Deciphering The Early Evolution of Echinoderms with Cambrian Fossils

Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister‐group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm‐type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan

A shallow-water cyrtocrinid crinoid (Articulata) from the upper Albian of the Western Pyrenees, North Spain

The cyrtocrinid crinoid Proholopus holopiformis (Remeš, 1902) is described from the upper Albian succession that may be included in the Albeniz unit to the east of Iruñea-Pamplona (Navarre, Western Pyrenees, Spain). Although based on partially disarticulated material, this taxon preserve calyx, stem, attachment structure, and brachial plates from the arms. Proholopus holopiformis was previously described from the Upper Jurassic-Lower Cretaceous of the Czech Republic and Lower Cretaceous of France and Crimea, thus the occurrence from Spain represents the youngest of the species. This allows a better characterization of the family Proholopodidae that was originally described based on calyx morphology only and expands its distribution to the upper Albian. Proholopus holopiformis inhabited the fore-reef areas of coral-sponge bioconstructions. Based on coral types, crinoids are thought to have dwelled near the euphotic-oligophotic zones transition and, thus, thrive in shallow depths. Most specimens have bite marks compatible with cidaroid predation. This represents one of the youngest occurrences of cyrtocrinids inhabiting shallow marine environments before their migration to the deep sea due to the ongoing Mesozoic Marine Revolution

Palaeoebiological implications of cuticle morphology, microstructure and formation in modern and fossil Daira (Decapoda, Brachyura, Dairoidea)

The origin and function of peculiar mushroom-shaped cuticular structures in some decapod crustaceans remains unknown. This ornamentation has appeared several times in widely disparate clades (in podotreme and heterotreme crabs, and pagurids). These structures are analysed in the modern genus Daira and compared with fossil material from the Eocene of Huesca and the Miocene of Alicante and Mallorca. A morphological and petrographic study is carried out using conventional microscopy and Scanning Electron Microscopy to understand the microstructure in modern and fossil representatives. This provides a clear view of the mushroom-like structures that cover the carapace of Daira and the distribution of the different layers of the exoskeleton. The results reveal a complex morphology, which involves all layers of the cuticle, with changes in the thickness of the exocuticle in different areas, and the presence of conical structures that especially affect the outer layers. These convolutions form a network of channels connected to the outside by pores. Finally, possible anti-predatory functions of these complex structures are proposed

Filling the early Eocene gap of paguroids (Decapoda, Anomura): a new highly diversified fauna from the Spanish Pyrenees (Serraduy Formation, Graus-Tremp Basin)

A highly diversified fauna of hermit crabs associated with reef environments from the Serraduy Formation (lower Eocene) in the southern Pyrenees (Huesca, Spain) is described. Other European Eocene outcrops have yielded paguroids associated with a single environment; however, the studied association represents one of the highest paguroid diversities in a single Eocene outcrop worldwide. The new material increases the diversity of known fossil paguroids, including eight species from which six are new: Clibanarius isabenaensis n. sp., Parapetrochirus serratus n. sp., Dardanus balaitus n. sp., ?Petrochirus sp., Eocalcinus veteris n. sp., ?Pagurus sp., Paguristes perlatus n. sp., and Anisopagurus primigenius n. sp. We erected a new combination for Paguristes sossanensis De Angeli and Caporiondo, 2009 and Paguristes cecconi De Angeli and Caporiondo, 2017 and transfer them to the genus Clibanarius. This association contains the oldest record of the genera Eocalcinus and Anisopagurus. Our data demonstrate that paguroids were diverse by the early Eocene in coral-reef environments and fill an important gap between the poorly known Paleocene assemblages and the more diverse mid- to late Eocene ones

A highly diverse dromioid crab assemblage (Decapoda, Brachyura) associated with pinnacle reefs in the lower Eocene of Spain

A highly diverse fauna of dromioid brachyurans from the Serraduy Formation (lower Eocene) in the western Pyrenees (Huesca, Spain) is described and illustrated. Recorded taxa are Mclaynotopus longispinosus new genus new species, Torodromia elongata n. gen. n. sp., Basidromilites glaessneri n. gen. n. sp., ?Basidromilites sp., Sierradromia gladiator n. gen. n. sp., Kromtitis isabenensis n. sp., and ?Basinotopus sp. Other European outcrops have yielded dromioids in association with specific environments, likely coral and sponge reef and siliciclastic soft bottoms; but the present material constitutes the most diverse dromioid assemblage from the lower Eocene worldwide. These dromioids co-occurred with a rich invertebrate fauna and lived near coral–algal reef mounds. Sedimentological data suggest that most of the fauna accumulated in fore reef settings as a result of storm activity. The present material greatly increases the diversity of known dromioid crabs associated with Eocene reef environments.UUID: http://zoobank.org/aed8cafa-7c64-4e70-bd45-9f357fc37a2

Cambrian cinctan echinoderms shed light on feeding in the ancestral deuterostome

Reconstructing the feeding mode of the latest common ancestor of deuterostomes is key to elucidating the early evolution of feeding in chordates and allied phyla; however, it is debated whether the ancestral deuterostome was a tentaculate feeder or a pharyngeal filter feeder. To address this, we evaluated the hydrodynamics of feeding in a group of fossil stem-group echinoderms (cinctans) using computational fluid dynamics. We simulated water flow past three-dimensional digital models of a Cambrian fossil cinctan in a range of possible life positions, adopting both passive tentacular feeding and active pharyngeal filter feeding. The results demonstrate that an orientation with the mouth facing downstream of the current was optimal for drag and lift reduction. Moreover, they show that there was almost no flow to the mouth and associated marginal groove under simulations of passive feeding, whereas considerable flow towards the animal was observed for active feeding, which would have enhanced the transport of suspended particles to the mouth. This strongly suggests that cinctans were active pharyngeal filter feeders, like modern enteropneust hemichordates and urochordates, indicating that the ancestral deuterostome employed a similar feeding strategy

Cambrian edrioasteroid reveals new mechanism for secondary reduction of the skeleton in echinoderms

Echinoderms are characterized by a distinctive high-magnesium calcite endoskeleton as adults, but elements of this have been drastically reduced in some groups. Herein, we describe a new pentaradial echinoderm, Yorkicystis haefneri n. gen. n. sp., which provides, to our knowledge, the oldest evidence of secondary non-mineralization of the echinoderm skeleton. This material was collected from the Cambrian Kinzers Formation in York (Pennsylvania, USA) and is dated as ca 510 Ma. Detailed morphological observations demonstrate that the ambulacra (i.e. axial region) are composed of flooring and cover plates, but the rest of the body (i.e. extraxial region) is preserved as a dark film and lacks any evidence of skeletal plating. Moreover, X-ray fluorescence analysis reveals that the axial region is elevated in iron. Based on our morphological and chemical data and on taphonomic comparisons with other fossils from the Kinzers Formation, we infer that the axial region was originally calcified, while the extraxial region was non-mineralized. Phylogenetic analyses recover Yorkicystis as an edrioasteroid, indicating that this partial absence of skeleton resulted from a secondary reduction. We hypothesize that skeletal reduction resulted from lack of expression of the skeletogenic gene regulatory network in the extraxial body wall during development. Secondary reduction of the skeleton in Yorkicystis might have allowed for greater flexibility of the body wall

The oldest dairoidid crab (Decapoda, Brachyura, Parthenopoidea) from the Eocene of Spain

Eubrachyurans, or ‘higher’ true crabs, are the most speciose group of decapod crustaceans and have a rich fossil record extending into the Early Cretaceous. However, most extant families are first found in the fossil record in the Palaeogene, and particularly in the Eocene. Unfortunately, fossils of many early eubrachyuran groups are often fragmentary, and only a few studies have combined extinct and extant taxa in a phylogenetic context using different optimality criteria. Here, we report the dairoidid crab Phrynolambrus sagittalis sp. nov., an enigmatic eubrachyuran from the upper Eocene of Huesca (northern Spain), whose completeness and exquisite preservation permit examination of its anatomy in a phylogenetic context. Dairoidids have previously been considered among the oldest stone crabs (Eriphioidea) or elbow crabs (Parthenopoidea), two disparate and distantly related groups of true crabs living today. Mechanical preparation and computed tomography of the fossil material revealed several diagnostic features that allow a detailed comparison with families across the crab tree of life, and test hypotheses about its phylogenetic affinities. Phrynolambrus sagittalis is the first record of the genus in the Iberian Peninsula, and represents one of the oldest crown parthenopoidean crabs worldwide, expanding our knowledge of the biogeographical distribution of elbow crabs during the Palaeogene, as well as their early origins, anatomical diversity and systematic affinities. Understanding the disparity of Eocene eubrachyurans is pivotal to disentangling the systematic relationships among crown families, and interpreting the spatio-temporal patterns leading to the evolution of modern faunas