A mid-Cambrian tunicate and the deep origin of the ascidiacean body plan.

Tunicates are an evolutionarily significant subphylum of marine chordates, with their phylogenetic position as the sister-group to Vertebrata making them key to unraveling our own deep time origin. Tunicates greatly vary with regards to morphology, ecology, and life cycle, but little is known about the early evolution of the group, e.g. whether their last common ancestor lived freely in the water column or attached to the seafloor. Additionally, tunicates have a poor fossil record, which includes only one taxon with preserved soft-tissues. Here we describe Megasiphon thylakos nov., a 500-million-year-old tunicate from the Marjum Formation of Utah, which features a barrel-shaped body with two long siphons and prominent longitudinal muscles. The ascidiacean-like body of this new species suggests two alternative hypotheses for early tunicate evolution. The most likely scenario posits M. thylakos belongs to stem-group Tunicata, suggesting that a biphasic life cycle, with a planktonic larva and a sessile epibenthic adult, is ancestral for this entire subphylum. Alternatively, a position within the crown-group indicates that the divergence between appendicularians and all other tunicates occurred 50 million years earlier than currently estimated based on molecular clocks. Ultimately, M. thylakos demonstrates that fundamental components of the modern tunicate body plan were already established shortly after the Cambrian Explosion.

New opabiniid diversifies the weirdest wonders of the euarthropod stem group.

Once considered 'weird wonders' of the Cambrian, the emblematic Burgess Shale animals Anomalocaris and Opabinia are now recognized as lower stem-group euarthropods and have provided crucial data for constraining the polarity of key morphological characters in the group. Anomalocaris and its relatives (radiodonts) had worldwide distribution and survived until at least the Devonian. However, despite intense study, Opabinia remains the only formally described opabiniid to date. Here we reinterpret a fossil from the Wheeler Formation of Utah as a new opabiniid, Utaurora comosa nov. gen. et sp. By visualizing the sample of phylogenetic topologies in treespace, our results fortify support for the position of U. comosa beyond the nodal support traditionally applied. Our phylogenetic evidence expands opabiniids to multiple Cambrian stages. Our results underscore the power of treespace visualization for resolving imperfectly preserved fossils and expanding the known diversity and spatio-temporal ranges within the euarthropod lower stem group.

Neuroanatomy in a middle Cambrian mollisoniid and the ancestral nervous system organization of chelicerates.

Recent years have witnessed a steady increase in reports of fossilized nervous tissues among Cambrian total-group euarthropods, which allow reconstructing the early evolutionary history of these animals. Here, we describe the central nervous system of the stem-group chelicerate Mollisonia symmetrica from the mid-Cambrian Burgess Shale. The fossilized neurological anatomy of M. symmetrica includes optic nerves connected to a pair of lateral eyes, a putative condensed cephalic synganglion, and a metameric ventral nerve cord. Each trunk tergite is associated with a condensed ganglion bearing lateral segmental nerves, and linked by longitudinal connectives. The nervous system is preserved as reflective carbonaceous films underneath the phosphatized digestive tract. Our results suggest that M. symmetrica illustrates the ancestral organization of stem-group Chelicerata before the evolution of the derived neuroanatomical characters observed in Cambrian megacheirans and extant representatives. Our findings reveal a conflict between the phylogenetic signals provided by neuroanatomical and appendicular data, which we interpret as evidence of mosaic evolution in the chelicerate stem-lineage.

New multipodomerous appendages of stem-group euarthropods from the Cambrian (Stage 4) Guanshan Konservat-Lagerstätte.

Stem-group euarthropods are important for understanding the early evolutionary and ecological history of the most species-rich animal phylum on Earth. Of particular interest are fossil taxa that occupy a phylogenetic position immediately crownwards of radiodonts, for this part of the euarthropod tree is associated with the appearance of several morphological features that characterize extant members of the group. Here, we report two new euarthropods from the Cambrian Stage 4 Guanshan Biota of South China. The fuxianhuiid Alacaris? sp. is represented by isolated appendages composed of a gnathobasic protopodite and an endite-bearing endopod of at least 20 podomeres. This material represents the youngest occurrence of the family Chengjiangocarididae, and its first record outside the Chengjiang and Xiaoshiba biotas. We also describe Lihuacaris ferox gen. et sp. nov. based on well-preserved and robust isolated appendages. Lihuacaris ferox exhibits an atypical combination of characters including an enlarged rectangular base, 11 endite-bearing podomeres and a hypertrophied distal element bearing 8-10 curved spines. Alacaris? sp. appendages display adaptations for macrophagy. Lihuacaris ferox appendages resemble the frontal appendages of radiodonts, as well as the post-oral endopods of chengjiangocaridid fuxianhuids and other deuteropods with well-documented raptorial/predatory habits. Lihuacaris ferox contributes towards the record of endemic biodiversity in the Guanshan Biota.

Cambrian comb jellies from Utah illuminate the early evolution of nervous and sensory systems in ctenophores.

Ctenophores are a group of predatory macroinvertebrates whose controversial phylogenetic position has prompted several competing hypotheses regarding the evolution of animal organ systems. Although ctenophores date back at least to the Cambrian, they have a poor fossil record due to their gelatinous bodies. Here, we describe two ctenophore species from the Cambrian of Utah, which illuminate the early evolution of nervous and sensory features in the phylum. Thalassostaphylos elegans has 16 comb rows, an oral skirt, and an apical organ with polar fields. Ctenorhabdotus campanelliformis has 24 comb rows, an oral skirt, an apical organ enclosed by a capsule and neurological tissues preserved as carbonaceous films. These are concentrated around the apical organ and ciliated furrows, which connect to a circumoral nerve ring via longitudinal axons. C. campanelliformis deviates from the neuroanatomy of living ctenophores and demonstrates a substantial complexity in the nervous system of Cambrian ctenophores.

Furongian (Jiangshanian) occurrences of radiodonts in Poland and South China and the fossil record of the Hurdiidae.

The Furongian period represents an important gap in the fossil record of most groups of non-biomineralizing organisms, owing to a scarcity of Konservat-Lagerstätten of that age. The most significant of these deposits, the Jiangshanian strata of the Sandu Formation near Guole Township (Guangxi, South China), have yielded a moderately abundant, but taxonomically diverse soft-bodied fossil assemblage, which provides rare insights into the evolution of marine life at that time. In this contribution, we report the first discovery of a radiodont fossil from the Guole Konservat-Lagerstätte. The specimen is an incomplete frontal appendage of a possibly new representative of the family Hurdiidae. It is tentatively interpreted as composed of seven podomeres, six of which bearing laminiform endites. The best preserved of these endites is especially long, and it bears short auxiliary spines that greatly vary in size. This is the second occurrence of hurdiids and more generally radiodonts in the Furongian, the first being the external mould of an oral cone from Jiangshanian strata of the Wisniówka Sandstone Formation in Poland. Restudy of this Polish specimen confirms that it belongs to a hurdiid radiodont and best compares to Peytoia. The family Hurdiidae includes the oldest (basal Cambrian Epoch 2) and youngest (Early Ordovician, possibly Early Devonian) representatives of the Radiodonta and as such, has the longest stratigraphical range of the group. Yet, hurdiids only became prominent components of marine ecosystems during the middle Cambrian (Miaolingian), and their fossil record in younger strata remains limited.

Protaspid larvae are unique to trilobites.

Trilobites represent a model for 'evo-devo' studies in fossil euarthropods, owing to a rare developmental trait: the biomineralization of the dorsal exoskeleton soon after hatching. Many fossilized trilobite ontogenies thus feature early stages - the protaspides - characterized by non-articulated, calcified dorsal exoskeletons. The recent discovery of a protaspid-like fossil occurring with aglaspidid remains in Middle Ordovician strata of Siberia has been interpreted as evidence for the presence of protaspides in these distant relatives of trilobites. In this contribution, we demonstrate that this Siberian protaspis actually belongs to the asaphid trilobite Isotelus (or a related taxon), a genus likely present in the same bed. We conclude that protaspid larvae still represent a developmental trait unique to Trilobita.

The diverse radiodont fauna from the Marjum Formation of Utah, USA (Cambrian: Drumian).

Radiodonts have long been known from Cambrian deposits preserving non-biomineralizing organisms. In Utah, the presence of these panarthropods in the Spence and Wheeler (House Range and Drum Mountains) biotas is now well-documented. Conversely, radiodont occurrences in the Marjum Formation have remained scarce. Despite the large amount of work undertaken on its diverse fauna, only one radiodont (Peytoia) has been reported from the Marjum Biota. In this contribution we quadruple the known radiodont diversity of the Marjum fauna, with the description of the youngest members of two genera, Caryosyntrips and Pahvantia, and that of a new taxon Buccaspinea cooperi gen. et sp. nov. This new taxon can be identified from its large oral cone bearing robust hooked teeth with one, two, or three cusps, and by the unique endite morphology and organisation of its frontal appendages. Appendages of at least 12 podomeres bear six recurved plate-like endites proximal to up to four spiniform distal endites. Pahvantia hastata specimens from the Marjum Formation are particularly large, but otherwise morphologically indistinguishable from the carapace elements of this species found in the Wheeler Formation. One of the two new Caryosyntrips specimens can be confidently assigned to C. camurus. The other bears the largest spines relative to appendage length recorded for this genus, and possesses endites of variable size and unequal spacing, making its taxonomic assignment uncertain. Caryosyntrips, Pahvantia, and Peytoia are all known from the underlying Wheeler Formation, whereas isolated appendages from the Spence Shale and the Wheeler Formation, previously assigned to Hurdia, are tentatively reidentified as Buccaspinea. Notably, none of these four genera occurs in the overlying Weeks Formation, providing supporting evidence of a faunal restructuring around the Drumian-Guzhangian boundary. The description of three additional nektonic taxa from the Marjum Formation further documents the higher relative proportion of free-swimming species in this biota compared to those of the Wheeler and Weeks Lagerstätten. This could be related to a moderate deepening of the basin and/or changing regional ocean circulation at this time.

Revision of the mollisoniid chelicerate(?) Thelxiope, with a new species from the middle Cambrian Wheeler Formation of Utah.

The recent re-interpretation of the Lower Palaeozoic euarthropod group Mollisonia as belonging to Chelicerata has triggered a renewed interest for the poorly known family Mollisoniidae. In this contribution, we revise the anatomy, taxonomic diversity, and systematics of Thelxiope, the sister-taxon of Mollisonia. This mollisoniid genus comprises four species, and is characterized by the presence of one cephalic, seven thoracic (one per tergite), and three pygidial long sagittal spines. The type species, T. palaeothalassia Simonetta & Delle Cave, is a rare taxon in the Wuliuan Burgess Shale Formation of Canada, which can be recognized by the hypertrophy of a single of its sagittal spines, the posteriomost one. T. spinosa (Conway Morris & Robison)-a species originally assigned to a distinct genus 'Ecnomocaris' herein synonymised with Thelxiope-is known from a single specimen found in the Drumian Wheeler Formation of the House Range of Utah. It differs from the type-species in the hypertrophy of both the anteriormost (cephalic) and the posteriormost (third pygidial) sagittal spines. The same Wheeler strata have also yielded a single specimen of a new taxon, T. holmani sp. nov., which lacks hypertrophied sagittal spines and features blunt thoracic tergopleural tips. A putative fourth species, referred to Thelxiope sp. nov. A, extends the stratigraphical range of Thelxiope to the Lower Ordovician (Tremadocian), and its palaeographic range to West Gondwana. Currently under study, this relatively common component of the lower Fezouata Shale fauna is only briefly discussed. Features characterizing the genus Thelxiope and its components almost exclusively pertain to the sagittal spines, for the scarcity and inconsistent preservation of the Cambrian materials as-yet available preclude a confident assessment of the variability of other morphological features. The pygidium in Thelxiope and Mollisonia is not composed of four, but three tergites essentially similar to thoracic ones, except for the lack of articulations.

Proclivity of nervous system preservation in Cambrian Burgess Shale-type deposits.

Recent investigations on neurological tissues preserved in Cambrian fossils have clarified the phylogenetic affinities and head segmentation in pivotal members of stem-group Euarthropoda. However, palaeoneuroanatomical features are often incomplete or described from single exceptional specimens, raising concerns about the morphological interpretation of fossilized neurological structures and their significance for early euarthropod evolution. Here, we describe the central nervous system (CNS) of the short great-appendage euarthropod Alalcomenaeus based on material from two Cambrian Burgess Shale-type deposits of the American Great Basin, the Pioche Formation (Stage 4) and the Marjum Formation (Drumian). The specimens reveal complementary ventral and lateral views of the CNS, preserved as a dark carbonaceous compression throughout the body. The head features a dorsal brain connected to four stalked ventral eyes, and four pairs of segmental nerves. The first to seventh trunk tergites overlie a ventral nerve cord with seven ganglia, each associated with paired sets of segmental nerve bundles. Posteriorly, the nerve cord features elongate thread-like connectives. The Great Basin fossils strengthen the original description-and broader evolutionary implications-of the CNS in Alalcomenaeus from the early Cambrian (Stage 3) Chengjiang deposit of South China. The spatio-temporal recurrence of fossilized neural tissues in Cambrian Konservat-Lagerstätten across North America (Pioche, Burgess Shale, Marjum) and South China (Chengjiang, Xiaoshiba) indicates that their preservation is consistent with the mechanism of Burgess Shale-type fossilization, without the need to invoke alternative taphonomic pathways or the presence of microbial biofilms.

Appendicular anatomy of the artiopod Emeraldella brutoni from the middle Cambrian (Drumian) of western Utah.

The non-biomineralized artiopod Emeraldella brutoni Stein, Church & Robison, from the middle Cambrian (Drumian) Wheeler Formation in Utah represents the only confirmed occurrence of the genus Emeraldella outside of the stratigraphically older (Wuliuan) Burgess Shale Konservat-Lagerstätte in British Columbia. The hitherto known sole specimen of this species is preserved in dorsal view and lacks critical information on the ventral appendages. Here, we redescribe E. brutoni based on a new completely articulated specimen that illustrates the appendage organization in exceptional detail. The main body consists of a cephalic region covered by a semicircular head shield, a trunk including 10 tergites with expanded pleurae plus a cylindrical terminal segment, and a long articulated tailspine. The head carries a pair of elongate and flexible antennae, a pair of lateral eyes, and three pairs of post-antennal appendages. We report the presence of eyes in Emeraldella for the first time. The first post-antennal limb solely consists of an endopod with well-developed paired spiniform endites. The remaining cephalic appendages and those associated with all but the last trunk segments possess exopods terminating in paddle-shaped, distal lobes fringed with robust setae. The cylindrical terminal segment bears a pair of posteriorly oriented caudal flaps reminiscent of trunk exopods, and a styliform, possibly uniarticulate tailspine longer than the main body. The new data on E. brutoni suggests an exopodal origin for the paired caudal structures in Vicissicaudata, and improve our understanding of the fundamental organization of this major clade within Artiopoda.

New suspension-feeding radiodont suggests evolution of microplanktivory in Cambrian macronekton.

The rapid diversification of metazoans and their organisation in modern-style marine ecosystems during the Cambrian profoundly transformed the biosphere. What initially sparked this Cambrian explosion remains passionately debated, but the establishment of a coupling between pelagic and benthic realms, a key characteristic of modern-day oceans, might represent a primary ecological cause. By allowing the transfer of biomass and energy from the euphotic zone-the locus of primary production-to the sea floor, this biological pump would have boosted diversification within the emerging metazoan-dominated benthic communities. However, little is known about Cambrian pelagic organisms and their trophic interactions. Here we describe a filter-feeding Cambrian radiodont exhibiting morphological characters that likely enabled the capture of microplankton-sized particles, including large phytoplankton. This description of a large free-swimming suspension-feeder potentially engaged in primary consumption suggests a more direct involvement of nekton in the establishment of an oceanic pelagic-benthic coupling in the Cambrian.

The Vicissicaudata revisited - insights from a new aglaspidid arthropod with caudal appendages from the Furongian of China.

Cambrian marine ecosystems were dominated by arthropods, and more specifically artiopods. Aglaspidids represent an atypical group amongst them, not the least because they evolved and rapidly diversified during the late Cambrian, a time interval between the two diversification events of the Early Palaeozoic. Recent phylogenetic analyses have retrieved aglaspidids within the Vicissicaudata, a potentially important, but difficult to define clade of artiopods. Here we describe a new aglaspidid from the Furongian Guole Konservat-Lagerstätte of South China. This taxon displays a pretelsonic segment bearing non-walking appendages, features as-yet known in all vicissicaudatans, but aglaspidids. A new comprehensive phylogenetic analysis provides strong support for the legitimacy of a monophyletic clade Vicissicaudata, and demonstrates the pertinence of new characters to define Aglaspidida. It also motivates important changes to the systematics of the phylum, including the elevation of Artiopoda to the rank of subphylum, and the establishment of a new superclass Vicissicaudata and a new aglaspidid family Tremaglaspididae. Two diversification pulses can be recognized in the early history of artiopods - one in the early Cambrian (trilobitomorphs) and the other in the late Cambrian (vicissicaudatans). The discrepancy between this pattern and that traditionally depicted for marine invertebrates in the Early Palaeozoic is discussed.

Reconstructing the diet of a 505-million-year-old arthropod: Sidneyia inexpectans from the Burgess Shale fauna.

The feeding ecology of the 505-million-year-old arthropod Sidneyia inexpectans from the middle Cambrian (Series 3, Stage 5) Burgess Shale fauna (British Columbia, Canada) is revealed by three lines of evidence: the structure of its digestive system, the fossilized contents of its gut and the functional anatomy of its appendages. The digestive tract of Sidneyia is straight, tubular and relatively narrow in the trunk region. It is enlarged into a pear-shaped area in the cephalic region and stretches notably to form a large pocket in the abdomen. The mouth is ventral, posteriorly directed and leads to the midgut via a short tubular structure interpreted as the oesophagus. Anteriorly, three pairs of glands with internal, branching tubular structures open into the digestive tract. These glands have equivalents in various Cambrian arthropod taxa (e.g. naraoiids) and modern arthropods. Their primary function was most likely to digest and assimilate food. The abdominal pocket of Sidneyia concentrates undigested skeletal elements and various residues. It is interpreted here as the functional analogue of the stercoral pocket of some extant terrestrial arachnids (e.g. Araneae, Solifugae), whose primary function is to store food residuals and excretory material until defecation. Analysis of the gut contents indicates that Sidneyia fed largely on small ptychopariid trilobites, brachiopods, possibly agnostids, worms and other undetermined animals. Sidneyia was primarily a durophagous carnivore with predatory and/or scavenging habits, feeding on small invertebrates that lived at the water-sediment interface. There is no evidence for selective feeding. Its food items (e.g. living prey or dead material) were grasped and manipulated ventrally by its anterior appendages, then macerated into ingestible fragments and conveyed to the mouth via the converging action of strong molar-like gnathobases. Digestion probably took place within the anterior midgut via enzymes secreted in the glands. Residues were transported through the digestive tract into the abdominal pocket. The storage of faeces suggests infrequent feeding. The early diagenetic three-dimensional preservation of the digestive glands and abdominal pocket may be due to the capacity of Sidneyia to store Phosphorus and Calcium (e.g. spherites) in its digestive tissues during life as do, for example, modern horseshoe crabs.

Sophisticated digestive systems in early arthropods.

Understanding the way in which animals diversified and radiated during their early evolutionary history remains one of the most captivating of scientific challenges. Integral to this is the 'Cambrian explosion', which records the rapid emergence of most animal phyla, and for which the triggering and accelerating factors, whether environmental or biological, are still unclear. Here we describe exceptionally well-preserved complex digestive organs in early arthropods from the early Cambrian of China and Greenland with functional similarities to certain modern crustaceans and trace these structures through the early evolutionary lineage of fossil arthropods. These digestive structures are assumed to have allowed for more efficient digestion and metabolism, promoting carnivory and macrophagy in early arthropods via predation or scavenging. This key innovation may have been of critical importance in the radiation and ecological success of Arthropoda, which has been the most diverse and abundant invertebrate phylum since the Cambrian.

The sensory dorsal organs of crustaceans.

The cuticle of crustaceans bears numerous organs, of which the functions of many are unknown. One of these, the sensory dorsal organ (SDO), is present in a wide diversity of taxa. Here we critically review the variability, ultrastructure, distribution, and possible function of this enigmatic cuticular organ. Previous data are complemented by new observations on larvae and adults of various malacostracans. The SDO is composed of four sensors arranged as the corners of a square, the centre of which is occupied by a gland. Pores or pegs surrounding this central complex may also form part of the organ. The arrangement and the external aspect of the five main elements varies greatly, but this apparently has little impact on their ultrastructural organisation. The sensors and the gland are associated with a particularly thin cuticle. Each sensor contains four outer dendritic segments and the central gland is made of a single large cell. It is not yet known what this large cell secretes. The SDO is innervated from the tritocerebrum and therefore belongs to the third cephalic segment. A similar organ, here called the posterior SDO, has been repeatedly observed more posteriorly on the carapace. It resembles the SDO but has a greater number of sensors (usually six, but up to ten) apparently associated with only two outer dendritic segments. The SDO and the posterior SDO are known in the Eumalacostraca, the Hoplocarida, and the Phyllocarida. Some branchiopods also possess a 'dorsal organ' resembling both the SDO and the ion-transporting organ more typical of this group. This may indicate a common origin for these two functionally distinct groups of organs. New observations on the posterior SDO support the hypothesis that the SDO and the posterior SDO are homologous to the lattice organ complexes of the costracans. However, the relationship between the SDO and the dorsal cephalic hump of calanoid copepods remains unclear. No correlation can be demonstrated between the presence of a SDO and a particular ecological or biological trait. In fossils, the most convincing examples of SDO-like organs are found in some Late Cambrian arthropods from the Alum Shale of southern Sweden. They suggest that related organs might have been present in non-crustacean Cambrian arthropods. The distribution of the SDO and posterior SDO in extant and fossil crustaceans strongly suggests that these organs originated early in the history of the group, and are crucial to the functioning of these organisms. However, except for knowing that the sensors are chemoreceptors and that in a given organ a functional relationship probably exists between them and the gland, little is known about this function. The description of a SDO in freshwater carideans, which can be easily reared in a laboratory, opens the way for behavioural and physiological experiments to be undertaken that could prove crucial for the determination of this function.

Controls on gut phosphatisation: the trilobites from the Weeks Formation Lagerstätte (Cambrian; Utah).

Despite being internal organs, digestive structures are frequently preserved in Cambrian Lagerstätten. However, the reasons for their fossilisation and their biological implications remain to be thoroughly explored. This is particularly true with arthropods--typically the most diverse fossilised organisms in Cambrian ecosystems--where digestive structures represent an as-yet underexploited alternative to appendage morphology for inferences on their biology. Here we describe the phosphatised digestive structures of three trilobite species from the Cambrian Weeks Formation Lagerstätte (Utah). Their exquisite, three-dimensional preservation reveals unique details on trilobite internal anatomy, such as the position of the mouth and the absence of a differentiated crop. In addition, the presence of paired pygidial organs of an unknown function is reported for the first time. This exceptional material enables exploration of the relationships between gut phosphatisation and the biology of organisms. Indeed, soft-tissue preservation is unusual in these fossils as it is restricted to the digestive structures, which indicates that the gut played a central role in its own phosphatisation. We hypothesize that the gut provided a microenvironment where special conditions could develop and harboured a source of phosphorus. The fact that gut phosphatization has almost exclusively been observed in arthropods could be explained by their uncommon ability to store ions (including phosphorous) in their digestive tissues. However, in some specimens from the Weeks Formation, the phosphatisation extends to the entire digestive system, suggesting that trilobites might have had some biological particularities not observed in modern arthropods. We speculate that one of them might have been an increased capacity for ion storage in the gut tissues, related to the moulting of their heavily-mineralised carapace.