An abundant and well preserved trilobite fauna is described from Upper Cambrian calcareous mudstones of the McKay Group, near Cranbrook in southeastern British Columbia. The trilobites are mostly articulated, consisting of similar numbers of molts and carcasses. They are representative of a new deep water biofacies, the Wujiajiania Biofacies, and a new Wujiajiania sutherlandi Fauna of late Steptoean age. The trilobites were collected from a narrow interval (<20 m thick) of richly fossiliferous strata, in a thick sequence of unfossiliferous to sparsely fossiliferous strata of similar lithology. The fauna includes fourteen species, six of which are new: Aciculolenus palmeri, Burnetiella leechi, Hedinaspis canadensis, Labiostria westropi, Pterocephalia norfordi, and Wujiajiania sutherlandi.
Protaspides and later growth stages are described and discussed for the Ordovician trinucleid trilobite Cryptolithus tesselatus Green and the raphiophorid Lonchodomas chaziensis Shaw. A small triangular rostral plate is described for a single protaspid instar in both Cryptolithus and Lonchodomas. The presence of this small sclerite in the ontogeny of these taxa supports the origin of the Trinucleoidea from the Ptychopariida through reduction in and then fusion or loss of the rostral plate. Earlier and later growth stages have fused ventral sclerites, with no signs of connective sutures. All members of the Trinucleoidea with known ontogenies share similar small, ovoid asaphoid protaspides, with distinct axes and varying numbers of sharp, conical to subtubular, submarginal spines on the dorsal exoskeleton and marginal spines on the hypostome. Two protaspid instars are identified in both Cryptolithus and Lonchodomas, sharing many characters that indicate homologous levels of development. A single origin for the median suture of the Asaphida is supported, with an anterior rostellum as its precursor.
Differences in the larval ecology of Ordovician trilobites directly influenced the outcome of the Ashgill extinction (latest Ordovician) and indirectly governed the pattern of evolution in post-Ordovician trilobites. Larval ecology also affected survivorship patterns within the Ordovician, particularly between the Llandeilo and the Caradoc stages. All taxa with pelagic adults became extinct by the end of the Ordovician. Similarly, trilobites with entirely planktonic larvae had all but disappeared by the end of the Ordovician. Although suffering a significant loss in diversity, taxa with benthic larvae provided the ancestral stock for the majority of post-Ordovician lineages. Trilobites with a two-stage protaspid period (single planktonic followed by benthic larvae) suffered least during the Ashgill extinction, giving rise to approximately 23% of the new genera appearing in the early Silurian.Patterns of extinction/survivorship among trilobite taxa with different developmental strategies indicate that the so-called Ashgill extinction was most likely the result of a composite phenomenon, including environmental perturbations, ecosystem breakdown and biogeographic restriction, and was not the consequence of a single catastrophic event (e.g., bolide collision). Indeed, our results are consistent with extinction models which invoke periodic global cooling and sea level regression associated with glaciation. Taxa with planktonic larvae first suffered a marked decline in diversity between the Llandeilo and the Caradoc (end of Middle Ordovician), coincident with the onset of the Ordovician-Silurian glaciation. During this time interval, the paleogeographic ranges of most surviving genera with solely planktonic larvae were severely constricted to lower latitude paleobiogeographic provinces. During the Ashgill (Rawtheyan to Hirnantian), climatic and sea level fluctuations were most extreme because of continued and more extensive glaciation. At this time, when extinctions were particularly severe, there was also a significant effect upon the survivorship of taxa with benthic larvae.Our results indicate that the proportion of the life cycle spent in the water column, dependent upon a planktonic food source, was a critical factor in the survivorship of trilobite taxa during Middle to Late Ordovician time. Consequently, differences in life history patterns predisposed individual taxa to survival or extinction.
Three types of alimentary canals (=midgut) occur in the Annelida and non-trilobite Arthropoda: 1) a sagittal tube with metamerically paired diverticula related to the number of somites; 2) a tube that is constricted slightly between somites; and 3) a simple tubular gut that may taper slightly backwards to the anus. At least two of these three types (1 and 3) occur in the Trilobita. Pterocephalia and Olenoides share the first type with the probable sister taxon to the Trilobita, Naraoia (Nectaspida), and this is probably the plesiomorphic condition for the class. Varying feeding habits may well have made this character homoplastic within each of these groups. The preservation of parts of the alimentary tract in specimens of Upper Cambrian Pterocephalia n. sp. (McKay Group, British Columbia) was probably a function of taphonomic and/or very early diagenetic changes that resulted from the type of food preferred by that trilobite. Other trilobites from the same beds do not have their soft parts preserved. The alimentary structures are preserved in a different fashion from, apparently unattached to, and an order of magnitude larger than genal caeca that occur in this taxon. Thus, genal caeca are regarded as imprints of circulatory rather than alimentary structures.Energy dispersive analysis of a fragment of preserved alimentary tract of Pterocephalia n. sp. showed the presence of Ca, Si, Al, Fe, P, K, Na, and Cl. These alimentary tracts are composed of a complex mixture of minerals that probably includes clays, detrital quartz, carbonates, phosphates, and oxides or hydroxides. The structure of these dark fillings is microcrystalline. The presence of detrital minerals as part of this mixture would suggest that this trilobite was a deposit feeder.
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