Trilobite tridents demonstrate sexual combat at 400 Mya.

The Devonian trilobite Walliserops carries a remarkable anterior cephalic trident posing a challenge to functional interpretation. A unique teratological specimen of Walliserops trifurcatus showing four, rather than three tines, is inconsistent with possible hypotheses connecting the trident to feeding techniques and suggests a sexually selected function. Malformations in a variety of living organisms support this conclusion. Morphometric comparisons to similar structures used for intraspecific combat in dynastine beetles show that the trident occupies a comparable shape space consistent with the hypothesis that it was a sexual combat weapon, the oldest reported example of its kind. This lends further credibility to the idea that some trilobites may have been strongly sexually dimorphic.

Revision of F. R. C. Reeds Ordovician trilobite types from Myanmar (Burma) and western Yunnan Province, China.

The field collections made from Burma (Myanmar) by the Geological Survey of India, and described by F.R.C. Reed more than a century ago, still provide the only ground truthing for an important region of the Ordovician marginal terranes fringing Gondwana. A revision of these faunas is overdue, particularly as it is likely that further collections cannot be made in the northern Shan State in the near future. The specimens, stored in the Geological Survey of India collections in Kolkata, cannot be loaned. Sixteen species are fully revised herein; another twelve species are left under open nomenclature because of inadequacies in the material. Several of Reeds species subsequently became type species of genera that have proved to be widespread: Birmanites Sheng, 1934, Encrinurella Reed, 1915, Neseuretinus Dean, 1967, and Pliomerina Chugaeva, 1956. Reeds Ordovician trilobite collections came from two main areas: northern Shan State (Myanmar), and westernmost Yunnan (China). The Burmese (Myanmar) collections are from the Upper Ordovician (Katian) while Yunnan specimens are from the Middle Ordovician (Darriwilian), though Upper Ordovician trilobites also occur in the area. Both collections are predominantly from clastic strata. Based on a small new Katian collection from Pupiao, we report Neseuretinus birmanicus (Reed, 1906) in common between the northern part of the Shan State and western Yunnan. A few genera (Dionide Barrande, 1847, Phorocephala Lu, 1957, Lonchodomas Angelin, 1854, Nileus Dalman, 1827) are distributed worldwide, and include pelagic (Phorocephala) or deeper benthic (Dionide) taxa. The palaeogeographic comparisons offered by the other taxa are mostly peri-Gondwanan and extend from southwest China westwards (present geography) as far as the Iberian Pennsula. Birmanites is the type genus of a subfamily (Birmanitinae Kobayashi, 1960, revived herein) widely distributed over Ordovician Gondwana, and absent from Laurentia, Baltica and North China/Siberia. Mioptychopyge Zhou, Dean, Yuan Zhou, 1998, probably belongs with the same group and is otherwise known from South China. Parillaenus Jaanusson, 1954, is also peripheral Gondwanan, as is Prionocheilus Rouault, 1847. The Reedocalymeninae Kobayashi, 1951 (Neseuretinus, Reedocalymene Kobayashi, 1951) are similarly diagnostic of periGondwanan sites. However, some genera (Pliomerina, Encrinurella, Ovalocephalus Koroleva, 1959) have been associated with other oriental and Australian occurrences in particular, with outliers in certain terranes in Kazakhstan, i.e. palaeotropical Gondwana.

A Silurian 'marrellomorph' arthropod.

Xylokorys chledophilia, a new arthropod with three-dimensionally preserved soft tissues, is described from the Herefordshire (Silurian) Lagerstätte of England. The head and trunk are covered by a relatively featureless ovoid carapace, which comprises a domed central part and a flange-like border. The head bears five pairs of appendages. The first is uniramous, with dorsal and ventral projections distally. Appendages two to four are biramous and each endopod terminates in two projections. Appendage five is possibly biramous. The hypostome is very long and subrectangular in outline. There are approximately 35 pairs of biramous trunk appendages. Each exopod comprises a long slender shaft bearing numerous fine filaments; each endopod comprises a ribbon-like shaft bearing paddle-like endites. Morphological comparisons and cladistic analyses of X. chledophilia indicate affinity with Vachonisia rogeri from the Lower Devonian Hunsrück Slate, within the marrellomorphs, but assignment to Marrellomorpha is provisional pending revision of other members of this clade. Xylokorys is the first 'marrellomorph' to be reported from the Silurian. It is interpreted as a benthic particle filter feeder, which may also have consumed prey items.

Arthropod eyes: The early Cambrian fossil record and divergent evolution of visual systems.

Four types of eyes serve the visual neuropils of extant arthropods: compound retinas composed of adjacent facets; a visual surface populated by spaced eyelets; a smooth transparent cuticle providing inwardly directed lens cylinders; and single-lens eyes. The first type is a characteristic of pancrustaceans, the eyes of which comprise lenses arranged as hexagonal or rectilinear arrays, each lens crowning 8-9 photoreceptor neurons. Except for Scutigeromorpha, the second type typifies Myriapoda whose relatively large eyelets surmount numerous photoreceptive rhabdoms stacked together as tiers. Scutigeromorph eyes are facetted, each lens crowning some dozen photoreceptor neurons of a modified apposition-type eye. Extant chelicerate eyes are single-lensed except in xiphosurans, whose lateral eyes comprise a cuticle with a smooth outer surface and an inner one providing regular arrays of lens cylinders. This account discusses whether these disparate eye types speak for or against divergence from one ancestral eye type. Previous considerations of eye evolution, focusing on the eyes of trilobites and on facet proliferation in xiphosurans and myriapods, have proposed that the mode of development of eyes in those taxa is distinct from that of pancrustaceans and is the plesiomorphic condition from which facetted eyes have evolved. But the recent discovery of enormous regularly facetted compound eyes belonging to early Cambrian radiodontans suggests that high-resolution facetted eyes with superior optics may be the ground pattern organization for arthropods, predating the evolution of arthrodization and jointed post-protocerebral appendages. Here we provide evidence that compound eye organization in stem-group euarthropods of the Cambrian can be understood in terms of eye morphologies diverging from this ancestral radiodontan-type ground pattern. We show that in certain Cambrian groups apposition eyes relate to fixed or mobile eyestalks, whereas other groups reveal concomitant evolution of sessile eyes equipped with optics typical of extant xiphosurans. Observations of fossil material, including that of trilobites and eurypterids, support the proposition that the ancestral compound eye was the apposition type. Cambrian arthropods include possible precursors of mandibulate eyes. The latter are the modified compound eyes, now sessile, and their underlying optic lobes exemplified by scutigeromorph chilopods, and the mobile stalked compound eyes and more elaborate optic lobes typifying Pancrustacea. Radical divergence from an ancestral apposition type is demonstrated by the evolution of chelicerate eyes, from doublet sessile-eyed stem-group taxa to special apposition eyes of xiphosurans, the compound eyes of eurypterids, and single-lens eyes of arachnids. Different eye types are discussed with respect to possible modes of life of the extinct species that possessed them, comparing these to extant counterparts and the types of visual centers the eyes might have served.

Trilobite spines and beetle horns: sexual selection in the Palaeozoic?

Raphiophorid trilobites commonly bore median cephalic protuberances such as spines or bulbs, showing a remarkable variety of form. It is unlikely that their primary function was for protection or in hydrodynamics. A case is made that they were secondary sexual features, by comparison with similar morphological structures developed on rhinoceros beetles and other arthropods. This interpretation is supported by four lines of evidence: their ontogeny, their diversity, the existence of plausible examples of sexual dimorphs in some cases and the fact that they show positive allometry.

Taxonomy and fossils: a critical appraisal.

Many compendia at the species, genus and family levels document the fossil record, but these are not standardized, nor usually critical in content, and few are available on the World Wide Web. The sampling of the available record is good for organisms with fossilizable parts, but preservational constraints on the entire morphology, life history and geographical distribution lead to difficulties in recognizing and naming species. We recommend abandoning some of the palaeontological species concepts such as chronospecies and stratospecies, and we advocate species recognition based on unique combinations of characters. The compilation of species lists is extremely time consuming, and given the inherent problems we suggest that compilation of generic lists is a more achievable goal because genera are recognized by definitive morphological characters. In calculating taxon duration, care must be taken to distinguish between mono-, para- and polyphyletic groups, the first being the only reliable unit for use in calculating diversity curves. We support the inclusion of fossils into classifications based on Recent organisms, but we recognize some of the problems this may pose for standard Linnaean classifications. Web-based taxonomy is the way forward, having the advantages of speed and currency of information dissemination, universal access with links to primary literature and increasingly sophisticated imagery. These advantages over conventional outlets will only be realized with careful Web design and a commitment to maintenance.