The Fossil Record of the Clam Shrimp (Crustacea; Branchiopoda).

Clam shrimp (the paraphyletic assemblage of spinicaudatans, laevicaudatans, cyclestherids and the extinct leaiins) are small, bivalved branchiopod crustaceans that specialize in ephemeral freshwater habitats. They have a long fossil record (Devonian onward) that has often been overlooked. Here we briefly review the fossil record of the major groups of clam shrimp and clear up some misconceptions in the literature as to their origin. The dominant group of clam shrimp in the fossil record is the Spinicaudata, which have a diverse fossil record beginning in the Devonian. The clam shrimp suborder Laevicaudata are known from the Permian, with possible soft-part preservation from the Jurassic. However, owing the character-poor nature of these fossils, it is impossible to tell if they represent crown group or stem group laevicaudatans. In contrast, the total group Spinicaudata have a rich record of mostly carapace fossils- the earliest from the Early Devonian. The leaiins are an enigmatic extinct diplostracan lineage thought to be closely related to the spinicaudatans. They have a record that extends from the Middle Devonian to the Permian. The Cyclestherida have a somewhat problematic fossil record: there are no examples of cyclestherids preserved with soft-parts, so the only character used to assign fossils to this lineage is the carapace shape. According to that metric, cyclestherids have a record that begins in the Middle Devonian. Exceptionally preserved clam shrimp are found in the Paleozoic and Mesozoic. Assessing holistically what is known about the clam shrimp fossil record along with carapace morphology, carapace ornamentation and examples of exceptional preservation will ultimately contribute to a synthetic paleontological and neontological understanding of the group, its systematics and evolution.

The World's First Clam Shrimp Symposium: Drawing Paleontology and Biology Together.

After a symposium and special issue devoted to the study of clam shrimp, it is tempting to ask what is next... where is the study of clam shrimp going? Rather than try to read the tea leaves to predict the future, we will instead offer some closing thoughts on where the study of clam shrimp should go and what areas are ripe for investigation. Many of these ideas integrate both fossil and modern clam shrimp to get at a more complete view of their evolution and ecology.

Fossil and Modern Clam Shrimp (Branchiopoda: Spinicaudata, Laevicaudata): The World's First Clam Shrimp Symposium and a Celebration of Brian V. Timms.

This special volume of Zoological Studies is the result of a symposium entitled "Fossil and Modern Clam Shrimp" held at the midyear meeting of The Crustacean Society in May of 2019. This symposium is the first ever focusing on clam shrimp, and the first conference where both palaeontologists and biologists specialising in these animals were able to come together. The papers presented here provide insight into the palaeontology, biology, ecology, taxonomy and phylogeny of the clam shrimp. This chapter introduces the symposium, its aims, and the resulting research, presented in the subsequent chapters. In addition, in this symposium we celebrate our great friend Brian V. Timms, who has mentored so many of us, brought us on various excursions across Australia, and has done more to advance Australian branchiopod studies than anyone else in history.

The phylogeny of fossil whip spiders.

BACKGROUND: Arachnids are a highly successful group of land-dwelling arthropods. They are major contributors to modern terrestrial ecosystems, and have a deep evolutionary history. Whip spiders (Arachnida, Amblypygi), are one of the smaller arachnid orders with ca. 190 living species. Here we restudy one of the oldest fossil representatives of the group, Graeophonus anglicus Pocock, 1911 from the Late Carboniferous (Duckmantian, ca. 315 Ma) British Middle Coal Measures of the West Midlands, UK. Using X-ray microtomography, our principal aim was to resolve details of the limbs and mouthparts which would allow us to test whether this fossil belongs in the extant, relict family Paracharontidae; represented today by a single, blind species Paracharon caecus Hansen, 1921. RESULTS: Tomography reveals several novel and significant character states for G. anglicus; most notably in the chelicerae, pedipalps and walking legs. These allowed it to be scored into a phylogenetic analysis together with the recently described Paracharonopsis cambayensis Engel & Grimaldi, 2014 from the Eocene (ca. 52 Ma) Cambay amber, and Kronocharon prendinii Engel & Grimaldi, 2014 from Cretaceous (ca. 99 Ma) Burmese amber. We recovered relationships of the form ((Graeophonus (Paracharonopsis + Paracharon)) + (Charinus (Stygophrynus (Kronocharon (Charon (Musicodamon + Paraphrynus)))))). This tree largely reflects Peter Weygoldt's 1996 classification with its basic split into Paleoamblypygi and Euamblypygi lineages; we were able to score several of his characters for the first time in fossils. Our analysis draws into question the monophyly of the family Charontidae. CONCLUSIONS: Our data suggest that Graeophonus is a crown group amblypygid, and falls within a monophyletic Paleoamblypgi clade, but outside the family Paracharontidae (= Paracharonopsis + Paracharon). Our results also suggest a new placement for the Burmese amber genus Kronocharon, a node further down from its original position. Overall, we offer a broad phylogenetic framework for both the fossil and Recent whip spiders against which future discoveries can be tested.

Testing the phylogenetic position of Cambrian pancrustacean larval fossils by coding ontogenetic stages.

The study of ontogeny as an integral part of understanding the pattern of evolution dates back over 200 years, but only recently have ontogenetic data been explicitly incorporated into phylogenetic analyses. Pancrustaceans undergo radical ontogenetic changes. The spectacular upper Cambrian "Orsten" fauna preserves phosphatized fossil larvae, including putative crown-group pancrustaceans with amazingly complete developmental sequences. The putative presence and nature of adult stages remains a source of debate, causing spurious placements in a traditional morphological analysis. We introduce a new coding method where each semaphoront (discrete larval or adult stage) is considered an operational taxonomic unit. This avoids a priori assumptions of heterochrony. Characters and their states are defined to identify changes in morphology throughout ontogeny. Phylogenetic analyses of semaphoronts produced possible relationships of each Orsten fossil to the crown-group clade expected from morphology shared with extant larvae. Bredocaris is a member of the stem lineage of Thecostraca or (Thecostraca + Copepoda), and Yicaris and Rehbachiella are probably members of the stem lineage of Cephalocarida. These placements rely directly on comparisons between extant and fossil larval character states. The position of Phosphatocopina remains unresolved. This method may have broader applications to other phylogenetic problems which may rely on ontogenetically variable homology statements.

Decoupling elongation and segmentation: notch involvement in anostracan crustacean segmentation.

Repeated body segments are a key feature of arthropods. The formation of body segments occurs via distinct developmental pathways within different arthropod clades. Although some species form their segments simultaneously without any accompanying measurable growth, most arthropods add segments sequentially from the posterior of the growing embryo or larva. The use of Notch signaling is increasingly emerging as a common feature of sequential segmentation throughout the Bilateria, as inferred from both the expression of proteins required for Notch signaling and the genetic or pharmacological disruption of Notch signaling. In this study, we demonstrate that blocking Notch signaling by blocking γ-secretase activity causes a specific, repeatable effect on segmentation in two different anostracan crustaceans, Artemia franciscana and Thamnocephalus platyurus. We observe that segmentation posterior to the third or fourth trunk segment is arrested. Despite this marked effect on segment addition, other aspects of segmentation are unaffected. In the segments that develop, segment size and boundaries between segments appear normal, engrailed stripes are normal in size and alignment, and overall growth is unaffected. By demonstrating Notch involvement in crustacean segmentation, our findings expand the evidence that Notch plays a crucial role in sequential segmentation in arthropods. At the same time, our observations contribute to an emerging picture that loss-of-function Notch phenotypes differ significantly between arthropods suggesting variability in the role of Notch in the regulation of sequential segmentation. This variability in the function of Notch in arthropod segmentation confounds inferences of homology with vertebrates and lophotrochozoans.

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.