Preserved appendages in a Silurian binodicope: implications for the evolutionary history of ostracod crustaceans.

Ostracod crustaceans originated at least 500 Ma ago. Their tiny bivalved shells represent the most species-abundant fossil arthropods, and ostracods are omnipresent in a wide array of freshwater and marine environments today and in the past. Derima paparme gen. et sp. nov. from the Herefordshire Silurian Lagerstätte (~430 Ma) in the Welsh Borderland, UK, is one of only a handful of exceptionally preserved ostracods (with soft parts as well as the shell) known from the Palaeozoic. A male specimen provides the first evidence of the appendages of Binodicopina, a major group of Palaeozoic ostracods comprising some 135 Ordovician to Permian genera. The appendage morphology of D. paparme, but not its shell, indicates that binodicopes belong to Podocopa. The discovery that the soft-part morphology of binodicopes allies them with podocopes affirms that using the shell alone is an unreliable basis for classifying certain fossil ostracods, and knowledge of soft-part morphology is critical for the task. Current assignment of many fossil ostracods to higher taxa, and therefore the evolutionary history of the group, may require reconsideration.

A vicissicaudatan arthropod from the Silurian Herefordshire Lagerstätte, UK.

A new arthropod, Carimersa neptuni gen. et sp. nov., is described from the Silurian (Wenlock Series) Herefordshire Konservat-Lagerstätte, UK. The head bears pedunculate eyes and five pairs of appendages. Triflagellate antennae are followed by two pairs of uniramous limbs each with an endopod bearing a pronounced gnathobasic basipod. The posterior two pairs of head limbs and all trunk limbs bear an endopod, exopod and filamentous exite. The trunk consists of 10 appendage-bearing segments followed by an apodous abdomen of four segments. The arthropod resolves as sister taxon to Kodymirus and Eozetetes + Aglaspidida. It is the first representative of Vicissicaudata reported from the Herefordshire Lagerstätte and the first Silurian example with well-preserved appendages. The preservation of a cluster of radiolarians apparently captured by the trunk appendages is the first direct association of predator and prey discovered in the Herefordshire fauna, and suggests that Carimersa was a nektobenthic form that used its gnathobasic basipods in microdurophagy.

Variation in appendages in early Cambrian bradoriids reveals a wide range of body plans in stem-euarthropods.

Traditionally, the origin and evolution of modern arthropod body plans has been revealed through increasing levels of appendage specialisation exhibited by Cambrian euarthropods. Here we show significant variation in limb morphologies and patterns of limb-tagmosis among three early Cambrian arthropod species conventionally assigned to the Bradoriida. These arthropods are recovered as a monophyletic stem-euarthropod group (and sister taxon to crown-group euarthropods, i.e. Chelicerata, Mandibulata and their extinct relatives), thus implying a radiation of stem-euarthropods where trends towards increasing appendage specialisation were explored convergently with other euarthropod groups. The alternative solution, where bradoriids are polyphyletic, representing several independent origins of a small, bivalved body plan in lineages from diverse regions of the euarthropod and mandibulate stems, is only marginally less parsimonious. The new data reveal a previously unknown disparity of body plans in stem-euarthropods and both solutions support remarkable evolutionary convergence, either of fundamental body plans or appendage specialization patterns.

Three-dimensionally preserved soft tissues and calcareous hexactins in a Silurian sponge: implications for early sponge evolution.

Sponges (Porifera), as one of the earliest-branching animal phyla, are crucial for understanding early metazoan phylogeny. Recent studies of Lower Palaeozoic sponges have revealed a variety of character states and combinations unknown in extant taxa, challenging our views of early sponge morphology. The Herefordshire Konservat-Lagerstätte yields an abundant, diverse sponge fauna with three-dimensional preservation of spicules and soft tissue. Carduispongia pedicula gen. et sp. nov. possesses a single layer of hexactine spicules arranged in a regular orthogonal network. This spicule type and arrangement is characteristic of the reticulosans, which have traditionally been interpreted as early members of the extant siliceous Class Hexactinellida. However, the unusual preservation of the spicules of C. pedicula reveals an originally calcareous composition, which would be diagnostic of the living Class Calcarea. The soft tissue architecture closely resembles the complex sylleibid or leuconid structure seen in some modern calcareans and homoscleromorphs. This combination of features strongly supports a skeletal continuum between primitive calcareans and hexactinellid siliceans, indicating that the last common ancestor of Porifera was a spiculate, solitary, vasiform animal with a thin skeletal wall.

A new ophiocistioid with soft-tissue preservation from the Silurian Herefordshire Lagerstätte, and the evolution of the holothurian body plan.

Reconstructing the evolutionary assembly of animal body plans is challenging when there are large morphological gaps between extant sister taxa, as in the case of echinozoans (echinoids and holothurians). However, the inclusion of extinct taxa can help bridge these gaps. Here we describe a new species of echinozoan, Sollasina cthulhu, from the Silurian Herefordshire Lagerstätte, UK. Sollasina cthulhu belongs to the ophiocistioids, an extinct group that shares characters with both echinoids and holothurians. Using physical-optical tomography and computer reconstruction, we visualize the internal anatomy of S. cthulhu in three dimensions, revealing inner soft tissues that we interpret as the ring canal, a key part of the water vascular system that was previously unknown in fossil echinozoans. Phylogenetic analyses strongly suggest that Sollasina and other ophiocistioids represent a paraphyletic group of stem holothurians, as previously hypothesized. This allows us to reconstruct the stepwise reduction of the skeleton during the assembly of the holothurian body plan, which may have been controlled by changes in the expression of biomineralization genes.

A well-preserved respiratory system in a Silurian ostracod.

Ostracod crustaceans are diverse and ubiquitous in aqueous environments today but relatively few known species have gills. Ostracods are the most abundant fossil arthropods but examples of soft-part preservation, especially of gills, are exceptionally rare. A new ostracod, Spiricopia aurita (Myodocopa), from the marine Silurian Herefordshire Lagerstätte (430 Mya), UK, preserves appendages, lateral eyes and gills. The respiratory system includes five pairs of gill lamellae with hypobranchial and epibranchial canals that conveyed haemolymph. A heart and associated vessels had likely evolved in ostracods by the Mid-Silurian.

A three-dimensionally preserved lobopodian from the Herefordshire (Silurian) Lagerstätte, UK.

The Herefordshire (Silurian) Lagerstätte (approx. 430 Myr BP) has yielded, among many exceptionally preserved invertebrates, a wide range of new genera belonging to crown-group Panarthropoda. Here, we increase this panarthropod diversity with the lobopodian Thanahita distos, a new total-group panarthropod genus and species. This new lobopodian preserves at least nine paired, long, slender appendages, the anterior two in the head region and the posterior seven representing trunk lobopods. The body ends in a short post-appendicular extension. Some of the trunk lobopods bear two claws, others a single claw. The body is covered by paired, tuft-like papillae. Thanahita distos joins only seven other known three-dimensionally preserved lobopodian or onychophoran (velvet worm) fossil specimens and is the first lobopodian to be formally described from the Silurian. Phylogenetic analysis recovered it, together with all described Hallucigenia species, in a sister-clade to crown-group panarthropods. Its placement in a redefined Hallucigeniidae, an iconic Cambrian clade, indicates the survival of this clade to Silurian times.

Naked chancelloriids from the lower Cambrian of China show evidence for sponge-type growth.

Chancelloriids are an extinct group of spiny Cambrian animals of uncertain phylogenetic position. Despite their sponge-like body plan, their spines are unlike modern sponge spicules, but share several features with the sclerites of certain Cambrian bilaterians, notably halkieriids. However, a proposed homology of these 'coelosclerites' implies complex transitions in body plan evolution. A new species of chancelloriid, Allonnia nuda, from the lower Cambrian (Stage 3) Chengjiang Lagerstätte is distinguished by its large size and sparse spination, with modified apical sclerites surrounding an opening into the body cavity. The sclerite arrangement in A. nuda and certain other chancelloriids indicates that growth involved sclerite addition in a subapical region, thus maintaining distinct zones of body sclerites and apical sclerites. This pattern is not seen in halkieriids, but occurs in some modern calcarean sponges. With scleritome assembly consistent with a sponge affinity, and in the absence of cnidarian- or bilaterian-grade features, it is possible to interpret chancelloriids as sponges with an unusually robust outer epithelium, strict developmental control of body axis formation, distinctive spicule-like structures and, by implication, minute ostia too small to be resolved in fossils. In this light, chancelloriids may contribute to the emerging picture of high disparity among early sponges.

Host-specific infestation in early Cambrian worms.

Symbiotic relationships are widespread in terrestrial and aquatic animals today, but evidence of symbiosis in the fossil record between soft-bodied bilaterians where the symbiont is intimately associated with the integument of the host is extremely rare. The radiation of metazoan life apparent in the Ediacaran (~635-541 million years ago) and Cambrian (~541-488 million years ago) periods is increasingly accepted to represent ecological diversification resulting from earlier key genetic developmental events and other innovations that occurred in the late Tonian and Cryogenian periods (~850-635 million years ago). The Cambrian has representative animals in each major ecospace category, the early Cambrian in particular having witnessed the earliest known complex animal communities and trophic structures, including symbiotic relationships. Here we report on newly discovered Cricocosmia and Mafangscolex worms that are hosts to aggregates of a new species of tiny worm in the lower Cambrian (Series 2, Stage 3) Chengjiang Lagerstätte of Yunnan Province, southwest China. The worm associations suggest the earliest known record of aggregate infestation of the integument of a soft-bodied bilaterian, host specificity and host shift.

An edrioasteroid from the Silurian Herefordshire Lagerstätte of England reveals the nature of the water vascular system in an extinct echinoderm.

Echinoderms are unique in having a water vascular system with tube feet, which perform a variety of functions in living forms. Here, we report the first example of preserved tube feet in an extinct group of echinoderms. The material, from the Silurian Herefordshire Lagerstätte, UK, is assigned to a new genus and species of rhenopyrgid edrioasteroid, Heropyrgus disterminus The tube feet attach to the inner surface of compound interradial plates and form two sets, an upper and a lower, an arrangement never reported previously in an extant or extinct echinoderm. Cover plates are absent and floor plates are separated creating a large permanent entrance to the interior of the oral area. The tube feet may have captured food particles that entered the oral area and/or enhanced respiration. The pentameral symmetry of the oral surface transitions to eight columns in which the plates are vertically offset resulting in a spiral appearance. This change in symmetry may reflect flexibility in the evolutionary development of the axial and extraxial zones in early echinoderm evolution.

A new crustacean from the Herefordshire (Silurian) Lagerstätte, UK, and its significance in malacostracan evolution.

Cascolus ravitis gen. et sp. nov. is a three-dimensionally preserved fossil crustacean with soft parts from the Herefordshire (Silurian) Lagerstätte, UK. It is characterized by a head with a head shield and five limb pairs, and a thorax (pereon) with nine appendage-bearing segments followed by an apodous abdomen (pleon). All the appendages except the first are biramous and have a gnathobase. The post-mandibular appendages are similar one to another, and bear petal-shaped epipods that probably functioned as a part of the respiratory-circulatory system. Cladistic analysis resolves the new taxon as a stem-group leptostracan (Malacostraca). This well-preserved arthropod provides novel insights into the evolution of appendage morphology, tagmosis and the possible respiratory-circulatory physiology of a basal malacostracan.

Tiny individuals attached to a new Silurian arthropod suggest a unique mode of brood care.

The ∼430-My-old Herefordshire, United Kingdom, Lagerstätte has yielded a diversity of remarkably preserved invertebrates, many of which provide fundamental insights into the evolutionary history and ecology of particular taxa. Here we report a new arthropod with 10 tiny arthropods tethered to its tergites by long individual threads. The head of the host, which is covered by a shield that projects anteriorly, bears a long stout uniramous antenna and a chelate limb followed by two biramous appendages. The trunk comprises 11 segments, all bearing limbs and covered by tergites with long slender lateral spines. A short telson bears long parallel cerci. Our phylogenetic analysis resolves the new arthropod as a stem-group mandibulate. The evidence suggests that the tethered individuals are juveniles and the association represents a complex brooding behavior. Alternative possibilities-that the tethered individuals represent a different epizoic or parasitic arthropod-appear less likely.

A 425-million-year-old silurian pentastomid parasitic on ostracods.

Pentastomids (tongue worms) are worm-like arthropods known today from ∼140 species [1]. All but four are parasitic on vertebrates. Their life cycle typically involves larval development in an intermediate host followed by maturation in the respiratory tract of a definitive terrestrial host. Fossil pentastomids are exceedingly rare and are known only from isolated juveniles [2-6]. The identity of the possible hosts of fossil pentastomids and the origin of their lifestyle have generated much debate. A new, exceptionally preserved species, described based on adults from 425-million-year-old marine rocks, is the only known fossil pentastomid associated with a host, in this case a species of ostracod crustacean. The pentastomids are preserved near eggs within the ostracod and also, uniquely for any fossil or living pentastomid, are attached externally to the host. This discovery affirms the origin of pentastomids as ectoparasitic on marine invertebrates. The terrestrialization of pentastomids may have occurred in parallel with the vertebrate invasion of land.

A chancelloriid-like metazoan from the early Cambrian Chengjiang Lagerstätte, China.

Nidelric pugio gen. et sp. nov. from the Cambrian Series 2 Heilinpu Formation, Chengjiang Lagerstätte, Yunnan Province, China, is an ovoid, sac-like metazoan that bears single-element spines on its surface. N. pugio shows no trace of a gut, coelom, anterior differentiation, appendages, or internal organs that would suggest a bilateral body plan. Instead, the sac-like morphology invites comparison with the radially symmetrical chancelloriids. However, the single-element spines of N. pugio are atypical of the complex multi-element spine rosettes borne by most chancelloriids and N. pugio may signal the ancestral chancelloriid state, in which the spines had not yet fused. Alternatively, N. pugio may represent a group of radial metazoans that are discrete from chancelloriids. Whatever its precise phylogenetic position, N. pugio expands the known disparity of Cambrian scleritome-bearing animals, and provides a new model for reconstructing scleritomes from isolated microfossils.

Exceptionally preserved 450-million-year-old ordovician ostracods with brood care.

Ostracod crustaceans are the most abundant fossil arthropods and are characterized by a long stratigraphic range. However, their soft parts are very rarely preserved, and the presence of ostracods in rocks older than the Silurian period [1-5] was hitherto based on the occurrence of their supposed shells. Pyritized ostracods that preserve limbs and in situ embryos, including an egg within an ovary and possible hatched individuals, are here described from rocks of the Upper Ordovician Katian Stage Lorraine Group of New York State, including examples from the famous Beecher's Trilobite Bed [6, 7]. This discovery extends our knowledge of the paleobiology of ostracods by some 25 million years and provides the first unequivocal demonstration of ostracods in the Ordovician period, including the oldest known myodocope, Luprisca incuba gen. et sp. nov. It also provides conclusive evidence of a developmental brood-care strategy conserved within Ostracoda for at least 450 million years.

A Silurian short-great-appendage arthropod.

A new arthropod, Enalikter aphson gen. et sp. nov., is described from the Silurian (Wenlock Series) Herefordshire Lagerstätte of the UK. It belongs to the Megacheira (=short-great-appendage group), which is recognized here, for the first time, in strata younger than mid-Cambrian age. Discovery of this new Silurian taxon allows us to identify a Devonian megacheiran representative, Bundenbachiellus giganteus from the Hunsrück Slate of Germany. The phylogenetic position of megacheirans is controversial: they have been interpreted as stem chelicerates, or stem euarthropods, but when Enalikter and Bundenbachiellus are added to the most comprehensive morphological database available, a stem euarthropod position is supported. Enalikter represents the only fully three-dimensionally preserved stem-group euarthropod, it falls in the sister clade to the crown-group euarthropods, and it provides new insights surrounding the origin and early evolution of the euarthropods. Recognition of Enalikter and Bundenbachiellus as megacheirans indicates that this major arthropod group survived for nearly 100 Myr beyond the mid-Cambrian.

A Silurian myodocope with preserved soft-parts: cautioning the interpretation of the shell-based ostracod record.

Ostracod crustaceans are the most abundant fossil arthropods. The Silurian Pauline avibella gen. et sp. nov., from the Herefordshire Lagerstätte, UK, is an extremely rare Palaeozoic example with soft-part preservation. Based on its soft-part morphology, especially the exceptionally preserved limbs and presence of lateral eyes, it is assigned to the myodocopid myodocopes. The ostracod is very large, with an epipod on the fifth limb pair, as well as gills implying the presence of a heart and an integrated respiratory-circulatory system as in living cylindroleberidid myodocopids. Features of its shell morphology, however, recall halocyprid myodocopes and palaeocopes, encouraging caution in classifying ostracods based on the carapace alone and querying the interpretation of their shell-based fossil record, especially for the Palaeozoic, where some 500 genera are presently assigned to the Palaeocopida.

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.