Rudy Lerosey-Aubril

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.

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.

Occurrence of the Ordovician-type aglaspidid Tremaglaspis in the Cambrian Weeks Formation (Utah, USA)

The Guzhangian Weeks Formation preserves a diverse, yet virtually unstudied, non-trilobite arthropod fauna. Here we describe Tremaglaspis vanroyi sp. nov., the oldest representative of an enigmatic group of extinct arthropods, the Aglaspidida. Tremaglaspis was previously known from the Lower Ordovician and its morphology was regarded as particularly derived within the clade. Its occurrence in the Cambrian of Utah suggests that much of the early evolutionary history of the Aglaspidida remains unknown. A review of the environmental settings of previous aglaspidid findings suggests that these arthropods preferentially inhabited shallow-water environments, which may partially explain their limited fossil record.

The first aglaspidid sensu stricto from the Cambrian of China (Sandu Formation, Guangxi)

Aglaspidids represent an obscure group of lower Palaeozoic arthropods with a patchy biogeographic distribution. Before the recent description of a representative from Tasmania, these arthropods were exclusively known from Laurentia during the late Cambrian. Here we describe a new species, Aglaspella sanduensis sp. nov., from the Furongian of China, confirming that aglaspidids sensu stricto were already widely distributed worldwide by the late Cambrian; this demonstrates that some aglaspidids had great dispersal capabilities. A new diagnosis of the genus Aglaspella is proposed and the species formerly known as Aglaspella eatoni is assigned to a new taxon, Hesselbonia gen. nov.

A new aglaspidid euarthropod with a six-segmented trunk from the Lower Ordovician Fezouata Konservat-Lagerstätte, Morocco

A new euarthropod with an uncommon morphology, Brachyaglaspis singularis gen. et sp. nov., is described from the Early Ordovician (middle Floian) Fezouata biota of Morocco. The presence of a pair of postventral plates, widely attached to each other and located under the posterior-most trunk tergite and the base of the tailspine, indicates a phylogenetic relationship with the enigmatic group Aglaspidida. The overall morphology of Brachyaglaspis most closely resembles that of the ‘Ordovician-type’ aglaspidids, more specifically the late Cambrian – Early Ordovician genus Tremaglaspis . However, the presence of a prominent cephalon and only six trunk tergites in the new genus deviates from the organization of all other known aglaspidid species, notably extending the known range of morphological disparity of the group. A taxonomic revision of this euarthropod group indicates that the most accurate name and authorship combination correspond to Aglaspidida Walcott, 1912.

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 thecostracans. 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.

Notchia weugi gen. et sp. nov.: a new short-headed arthropod from the Weeks Formation Konservat-Lagerstätte (Cambrian; Utah)

The Weeks Formation preserves a diverse, yet largely undescribed, exceptionally preserved fauna of late Guzhangian age. Here I describe Notchia weugi gen. et sp. nov., a new arthropod characterized by a short cephalon, a trunk with 12 tergites and weakly differentiated into two morphological regions, and a spine-bearing rectangular telson. This combination of characters is incompatible with its assignment to any known groups. The new taxon also adds to examples of convergent evolution of ramified digestive glands in arthropods, possibly as an adaptation to infrequent feeding.

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.

Laurentian origin of solutan echinoderms: new evidence from the Guzhangian (Cambrian Series 3) Weeks Formation of Utah, USA

A new solutan echinoderm, Pahvanticystis utahensi s gen. et sp. nov. is described from the upper part of the Weeks Formation (Guzhangian). The Cambrian (Series 3) succession of the central House Range in western Utah documents the early diversification of the class Soluta, which is characterized by a major ecological transition from sessile, ‘pelmatozoan’ primitive taxa ( Coleicarpus , Wheeler Formation), to more and more vagile, temporarily attached ( Castericystis , Marjum Formation), to mostly unattached, ‘homalozoan’ derived forms ( Pahvanticystis , Weeks Formation). The morphology of Pahvanticystis is remarkably intermediate between those of Castericystis and Minervaecystis . Its twisted, flattened dististele possibly represents an adaptation for a more efficient crawling atop soft substrates. This morphological feature also questions the phylogenetic relationships between syringocrinid and dendrocystitid solutans, and the possible evolution of the latter from basal, Pahvanticystis - or Minervaecystis -like syringocrinids by paedomorphosis.

The Late Palaeozoic trilobites of Iran and Armenia and their palaeogeographical significance

The Iranian territory is composed of a mosaic of tectonic units, several of which underwent in the Permian and Triassic periods a migration from northern Gondwana to southern Laurussia associated with the opening of the Neo-Tethys Ocean. Although this broad outline of Permo-Triassic palaeogeographical evolution of Iranian microplates is now widely accepted, the individual timing of migration of these blocks, and their biogeographical relationships, remain insufficiently known. Here I review the Late Palaeozoic record of trilobites in Iran and Armenia, and discuss their palaeobiogeographical affinities in an attempt to shed light on the Permian palaeogeographical evolution of Iranian and Armenian terranes. Seven Iranian or Armenian localities, representative of five tectonic units, have yielded Carboniferous and Permian trilobites. Ten species are recognized, including two new taxa, Persia praecox gen. nov. sp. nov. and Pseudophillipsia ( s.l. ) parvizii sp. nov. P. praecox is the only Carboniferous (Tournaisian) species. The others are Wordian to Wuchiapingian in age and can be separated into three morphological groups, probably representing clades. One is composed of representatives of Acropyge , while the two others ( armenica -group and paffenholzi -group) comprise species of Pseudophillipsia . Only P. ( s.l. ) parvizii sp. nov. from the Zagros Mountains (Arabian Plate) is not attributed to one of these groups. The distribution of trilobites in Iran and Armenia strongly suggests that the Alborz, Central Iran and Transcaucasia microplates represented a single biogeographical unit in Middle and Late Permian times. Special relationships of this biochore with South China can also be stressed.