Trilobite monophyly revisited

Fortey's and Whittington's recent refutation of Lauterbach's hypothesis of a paraphyletic Trilobita is supported. However, much of the character evidence raised by Fortey and Whittington to substantiate the monophyly of the Trilobita (including, inter alia, "Olenellinae”; and Agnostoidea) is ambiguous. Of seven proposed synapomorphies, only one (dorsal cuticle calcification) may be maintained at that node after testing within a cladistic framework. The other six characters are either constrained by calcification or define nodes up or down the cladogram. As positioned by Fortey's and Whittington's characters, Agnostoidea could be regarded either as the most primitive trilobites, or as being outside that clade. Lauterbach's support for an “olenelline"‐chelicerate clade is found to include interdependent characters which are reduced here to two testable derived similarities. Only one of these may conform to general criteria indicative of homology, such as detailed similarity and topology. It is, however, rejected on the basis of parsimony. We emphasize that resolution of the chelicerate‐"olenelline"‐trilobite three‐taxon problem must be based on recognition of homologies among each of these taxa. Nectaspida are excluded from Trilobita as defined by cuticle calcification, but as ingroup “Arachnata”; (sensu Lauterbach) they are important for determining character generality in this clade.     

Trilobite larvae and larval ecology

Silicified trilobite faunas yield a well preserved and abundant record of early development in many taxa. Although their morphologies are well known, protaspid larvae are poorly understood in terms of trilobite life history and developmental biology. Two distinct morphologic types are recognized among the great diversity of protaspides studied; these are termed adult‐like and nonadult‐tike according to gross similarity to later developmental stages. Transitions between nonadult‐like and adult‐like morphologies are abrupt, occurring between successsive instars, and, thereby, constitute metamorphoses.

By analogy with developmental patterns among modern marine arthropod taxa, metamorphoses during early trilobite ontogeny correspond to radical modifications in life mode and ecology. Adult‐like trilobite protaspides possess a dorso‐ventrally flattened tergum which displays a coarsely featured prosopon and surface‐parallel spines; these are interpreted as benthic larvae. In contrast, nonadult‐like protaspid larvae display an ovoid to spheroidal shape with spine pairs that project at right angles to one another; these are considered to have been planktonic. Protaspid morphologies are compared and discussed in light of these inferred modes of life.     

Trilobite cuticle thickness in relation to palaeoenvironment

100 thickness measurements from thin sections of cephala or pygidia of early Ordovician trilobites occurring across an onshore to offshore environmental gradient show that progressively greater maximum cuticle thickness was characteristic of increasingly inshore sites. There is a 40-fold difference between the thinnest and thickest cuticles, and exclusively thin cuticles are confined to the offshore Olenid Biofacies. Variability in cuticle thickness increases offshore to onshore. Environmental control is shown to be more influential on cuticle thickness than is the overall length of the trilobite: some comparatively large trilobites having thin cuticles and small trilobites thick cuticles. The environmental factors which might be responsible for the pattern are briefly discussed. The thin cuticles dominating the offshore Olenid Biofacies were probably appropriate for dysaerobic conditions. Thick cuticles in the most inshore biofacies may have offered protection against predators and turbulence, but the additional presence there of trilobites with thinner cuticles is considered to reflect the greater heterogeneity of the epeiric habitat.     

The Trilobite Subfamily Monorakinae (Pterygometopidae)

The Monorakinae is a subfamily of the Pterygometopidae characterised by the fusion of L2 and L3 in the glabella. The resulting bicomposite lobe is expanded backwards to reach the occipital furrow, displacing L1 from contact with the axial furrow and causing the realignment of S1 to a markedly oblique orientation. The bicomposite lobe is commonly bounded adaxially by a longitudinal furrow containing three pairs of apodemal pits. The Monorakinae was probably derived from the Pterygometopinae, and includes the genera and subgenera Monorakos , Carinopyge , Ceratevenkaspis , Elasmaspis , Evenkaspis ( Evenkaspis ) and E . ( Parevenkaspis ), of which Carinopyge , Elasmaspis and Evenkaspis ( Parevenkaspis ) are known only from limited parts of the exoskeleton. Monorakines have a stratigraphical range of Caradoc–Ashgill. Their known geographical distribution in the Siberian Platform, Taimyr, the Russian Far East, and the Seward Peninsula of Alaska is restricted to areas that in the Ordovician were part of the palaeocontinents of Siberia and Arctida, which must have been connected or situated close together at that time. The occurrence of monorakines in the Taimyr Peninsula but their absence from Baltica does not support the suggestion of some workers that Taimyr was part of Baltica in the Ordovician.     

A Middle Devonian lichid Trilobite from South-Eastern Australia

A new lichid trilobite belonging to the subgenusMephiarges is described from delicately silicified fragments present in a massive Middle Devonian limestone in South-eastern Australia. Previously this subgenus was limited to one species of which only the cephalon is known.     

Trilobite body patterning and the evolution of arthropod tagmosis

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased.     

Phylogeny of the Trilobite Subgenus Acanthopyge (Lobopyge)

Cladistic analysis of the trilobite subgenus Acanthopyge (Lobopyge) has not been previously attempted, apart from an inferred phylogeny of the Lichida by Thomas and Holloway (1988, Philos. Trans. R. Soc. London B Biol. Sci. 321, 179–262). Results of two separate analyses with variable taxonomic sampling show a possible cosmopolitan affinity for Australian Devonian species of A. (Lobopyge) and corroborate the placement of A. (Lobopyge) rohri in A. (Lobopyge) . Benelopyge is a junior subjective synonym of A. (Lobopyge) .     

Trilobite tagmosis and body patterning from morphological and developmental perspectives

The Trilobita were characterized by a cephalic region in whichthe biomineralized exoskeleton showed relatively high morphologicaldifferentiation among a taxonomically stable set of well definedsegments, and an ontogenetically and taxonomically dynamic trunkregion in which both exoskeletal segments and ventral appendageswere similar in overall form. Ventral appendages were homonomousbiramous limbs throughout both the cephalon and trunk, exceptfor the most anterior appendage pair that was antenniform, preoral,and uniramous, and a posteriormost pair of antenniform cerci,known only in one species. In some clades trunk exoskeletalsegments were divided into two batches. In some, but not all,of these clades the boundary between batches coincided withthe boundary between the thorax and the adult pygidium. Therepeated differentiation of the trunk into two batches of segmentsfrom the homonomous trunk condition indicates an evolutionarytrend in aspects of body patterning regulation that was achievedindependently in several trilobite clades. The phylogeneticplacement of trilobites and congruence of broad patterns oftagmosis with those seen among extant arthropods suggest thatthe expression domains of trilobite cephalic Hox genes may haveoverlapped in a manner similar to that seen among extant arachnates.This, coupled with the fact that trilobites likely possessedten Hox genes, presents one alternative to a recent model inwhich Hox gene distribution in trilobites was equated to eightputative divisions of the trilobite body plan.