. It is considered that during exuviation the old pygidium became wedged behind the cephalon. This facilitated its removal. An ecdysial configuration of Asaphiscus wheeleri is described which possesses inverted and partially rotated free cheeks. In addition, part of the thorax of the specimen is wedged obliquely behind the cephalon. This is considered to have aided withdrawal of the trilobite from its old exoskeleton. Two specimens of Encrinurus mitchelli are described which possess free cheeks inverted beneath the cranidium by lateral rotation, in a manner similar to that of A. wheeleri. Three examples of Ogygopsis klotzi are described, one a failed exuvia and two in which the free cheeks were inverted and rotated through 180° with respect to their original position and came to rest beneath the thorax. An identical exuvia of Paradoxides davidis trapezopyge is also described. Two specimens of Oryctocephalus exhibiting two different arrangements of inverted free cheeks are recorded. Possible mechanisms for each of these free cheek inversions are proposed.
Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Osnabrück, Germany, represent a relatively complete growth series of 10 stages. Based on this growth sequence, morphological changes throughout the ontogeny can be identified. The major change affects the shape of the epimera of the opisthosoma. In earlier stages, they appear very spine-like, whereas in later stages the bases of these spine-like structures become broader; the broadened bases are then successively drawn out distally. In the most mature stage known, the epimera are of trapezoidal shape and approach each other closely to form a complete flange around the thoracetron (=fused tergites of the opisthosoma). These ontogenetic changes question the taxonomic status of different species of Euproops, as the latter appear to correspond to different stages of the ontogenetic series reconstructed from the Piesberg specimens. This means that supposed separate species could, in fact, represent different growth stages of a single species. It could alternatively indicate that heterochrony (=evolutionary change of developmental timing) plays an important role in the evolution of Xiphosura. We propose a holomorph approach, i.e., reconstructing ontogenetic sequences for fossil and extant species as a sound basis for a taxonomic, phylogenetic, and evolutionary discussion of Xiphosura.
One of the oldest known fossil scorpions, a new species from the mid-Silurian Eramosa Formation (430 myr) of Ontario, Canada, exhibits several surprising features. The depositional environment and associated biota indicate a marine habitat; however, the leg morphology of this scorpion, which has a short tarsus in common with all Recent scorpions, suggests that a key adaptation for terrestrial locomotion, the ability to support its weight on a subterminal 'foot', appeared remarkably early in the scorpion fossil record. Specimens are preserved intact and undisturbed in a splayed posture typical of moults rather than carcasses. We postulate that these animals were aquatic, but occasionally ventured into extremely shallow water, or onto a transient subaerially exposed surface while moulting, before returning to deeper water. Shed exuviae were preserved in situ by rapid overgrowth of bacterial biofilm.
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