Mark D. Sutton

Arthropod fossil data increase congruence of morphological and molecular phylogenies

The relationships of major arthropod clades have long been contentious, but refinements in molecular phylogenetics underpin an emerging consensus. Nevertheless, molecular phylogenies have recovered topologies that morphological phylogenies have not, including the placement of hexapods within a paraphyletic Crustacea, and an alliance between myriapods and chelicerates. Here we show enhanced congruence between molecular and morphological phylogenies based on 753 morphological characters for 309 fossil and Recent panarthropods. We resolve hexapods within Crustacea, with remipedes as their closest extant relatives, and show that the traditionally close relationship between myriapods and hexapods is an artefact of convergent character acquisition during terrestrialisation. The inclusion of fossil morphology mitigates long-branch artefacts as exemplified by pycnogonids: when fossils are included, they resolve with euchelicerates rather than as a sister taxon to all other euarthropods.

Tomographic techniques for the study of exceptionally preserved fossils

Three-dimensional fossils, especially those preserving soft-part anatomy, are a rich source of palaeontological information; they can, however, be difficult to work with. Imaging of serial planes through an object (tomography) allows study of both the inside and outside of three-dimensional fossils. Tomography may be performed using physical grinding or sawing coupled with photography, through optical techniques of serial focusing, or using a variety of scanning technologies such as neutron tomography, magnetic resonance imaging and most usefully X-ray computed tomography. This latter technique is applicable at a variety of scales, and when combined with a synchrotron X-ray source can produce very high-quality data that may be augmented by phase-contrast information to enhance contrast. Tomographic data can be visualized in several ways, the most effective of which is the production of isosurface-based ‘virtual fossils’ that can be manipulated and dissected interactively.

Brood care in a Silurian ostracod

An exceptionally preserved new ostracod crustacean from the Silurian of Herefordshire, England, preserves eggs and possible juveniles within its carapace, providing an unequivocal and unique view of parental brood care in the invertebrate fossil record. The female fossil is assigned to a new family and superfamily of myodocopids based on its soft-part anatomy. It demonstrates a remarkably conserved egg-brooding reproductive strategy within these ostracods over 425 Myr. The soft-tissue anatomy urges extreme caution in classifying ‘straight-hinged’ Palaeozoic ostracods based on the carapace alone and fundamentally questions the nature of the shell-based Palaeozoic ostracod record.

Cambrian bivalved arthropod reveals origin of arthrodization

Extant arthropods are diverse and ubiquitous, forming a major constituent of most modern ecosystems. Evidence from early Palaeozoic Konservat Lagerstätten indicates that this has been the case since the Cambrian. Despite this, the details of arthropod origins remain obscure, although most hypotheses regard the first arthropods as benthic predators or scavengers such as the fuxianhuiids or megacheirans (‘great-appendage’ arthropods). Here, we describe a new arthropod from the Tulip Beds locality of the Burgess Shale Formation (Cambrian, series 3, stage 5) that possesses a weakly sclerotized thorax with filamentous appendages, encased in a bivalved carapace, and a strongly sclerotized, elongate abdomen and telson. A cladistic analysis resolved this taxon as the basal-most member of a paraphyletic grade of nekto-benthic forms with bivalved carapaces. This grade occurs at the base of Arthropoda (panarthropods with arthropodized trunk limbs) and suggests that arthrodization (sclerotization and jointing of the exoskeleton) evolved to facilitate swimming. Predatory and fully benthic habits evolved later in the euarthropod stem-lineage and are plesiomorphically retained in pycnogonids (sea spiders) and euchelicerates (horseshoe crabs and arachnids).

The arthropod Offacolus kingi (Chelicerata) from the Silurian of Herefordshire, England: computer based morphological reconstructions and phylogenetic affinities

The small, non–biomineralized, three–dimensionally preserved arthropod Offacolus kingi Orr et al. from the Wenlock Series (Silurian) of Herefordshire, England, is re–evaluated, and the new family Offacolidae erected. This new study is based on specimens which have been serially ground, reconstructed by computer and rendered in the round as coloured models. Offacolus possesses a prosomal appendage array similar to that of Limulus, but also bears robust and setose exopods on appendages II–V which are unlike those found in any other arthropods. Opisthosomal appendages are similar in number and morphology to the book–gills of Limulus. Cladistic analysis places Offacolus basally within the Chelicerata, as a sister taxon to the eurypterids and extant chelicerates, but more derived than the Devonian Weinbergina.

Deep molluscan phylogeny: synthesis of palaeontological and neontological data.

The position of the earliest-derived living molluscs, the Polyplacophora (chitons) and shell-less vermiform Aplacophora, remains highly contentious despite many morphological, developmental and molecular studies of extant organisms. These two groups are thought to represent either a basal molluscan grade or a clade (Aculifera) sister to the ‘higher’ molluscs (Conchifera). These incompatible hypotheses result in very different predictions about the earliest molluscs. A new cladistic analysis incorporating both Palaeozoic and extant molluscs is presented here. Our results support the monophyly of Aculifera and suggest that extant aplacophorans and polyplacophorans both derive from a disparate group of multivalved molluscs in two major clades. Reanalysis of the critical Ordovician taxon ‘Helminthochiton’ thraivensis shows that this animal lacks a true foot despite bearing polyplacophoran-like valves. Its position within our phylogenetic reconstruction indicates that many fossil ‘polyplacophorans’ in the order Palaeoloricata are likely to represent footless stem-group aplacophorans. ‘H.’ thraivensis and similar forms such as Acaenoplax may be morphological stepping stones between chitons and the shell-less aplacophorans. Our results imply that crown-group molluscan synapomorphies include serial repetition, the presence of a foot, a mineralized scleritome and a creeping rather than worm-like mode of life.

A Silurian sea spider

Pycnogonids (sea spiders) are marine arthropods numbering some 1,160 extant species. They are globally distributed in depths of up to 6,000 metres, and locally abundant; however, their typically delicate form and non-biomineralized cuticle has resulted in an extremely sparse fossil record that is not accepted universally. There are two opposing views of their phylogenetic position: either within Chelicerata as sister group to the euchelicerates, or as a sister taxon to all other euarthropods. The Silurian Herefordshire Konservat-Lagerstätte in England (∼ 425 million years (Myr) bp) yields exceptionally preserved three-dimensional fossils that provide unrivalled insights into the palaeobiology of a variety of invertebrates. The fossils are preserved as calcitic void in-fills in carbonate concretions within a volcaniclastic horizon, and are reconstructed digitally. Here we describe a new pycnogonid from this deposit, which is the oldest adult sea spider by ∼35 Myr and the most completely known fossil species. The large chelate first appendage is consistent with a chelicerate affinity for the pycnogonids. Cladistic analyses place the new species near the base of the pycnogonid crown group, implying that the latter had arisen by the Silurian period.

A new phyllocarid (Crustacea: Malacostraca) from the Silurian Fossil–Lagerstätte of Herefordshire, UK

A new three–dimensionally preserved arthropod, Cinerocaris magnifica, from the Wenlock Series (Silurian) of Herefordshire, UK, is described and assigned to the Phyllocarida (Crustacea). The description and reconstruction are based on specimens that have been serially ground, reconstructed by computer and rendered in three dimensions as coloured virtual models. Cinerocaris magnifica displayed the tagmosis characteristic of phyllocarids, with eight thoracic and seven abdominal somites, terminating in a telson with furca. The remarkable preservation of the appendages makes this the earliest completely known malacostracan crustacean. Two pairs of antennae (the first with two flagella) were followed by a mandible and first maxilla, each with a slender palp–like ramus. The second maxilla consisted of a limb stem with endites and an endopod that tapered distally. There was no exopod. The thoracopods comprised a limb stem with six or seven endites, an arrangement previously known only in entomostracans, and an endopod with about five endites. Flap–like outer rami correspond to an exopod and epipods. The pleopods bore two long slender oar–blade–like rami. Cladistic analysis places C. magnifica as a plesion within the Echinocaridina. It provides critical evidence of the limb morphology of an early malacostracan, which will be important in understanding crustacean evolution.

A Silurian armoured aplacophoran and implications for molluscan phylogeny

The Mollusca is one of the most diverse, important and well-studied invertebrate phyla; however, relationships among major molluscan taxa have long been a subject of controversy. In particular, the position of the shell-less vermiform Aplacophora and its relationship to the better-known Polyplacophora (chitons) have been problematic: Aplacophora has been treated as a paraphyletic or monophyletic group at the base of the Mollusca, proximate to other derived clades such as Cephalopoda, or as sister group to the Polyplacophora, forming the clade Aculifera. Resolution of this debate is required to allow the evolutionary origins of Mollusca to be reconstructed with confidence. Recent fossil finds support the Aculifera hypothesis, demonstrating that the Palaeozoic-era palaeoloricate ‘chitons’ included taxa combining certain polyplacophoran and aplacophoran characteristics. However, fossils combining an unambiguously aplacophoran-like body with chiton-like valves have remained elusive. Here we describe such a fossil, Kulindroplax perissokomos gen. et sp. nov., from the Herefordshire Lagerstätte (about 425 million years bp), a Silurian deposit preserving a marine biota in unusual three-dimensional detail. The specimen is reconstructed three-dimensionally through physical–optical tomography. Phylogenetic analysis indicates that this and many other palaeoloricate chitons are crown-group aplacophorans.

Silurian horseshoe crab illuminates the evolution of arthropod limbs

The basic arrangement of limbs in euarthropods consists of a uniramous head appendage followed by a series of biramous appendages. The body is divided into functional units or tagmata which are usually distinguished by further differentiation of the limbs. The living horseshoe crabs are remnants of a much larger diversity of aquatic chelicerates. The limbs of the anterior and posterior divisions of the body of living horseshoe crabs differ in the loss of the outer and inner ramus, respectively, of an ancestral biramous limb. Here we report a new fossil horseshoe crab from the mid-Silurian Lagerstätte in Herefordshire, United Kingdom (approximately 425 Myr B.P.), a site that has yielded a remarkably preserved assemblage of soft-bodied fossils. The limbs of the new form can be homologized with those of living Limulus, but retain an ancestral biramous morphology. Remarkably, however, the two limb branches originate separately, providing fossil evidence to suggest that repression or loss of gene expression might have given rise to the appendage morphology of Limulus. Both branches of the prosomal limbs of this new fossil are robust and segmented in contrast to their morphology in Cambrian arthropods, revealing that a true biramous limb was once present in chelicerates as well as in the mandibulates.