David M. Rudkin

A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale.

Odontogriphus omalus was originally described as a problematic non-biomineralized lophophorate organism. Here we re-interpret 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 probably originated in the Neoproterozoic Ediacaran Period, providing support for the retention of a biomat-based grazing community from the late Precambrian Period until at least the Middle Cambrian.

Exceptionally preserved Late Ordovician biotas from Manitoba, Canada

There are few body fossil biotas known from early Paleozoic accretionary shorelines, and very few examples of Ordovician soft-bodied assemblages. This study documents two recently discovered biotas from separate sedimentary basins in Manitoba, Canada, that provide unique information about tropical shoreline communities shortly before the Late Ordovician extinction event. Each site represents a distinct depositional environment, but they share biotic elements, including eurypterids, xiphosurids, and large problematic tubes. The William Lake biota, representing more restricted conditions, includes jellyfish that are among the best hydromedusan body fossils known. Rocks at the Airport Cove site, deposited under more open circulation, contain scolecodonts and noncalcified algae. These biotas have some parallels with the recently described Middle Ordovician Winneshiek Lagerstatte, but are also similar to some Late Silurian assemblages. Considered together, early Paleozoic marginal marine deposits are a rich but as yet poorly known source of paleobiodiversity data.

A holomorph approach to xiphosuran evolution—a case study on the ontogeny of Euproops

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.

THE WORLD'S BIGGEST TRILOBITE—ISOTELUS REX NEW SPECIES FROM THE UPPER ORDOVICIAN OF NORTHERN MANITOBA, CANADA

Abstract The largest known trilobite fossil, a virtually complete articulated dorsal shield of the asaphid Isotelus rex new species, has been recovered from Upper Ordovician (Cincinnatian, Richmondian) nearshore carbonates of the Churchill River Group in northern Manitoba. At over 700 mm in length, it is almost 70 percent longer than the largest previously documented complete trilobite, and provides the first unequivocal evidence of maximum trilobite length in excess of one-half metre. Comparisons with other fossil and extant members of the phylum suggest that in terms of maximum linear dimensions it was among the biggest arthropods ever to have lived. Sediments of the Churchill River Group were deposited in an equatorial epeiric setting and the extremely large size of I. rex n. sp. thus marks a striking example of low-latitude gigantism, in sharp contrast to the widespread phenomenon of “polar gigantism” in many modern marine benthic arthropods. Lack of extensive epibiontic colonization of the exoskeletal surface and the presence of large distinctive trace fossils in the same unit suggest that I. rex n. sp. may have been a semi-infaunal predator and scavenger that employed a shallow furrowing and probing mode of benthic feeding. The extinction of the isotelines (and virtually the entire asaphide lineage) at the end of the Ordovician cannot be related to the near contemporaneous achievement of exceptionally large adult size in some representatives. Failure to survive the terminal Ordovician extinction event was most likely a consequence of a pelagic larval life-style that proved ill-adapted to the rapid onset of global climatic cooling and loss of tropical shelf habitats.

A new mid-Silurian aquatic scorpion—one step closer to land?

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.

A NEW LATE SILURIAN (PRIDOLIAN) NARAOIID (EUARTHROPODA: NEKTASPIDA) FROM THE BERTIE FORMATION OF SOUTHERN ONTARIO, CANADA—DELAYED FALLOUT FROM THE CAMBRIAN EXPLOSION

Abstract The discovery of a new naraoiid nektaspid in the Upper Silurian (Pridolian) of southeastern Ontario significantly extends the range of this unusual group. Nektaspids are nonmineralized arthropods typical of Early and Middle Cambrian soft-bottom communities, but were thought to have become extinct in the Late Ordovician. The unique holotype specimen of Naraoia bertiensis n. sp. comes from a Konservat–Lagerstätte deposit renowned for its eurypterid fauna (the Williamsville Member of the Bertie Formation). Naraoia bertiensis lacks thoracic segments and is morphologically similar to Naraoia compacta from the Middle Cambrian Burgess Shale, save for the presence of a long ventral cephalic doublure and a subtly pointed posterior shield. To examine the phylogenetic relationships of the new naraoiid, we coded characters of the holotype specimen and of nine previously described nektaspids. The results confirm a sister taxon relationship between Naraoia compacta and Naraoia bertiensis and the monophyly of nektaspid forms lacking thoracic segments (family Naraoiidae). This latter group may have arisen from an ancestral segment-bearing form through heterochronic loss of thoracic segments early in the Cambrian. The disjunct occurrence of a naraoiid nektaspid in the Late Silurian resembles the reappearance of other “Lazarus taxa” that were thought to have been eliminated during mass extinction events. The naraoiid lineage survived the Late Ordovician biotic crisis, but in this case the “Lazarus effect” seems likely to be taphonomic in origin.

Phyllocarid Crustaceans from the Silurian Eramosa Lagerstätte (Ontario, Canada): Taxonomy and Functional Morphology

Abstract Two species of well-preserved phyllocarid crustaceans, Ceratiocaris macroura n. sp. and Ceratiocaris papilio Salter, occur in the Eramosa Formation Lagerstätte of Ontario, Canada, represented by several dozen articulated fossils preserving limbs and thoracic segmentation. Ceratiocaris macroura n. sp. is distinguished by possession of an exceptionally elongate telson in relation to its body length. Ceratiocaris papilio preserves fine detail of the abdominal scale-like ornamentation, allowing reinterpretation and enhanced characterization of this key feature. Ceratiocaris papilio has a wide paleogeographic distribution across Laurentia but is restricted to shallow marine depositional settings within the middle Silurian.

Oldest Near-Complete Acanthodian: The First Vertebrate from the Silurian Bertie Formation Konservat-Lagerstätte, Ontario

Background The relationships between early jawed vertebrates have been much debated, with cladistic analyses yielding little consensus on the position (or positions) of acanthodians with respect to other groups. Whereas one recent analysis showed various acanthodians (classically known as ‘spiny sharks’) as stem osteichthyans (bony fishes) and others as stem chondrichthyans, another shows the acanthodians as a paraphyletic group of stem chondrichthyans, and the latest analysis shows acanthodians as the monophyletic sister group of the Chondrichthyes. Methodology/Principal Findings A small specimen of the ischnacanthiform acanthodian Nerepisacanthus denisoni is the first vertebrate fossil collected from the Late Silurian Bertie Formation Konservat-Lagerstätte of southern Ontario, Canada, a deposit well-known for its spectacular eurypterid fossils. The fish is the only near complete acanthodian from pre-Devonian strata worldwide, and confirms that Nerepisacanthus has dentigerous jaw bones, body scales with superposed crown growth zones formed of ondontocytic mesodentine, and a patch of chondrichthyan-like scales posterior to the jaw joint. Conclusions/Significance The combination of features found in Nerepisacanthus supports the hypothesis that acanthodians could be a group, or even a clade, on the chondrichthyan stem. Cladistic analyses of early jawed vertebrates incorporating Nerepisacanthus, and updated data on other acanthodians based on publications in press, should help clarify their relationships.

A Curious Rusophycus (Arthropod Ichnofossil) Assemblage from the Upper Ordovician of Ontario, Canada

The ichnogenus Rusophycus includes a wide range of short bilobate excavations generally attributed to variable feeding behaviors of arthropods, especially trilobites. An unusual Rusophycus assemblage from Upper Ordovician Georgian Bay Formation in Ontario departs radically from previously described examples and presents new challenges for understanding the behavior represented by these traces. This specimen is unique in the arrangement of multiple Rusophycus burrows in a circular, lens-shaped array (as opposed to a linear or random arrangement typical of other Rusophycus assemblages). The size and shape of the individual Rusophycus components are consistent with traces attributed to the coeval trilobite Flexicalymene. Multiple Rusophycus assemblages likely reflect aggregations of trilobites in response to a local concentration of food. The topology of this particular Rusophycus assemblage suggests that the trilobites opportunistically exploited a rich and narrowly restricted food source, perhaps the decaying remains of a buried organism.

Earth’s oldest ‘Bobbit worm’ – gigantism in a Devonian eunicidan polychaete

Whilst the fossil record of polychaete worms extends to the early Cambrian, much data on this group derive from microfossils known as scolecodonts. These are sclerotized jaw elements, which generally range from 0.1–2 mm in size, and which, in contrast to the soft-body anatomy, have good preservation potential and a continuous fossil record. Here we describe a new eunicidan polychaete, Websteroprion armstrongi gen. et sp. nov., based primarily on monospecific bedding plane assemblages from the Lower-Middle Devonian Kwataboahegan Formation of Ontario, Canada. The specimens are preserved mainly as three-dimensional moulds in the calcareous host rock, with only parts of the original sclerotized jaw walls occasionally present. This new taxon has a unique morphology and is characterized by an unexpected combination of features seen in several different Palaeozoic polychaete families. Websteroprion armstrongi was a raptorial feeder and possessed the largest jaws recorded in polychaetes from the fossil record, with maxillae reaching over one centimetre in length. Total body length of the species is estimated to have reached over one metre, which is comparable to that of extant ‘giant eunicid’ species colloquially referred to as ‘Bobbit worms’. This demonstrates that polychaete gigantism was already a phenomenon in the Palaeozoic, some 400 million years ago.