Timothy P. Topper

The scleritome of Paterimitra: an Early Cambrian stem group brachiopod from South Australia

Early Cambrian tommotiids are problematic fossil metazoans with external organophosphatic sclerites that have been considered to be basal members of the lophophorate stem group. Tommotiids are almost exclusively known from isolated or rarely fused individual sclerites, which made previous reconstructions of the actual organism highly conjectural. However, the recent discovery of the first articulated specimens of the tommotiid Eccentrotheca revealed a tubular sclerite arrangement (scleritome) that limited the possible life habit to sessile filter feeding and thus further supported a lophophorate affinity. Here, we report the first articulated specimens of a second tommotiid taxon, Paterimitra from the Early Cambrian of the Arrowie Basin, South Australia. Articulated specimens of Paterimitra are composed of two bilaterally symmetrical sclerite types and an unresolved number of small, asymmetrical and irregular crescent-shaped sclerites that attached to the anterior margin of the symmetrical sclerites. Together, the sclerites form an open cone in which the symmetrical sclerites are joined together and form a small posterior opening near the base of the scleritome, while the irregular crescent-shaped sclerites defined a broad anterior opening. The coniform scleritome of Paterimitra is interpreted to have attached to hard substrates via a pedicle that emerged through the small posterior opening (sometimes forming a tube) and was probably a sessile filter feeder. The scleritome of Paterimitra can be derived from the tubular scleritome of Eccentrotheca by modification of basal sclerites and reduction in tube height, and probably represents a more derived member of the brachiopod stem group with the paired symmetrical sclerites possibly homologous to brachiopod valves.

Palaeoscolecid scleritome fragments with Hadimopanella plates from the early Cambrian of South Australia

Phosphatized articulated palaeoscolecid scleritome fragments with attached Hadimopanella Gedik, 1977 plates are described from the lower Cambrian Mernmerna Formation of South Australia. Hadimopanella is principally known from single, isolated, button-shaped, phosphatic sclerites. The new articulated material from South Australia reveals for the first time the configuration of plates referable to Hadimopanella within the scleritome. The scleritome fragments represent the main trunk sections of the cuticle with anterior and posterior terminations lacking. Each annulus on the trunk is ornamented by rows of irregularly alternating Hadimopanella plates. The large majority of plates display a single, centrally located, conical node referable to the form species H. apicata Wrona, 1982. However, individual plates display considerable morphological variation with plates situated along the flattened trunk margin identical to the form species H. antarctica Wrona, 1987. The South Australian material displays the detailed scleritome configuration of cuticular plates and platelets and demonstrates irrefutably that plates of the form species H. apicata and H. antarctica occur as mineralized cuticular elements on the same palaeoscolecid scleritome.

The Oldest Brachiopods from the Lower Cambrian of South Australia

The morphology and organophosphatic shell structure of the paterinate brachiopod Askepasma is documented using new and previously collected specimens from the lower Cambrian of South Australia. Lack of adequately preserved material has seen the majority of paterinate specimens previously reported from South Australia referred to the genus Askepasma and treated under open nomenclature. Large collections of paterinates from the lower Cambrian Wilkawillina, Ajax, and Wirrapowie limestones in the Arrowie Basin, South Australia have prompted redescription of the type species Askepasma toddense and the erection of a new species, Askepasma saproconcha sp. nov. Askepasma saproconcha sp. nov. currently represents the oldest known brachiopod from the lower Cambrian successions in South Australia with a FAD in pre-trilobitic (Terreneuvian, Cambrian Stage 2, lower Atdabanian) strata in the basal part of the Wilkawillina and Wirrapowie limestones. Askepasma toddense predominantly occurs in Abadiella huoi Zone equivalent strata (Unnamed Cambrian Series 2, Stage 3, middle-upper Atdabanian) in the upper part of the lower Wilkawillina, Wirrapowie, and Ajax limestones. The shell microstructure of Askepasma suggests a proximal stem group position within the Brachiopoda and similarities with tommotiid taxa provides further evidence that the ancestry of crown group brachiopods is firmly entrenched within the Tommotiida.

Microdictyon plates from the lower Cambrian Ajax Limestone of South Australia : Implications for species taxonomy and diversity

A small assemblage of isolated Microdictyon plates is described from the lower Cambrian Ajax Limestone, Mt Scott Range (Flinders Ranges), South Australia. Microdictyon plates are primarily known from single, isolated, perforated phosphatic sclerites; only one species (M. sinicum) from the Chengjiang Lagerstätte is known from soft-bodied preservation of the complete organism. The isolated plates from South Australia display a wide range of morphologies potentially reflecting: 1, considerable diversification within the group at this time; 2, extensive intraspecific morphological variation; 3, different plate morphotypes along the trunk of the animal; or 4, significant ontogenetic variation in successive growth stages. The South Australian specimens are similar to several individual sclerites described from other Cambrian palaeocontinents, but appear closest to faunas described from South China. Problems associated with the taxonomy of isolated Microdictyon plates are discussed, including the lack of knowledge associated with intraspecific and/or ontogenetic variability and how individual plate morphology may relate to size or relative position along the length of the complete organism.

Brachiopods hitching a ride: an early case of commensalism in the middle Cambrian Burgess Shale

Ecological interactions, including symbiotic associations such as mutualism, parasitism and commensalism are crucial factors in generating evolutionary novelties and strategies. Direct examples of species interactions in the fossil record generally involve organisms attached to sessile organisms in an epibiont or macroboring relationship. Here we provide support for an intimate ecological association between a calcareous brachiopod (Nisusia) and the stem group mollusc Wiwaxia from the Burgess Shale. Brachiopod specimens are fixed to Wiwaxia scleritomes, the latter showing no signs of decay and disarticulation, suggesting a live association. We interpret this association as the oldest unambiguous example of a facultative ectosymbiosis between a sessile organism and a mobile benthic animal in the fossil record. The potential evolutionary advantage of this association is discussed, brachiopods benefiting from ease of attachment, increased food supply, avoidance of turbid benthic conditions, biofoul and possible protection from predators, suggesting commensalism (benefiting the symbiont with no impact for the host). While Cambrian brachiopods are relatively common epibionts, in particular on sponges, the association of Nisusia with the motile Wiwaxia is rare for a brachiopod species, fossil or living, and suggests that symbiotic associations were already well established and diversified by the “middle” (Series 3, Stage 5) Cambrian.

Competition and mimicry: the curious case of chaetae in brachiopods from the middle Cambrian Burgess Shale

BackgroundOne of the first phyla to acquire biomineralized skeletal elements in the Cambrian, brachiopods represent a vital component in unraveling the early evolution and relationships of the Lophotrochozoa. Critical to improving our understanding of lophotrochozoans is the origin, evolution and function of unbiomineralized morphological features, in particular features such as chaetae that are shared between brachiopods and other lophotrochozoans but are poorly understood and rarely preserved. Micromitra burgessensis and Paterina zenobia from the middle Cambrian Burgess Shale are among the most remarkable examples of fossilized chaetae-bearing brachiopods. The form, functional morphology, evolutionary and ecological significance of their chaetae are studied herein.ResultsLike in Recent forms, the moveable but semi-rigid chaetae fringe both the dorsal and ventral mantle margins, but in terms of length, the chaetae of Burgess Shale taxa can exceed twice the maximum length of the shell from which it projects. This is unique amongst Recent and fossil brachiopod taxa and given their size, prominence and energy investment to the organism certainly had an important functional significance. Micromitra burgessensis individuals are preserved on hard skeletal elements, including conspecific shells, Tubulella and frequently on the spicules of the sponge Pirania muricata, providing direct evidence of an ecological association between two species. Morphological analysis and comparisons with fossil and extant brachiopod chaetae point to a number of potential functions, including sensory, defence, feeding, defouling, mimicry and spatial competition.ConclusionsOur study indicates that it is feasible to link chaetae length to the lack of suitable substrate in the Burgess Shale environment and the increased intraspecific competition associated with this. Our results however, also lend support to the elongated chaetae as an example of Batesian mimicry, of the unpalatable sponge Pirania muricata. We also cannot discount brachiopod chaetae acting as a sensory grille, extending the tactile sensitivity of the mantle into the environment, as an early warning system to approaching predators.

Associated conchs and opercula of Triplicatella disdoma (Hyolitha) from the early Cambrian of South Australia

Internal moulds of complete shells, including conch and associated opercula, of the hyolith Triplicatella disdoma from Cambrian Series 2, Stages 3–4 of South Australia are described. The conch of T. disdoma is shown to be narrow and cone-shaped, and with a rounded triangular cross-section. The conch shows a gentle dorsal curvature in lateral view. The fossils lack evidence of helens, and the operculum was smaller than the apertural diameter of the conch and could be withdrawn a short distance into the conch. Triplicatella was probably closely related to orthothecid hyoliths.

Ancestral billingsellides and the evolution and phylogenetic relationships of early rhynchonelliform brachiopods

The relationship of many Cambrian rhynchonelliform brachiopods is poorly understood, with many genera displaying a combination of morphological features that are taxonomically confusing. The study of middle Cambrian–early Tremadocian brachiopods is critical because this interval is sandwiched directly between the two largest radiation phases in the early Palaeozoic and provides raw data for deciphering the events leading up to the explosion of brachiopod genera in the Ordovician. Here we present a parsimony analysis of a wide selection of Cambrian and Ordovician brachiopod genera with a particular focus on the evolution and phylogeny of Billingsellida. The billingselloids were widespread by the late Cambrian and the group was originally thought to represent the ancestral stock of many Ordovician brachiopod lineages. The phylogenetic analyses portray the polytoechioids as derived billingselloids separate from the clitambonitoids that form a sister group. The Gondwanan brachiopod Roanella is interpreted as ancestral to the clitambonitoids within the Billingsellida and is reassigned to Clitambonitoidea within a new monogeneric family, Roanellidae nov. Antigonambonites displays no obvious relationship with the clitambonitoids and should be formally transferred to the polytoechioids. The monogeneric family Chaniellidae exhibits characters reminiscent of members of the polytoechioids and is transferred to the superfamily Polytoechioidea. The recently reappraised clitambonitoid Arctohedra is interpreted as a basal member of the entire order Billingsellida.

Reappraisal of the brachiopod Acrotreta socialis von Seebach, 1865: clarifying 150 years of confusion

The acrotretide Acrotreta socialis von Seebach, 1865 has suffered a long and tumultuous history since its original description from the middle Cambrian (Guzhangian Stage) Andrarum Limestone of Bornholm, Denmark. Uncertainties regarding key morphological characters have resulted in numerous revisions and redescriptions of the species with much of the taxonomic shuffling occurring before the mid-20th century. A major problem with the early documentation of acrotretide brachiopods is that many species were described in the days before the advent of the Scanning Electron Microscope (SEM), resulting in dubious interpretations of significant morphological characteristics, like the position of the pedicle foramen and the internal morphology of both valves. A detailed revision of Acrotreta species resulted in the proposal that the genus is restricted solely to the Ordovician with the Bornholm acrotretide species subsequently left floating in taxonomic limbo. Recent taxonomic suggestions have been provisional, awaiting morphological information based on type or topotype material. The examination of topotype material in addition to supplementary material collected from southern Sweden has elucidated many of the previous ambiguous morphological characteristics of the species confirming that the original description was based on more than one brachiopod species. The new morphological information acquired here has resulted in the erection of a new acrotretid genus, Clupeafumosus gen. nov., to accommodate the material previously described as Acrotreta socialis. Co-occurring with Clupeafumosus socialis in southern Sweden is another new acrotretide brachiopod, Monophthalma andersoni sp. nov., which is additionally described.

The early Cambrian tommotiid Kulparina rostrata from South Australia

Abstract. The early Cambrian (Terreneuvian, Stage 2) tommotiid Kulparina rostrata Conway Morris and Bengtson in Bengtson et al., 1990 is revised. The pyramidal sclerites of K. rostrata are shown to be bilaterally symmetrical and homologues of the symmetrical S1 sclerites of Paterimitra pyramidalis Laurie, 1986. The scleritome of K. rostrata is also shown to include flattened asymmetrical sclerites that were originally described under the name Eccentrotheca guano Bengtson in Bengtson et al., 1990 and which correspond to the L-sclerites of Paterimitra. A modified tubular scleritome and a sessile filter-feeding mode of life is envisaged for Kulparina rostrata.