The Drumian Wheeler Konservat‐Lagerstätte of the House Range of Utah (Wheeler‐HR) has yielded one of the most diverse exceptionally preserved Cambrian biotas of North America. The discovery of soft‐bodied fossils invariably provides precious insights into this biota, given that most of its non‐biomineralizing components are known from very few specimens. This contribution describes some 30 new exceptionally preserved fossils of Wheeler panarthropods. Two new species are recognized, the radiodont Hurdia sp. nov. A and the megacheiran Kanoshoia rectifrons gen. et sp. nov. Along with a species of Leanchoilia, K. rectifrons represents the first confident megacheiran record in these strata. The presence of the radiodont genus Amplectobelua and the isoxyid species Isoxys longissimus is reported outside of the Burgess Shale in Laurentia. New specimens of Caryosyntrips serratus, Naraoia compacta, Messorocaris magna and Mollisonia symmetrica provide insights into the phylogenetic affinities, local spatial distribution and morphological variation of these species hitherto known by single specimens in the Wheeler‐HR. The same is true of new materials of the more common Pahvantia hastata and Perspicaris? dilatus. Formal descriptions of the order Mollisoniida ord. nov. and family Mollisoniidae fam. nov. are also provided. Last, the preservation of body structures other than the dorsal exoskeletons is illustrated for the first time in two common components of the fauna: the agnostid Itagnostus interstrictus and the bivalved euarthropod Pseudoarctolepis sharpi. The new material substantially improves our understanding of the diversity of the Wheeler‐HR biota, and provides new evidence of its distinctiveness relative to the Wheeler biota of the Drum Mountains.
Radiodonts are a diverse clade of Lower Palaeozoic stem-group euarthropods that played a key role in the emergence of complex marine trophic webs. The latest addition to the group, Cambroraster falcatus, was recently described from the Wuliuan Burgess Shale, and is characterized by a unique horseshoe-shaped central carapace element. Here we report the discovery of Cambroraster sp. nov. A, a new species from the Cambrian Stage 3 Chengjiang Lagerstätte of South China. The new occurrence of Cambroraster demonstrates that some of the earliest known radiodonts had already evolved a highly derived carapace morphology adapted to an essentially eudemersal life as sediment foragers.
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.
Trace fossils represent the primary source of information on the evolution of animal behaviour through deep time, and provide exceptional insights into complex life strategies that would be otherwise impossible to infer from the study of body parts alone. Here, we describe unusual trace fossils found in marginal-marine, storm- and river-flood deposits from the Middle Devonian Naranco Formation of Asturias (northern Spain) that constitute the first evidence for infaunal moulting in a non-trilobite euarthropod. The trace fossils are preserved in convex hyporelief, and include two main morphological variants that reflect a behavioural continuum. Morphotype 1 consists of a structure that superficially resembles a Rusophycus with an oval outline that possesses a distinctly three lobed axis with an elevated central ridge and regularly spaced transverse furrows that convey the appearance of discrete body segments. The anterior part is the most irregular region of the structure, and it is not always recorded. Morphotype 2 displays more elongated, tubular morphology. Careful observation, however, reveals that it comprises up to three successive morphotype 1 specimens organised in a linear fashion and partially truncating each other. Trilobate morphology and effaced transverse furrows are locally evident, but the predominant morphological feature is the continuous, elevated ridge. The detailed morphology of morphotype 1 and well-preserved, discrete segments of morphotype 2 closely resemble the dorsal exoskeleton of the enigmatic late Carboniferous euarthropod Camptophyllia, suggesting the possible affinities of the producer. Comparisons with patterns of Devonian phacopid trilobite exuviation suggest that the Naranco Formation trace fossils may have been produced by the infaunal activities of an euarthropod that anchored its dorsal exoskeleton in the firm sediment during the body inversion moult procedure. Our findings expand the phylogenetic and environmental occurrence of infaunal moulting in Palaeozoic euarthropods, and suggest a defensive strategy against predation, previously only known from trilobites preserved in open-marine deposits.
The non-biomineralized artiopod Emeraldella brutoni Stein, Church & Robinson, 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.
Artiopodan euarthropods represent common and abundant faunal components in sites with exceptional preservation during the Cambrian. The Chengjiang biota in South China contains numerous taxa that are exclusively known from this deposit, and thus offer a unique perspective on euarthropod diversity during the early Cambrian. One such endemic taxon is the non-trilobite artiopodan Sinoburius lunaris, which has been known for approximately three decades, but few details of its anatomy are well understood due to its rarity within the Chengjiang, as well as technical limitations for the study of these fossils. Furthermore, the available material does not provide clear information on the ventral organization of this animal, obscuring our understanding of phylogenetically significant details such as the appendages.
We employed X-ray computed tomography to study the non-biomineralized morphology of Sinoburius lunaris. Due to the replacement of the delicate anatomy with pyrite typical of Chengjiang fossils, computed tomography reveals substantial details of the ventral anatomy of Sinoburius lunaris, and allow us to observe in detail the three-dimensionally preserved appendicular organization of this taxon for the first time. The dorsal exoskeleton consists of a crescent-shaped head shield with well-developed genal spines, a thorax with seven freely articulating tergites, and a fused pygidium with lateral and median spines. The head bears a pair of ventral stalked eyes that are accommodated by dorsal exoskeletal bulges, and an oval elongate ventral hypostome. The appendicular organization of the head is unique among Artiopoda. The deutocerebral antennae are reduced, consisting of only five podomeres, and bear an antennal scale on the second podomere that most likely represents an exite rather than a true ramus. The head includes four post-antennal biramous limb pairs. The first two biramous appendages are differentiated from the rest. The first appendage pair consists of a greatly reduced endopod coupled with a greatly elongated exopod with a potentially sensorial function. The second appendage pair carries a more conventionally sized endopod, but also has an enlarged exopod. The remaining biramous appendages are homonomous in their construction, but decrease in size towards the posterior end of the body. They consist of a basipodite with ridge-like crescentic endites, an endopod with seven podomeres and a terminal claw, and a lamellae-bearing exopod with a slender shaft. Contrary to previous reports, we confirm the presence of segmental mismatch in Sinoburius lunaris, expressed as diplotergites in the thorax. Maximum parsimony and Bayesian phylogenetic analyses support the monophyly of Xandarellida within Artiopoda, and illuminate the internal relationships within this enigmatic clade. Our results allow us to propose a transformation series explaining the origin of archetypical xandarellid characters, such as the evolution of eye slits in Xandarella spectaculum and Phytophilaspis pergamena as derivates from the anterolateral notches in the head shield observed in Cindarella eucalla and Luohuilinella species. In this context, Sinoburius lunaris is found to feature several derived characters within the group, such as the secondary loss of eye slits and a high degree of appendicular tagmosis. Contrary to previous findings, our analyses strongly support close affinities between Sinoburius lunaris, Xandarella spectaculum and Phytophilaspis pergamena, although the precise relationships between these taxa are sensitive to different methodologies.
The revised morphology of Sinoburius lunaris, made possible through the use of computed tomography to resolve details of its three-dimensionally preserved appendicular anatomy, contributes towards an improved understanding of the morphology of this taxon and the evolution of Xandarellida more broadly. Our results indicate that Sinoburius lunaris possesses an unprecedented degree of appendicular tagmosis otherwise unknown within Artiopoda, with the implication that this iconic group of Palaeozoic euarthropods likely had a more complex ecology and functional morphology than previously considered. The application of computer tomographic techniques to the study of Chengjiang euarthropods holds exceptional promise for understanding the morphological diversity of these organisms, and also better reconstructing their phylogenetic relationships and evolutionary history.
Moulting is a fundamental component of the ecdysozoan life cycle, but the fossil record of this strategy is susceptible to preservation biases, making evidence of ecdysis in soft-bodied organisms extremely rare. Here, we report an exceptional specimen of the fuxianhuiid Alacaris mirabilis preserved in the act of moulting from the Cambrian (Stage 3) Xiaoshiba Lagerstätte, South China. The specimen displays a flattened and wrinkled head shield, inverted overlap of the trunk tergites over the head shield, and duplication of exoskeletal elements including the posterior body margins and telson. We interpret this fossil as a discarded exoskeleton overlying the carcass of an emerging individual. The moulting behaviour of A. mirabilis evokes that of decapods, in which the carapace is separated posteriorly and rotated forward from the body, forming a wide gape for the emerging individual. A. mirabilis illuminates the moult strategy of stem-group Euarthropoda, offers the stratigraphically and phylogenetically earliest direct evidence of ecdysis within total-group Euarthropoda, and represents one of the oldest examples of this growth strategy in the evolution of Ecdysozoa.
Palaeontological data plays a key role in reconstructing the biology, ecology, and evolution of the biosphere throughout the history of Life on Earth. Whereas modern molecular-based technologies have led to significant progress in understanding the developmental biology and phylogenetic relationships of extant animal phyla, Cambrian soft-bodied fossils represent the only way of studying the morphology of their extinct ancestors. This chapter provides an overview of the significance of exceptionally preserved Cambrian fossils and their contribution towards illuminating the origin of animal body plans. Over the last few decades, conceptual and technical advances have distanced Cambrian fossils away from their previous misconception as failed evolutionary experiments. Current effortsfocus instead on clarifying the evolutionary links between these fascinating organisms and their extant descendants, made possible in part by the discovery of new exceptional deposits, and the improved knowledge of their unusual geological history capable of preserving delicate but informative tissues such as guts and brains.
The Ecdysozoa is a major animalclade whose main uniting feature is a distinctive growth strategy that requires the periodical moulting of the external cuticle. The staggering diversity within Ecdysozoa has prompted substantial efforts to reconstruct their origin and early evolution. Based on palaentological and developmental data, we proposed a scenario for the early evolution of the ecdysozoan clade Panarthropoda (Onychophora, Tardigrada, Euarthropoda), and postulated that a terminal mouth is ancestral for this lineage. In light of the accompanying comment by Claus Nielsen, we take this opportunity to clarify the significance of our argumentation for Panarthropoda in the phylogenetic context of Ecdysozoa, and Bilateria more broadly. We conclude that the ancestral ecdysozoan most likely had an adult terminal mouth, and that the last common ancestors of all the phyla that constitute Ecdysozoa almost certainly also had an adult terminal mouth. The occurrence of a ventral-facing mouth in various adult ecdysozoans – particularly panarthropods – is the result of convergence. Despite the paucity of embryological data on fossiltaxa, we contemplate the likelihood that a developmentally early ventral mouth opening could be ancestral for Ecdysozoa, and if so, then this would represent a symplesiomorphy of Bilateria as a whole.
Vision allows animals to interact with their environment. Aquatic chelicerates dominate the early record of lateral compound eyes among non-biomineralizing crown-group euarthropods. Although the conservative morphology of lateral eyes in Xiphosura is potentially plesiomorphic for Euarthropoda, synziphosurine eye organization has received little attention despite their early diverging phylogenetic position. Here, we re-evaluate the fossil evidence for lateral compound eyes in the synziphosurines Bunodes sp., Cyamocephalus loganensis, Legrandella lombardii, Limuloides limuloides, Pseudoniscus clarkei, Pseudoniscus falcatus and Pseudoniscus roosevelti. We compare these data with lateral eyes in the euchelicerates Houia yueya, Kasibelinurus amicorum and Lunataspis aurora. We find no convincing evidence for lateral eyes in most studied taxa, and Pseudoniscus roosevelti and Legrandella lombardii are the only synziphosurines with this feature. Our findings support two scenarios for euchelicerate lateral eye evolution. The elongate-crescentic lateral eyes of Legrandella lombardii might represent the ancestral organization, as suggested by the phylogenetic position of this taxon in stem-group Euchelicerata. Alternatively, the widespread occurrence of kidney-shaped lateral eyes in stem-group Xiphosura and stem-group Arachnida could represent the plesiomorphic condition; Legrandella lombardii eyes would therefore be derived. Both evolutionary scenarios support the interpretation that kidney-shaped lateral eyes are ancestral for crown-group Euchelicerata and morphologically conserved in extant Limulus polyphemus.
The Aglaspidida is a major group of Palaeozoic arthropods traditionally regarded as problematic, but whose evolutionary history has attracted renewed attention over the last decade. These efforts have resulted in an improved understanding of various aspects of aglaspidid palaeobiology, including their environmental, geographical and stratigraphical distribution, and phylogenetic relationships. Here, we describe a new weakly biomineralized non-trilobite arthropod from the Furongian Guole Konservat–Lagerstätte (Sandu Formation) in South China that contributes towards a more complete reconstruction of aglaspidid evolution and diversity during the late Cambrian. Beckwithia? gracilis n. sp. resembles the type species B. typa in the possession of a reentrant posterior cephalic margin, general trunk appearance and coarse tuberculation, but differs in having a more slender construction and a homogeneous exoskeletal sculpture. A revision of putative Beckwithia from Australia and Russia argues against their inclusion in this taxon, making the new species the only representative of the genus outside Laurentia. After Aglaspella and Glypharthrus, also from the Guole biota, this is the third aglaspidid genus shared between Laurentia and South China, which supports the hypothesis that some aglaspidids had great dispersal capabilities.
We describe the exceptionally well‐preserved non‐trilobite artiopodan Zhiwenia coronata gen. et sp. nov. from the Cambrian Stage 3 Xiaoshiba Lagerstätte in Yunnan, China. The exoskeleton consists of a cephalic shield with dorsal sutures expressed as lateral notches that accommodate stalked lateral eyes, an elongate trunk composed of 20 tergites—the first of which is reduced—and a short tailspine with marginal spines. Appendicular data include a pair of multi‐segmented antennae, and homonomous biramous trunk limbs consisting of an endopod with at least seven podomeres and a flattened exopod with lamellae. Although the presence of cephalic notches and a reduced first trunk tergite invites comparisons with the petalopleurans Xandarella, Luohiniella and Cindarella, the proportions and exoskeletal tagmosis of Zhiwenia do not closely resemble those of any major group within Trilobitomorpha. Parsimony and Bayesian phylogenetic analyses consistently support Zhiwenia as sister‐taxon to the Emu Bay Shale artiopodan Australimicola spriggi, and both of them as closely related to Acanthomeridion from the Chengjiang. This new monophyletic clade, Protosutura nov., occupies a basal phylogenetic position within Artiopoda as sister‐group to Trilobitomorpha and Vicissicaudata, illuminates the ancestral organization of these successful euarthropods, and leads to a re‐evaluation of the evolution of ecdysial dorsal sutures within the group.
The euarthropod body consists of serially repeated segments with various degrees of functional specialization and regionalization  . Some representatives exhibit deviant metameric patterns expressed as an indirect correspondence between components of the exoskeleton, usually the number or position of dorsoventral sclerotized plates and walking legs ( Supplemental Information ) [1–3] . Segmental mismatch in the form of supernumerary walking legs per tergite (i.e. dorsal exoskeletal plate) is characteristic of fuxianhuiids, Cambrian euarthropods widely regarded as critical for understanding the origin of this phylum [4,5] . The broader significance of this organization remains obscure, however, due to the difficulty of distinguishing which components of the fuxianhuiid trunk reflect ancestral or derived traits. Here, we describe for the first time the presence of metameric midgut diverticulae in Fuxianhuia protensa from the Chengjiang Konservat-Lagerstätte and demonstrate that these digestive structures follow the segmentation pattern of the dorsal exoskeleton. Midgut diverticulae signal a predatory or scavenging ecology [6,7] , falsifying the view of fuxianhuiids as simple mud-feeders  . Comparison with other euarthropods [1–3,5] indicates that fuxianhuiids possessed a unique mode of exoskeletal and visceral segmental mismatch, in which the tergites and midgut were segmentally patterned independently from the walking legs and ventral nerve cord. Our findings provide direct evidence of substantial developmental flexibility among stem-group euarthropods during the Cambrian.
La Tierra ha sido habitada por organismos vivientes por al menos 3 mil 500 millones de años (Ma)1. La vida ha evolucionado de forma dramática durante este tiempo, diversificándose y adaptándose a ambientes altamente dinámicos, y simultáneamente transformando la estructura física y química de dichos hábitats. En este contexto, el registro fósil adquiere una importancia incomparable, dado que captura evidencia directa sobre la evolución y diversificación de la vida en la Tierra en una escala de tiempo profundo. El origen de los animales representa una de las mayores contribuciones del registro fósil, no solamente porque este grupo biológico incluye a los seres humanos como especie, sino por el tremendo efecto que estos complejos organismos han tenido en modelar las interacciones que definen el funcionamiento global de la biosfera durante los últimos 500 Ma y que se mantienen hoy en día.
Extended parental care (XPC) is a complex reproductive strategy in which progenitors actively look after their offspring up to - or beyond - the first juvenile stage in order to maximize their fitness. Although the euarthropod fossil record has produced several examples of brood-care, the appearance of XPC within this phylum remains poorly constrained given the scarcity of developmental data for Palaeozoic stem-group representatives that would link juvenile and adult forms in an ontogenetic sequence. Here, we describe the post-embryonic growth of Fuxianhuia protensa from the early Cambrian Chengjiang Lagerstätte, and show parental care in this stem-group euarthropod. We recognize fifteen distinct ontogenetic stages based on the number and shape of the trunk tergites, and their allocation between the morphologically distinct thorax and abdomen. Our data demonstrate anamorphic post-embryonic development in F. protensa, in which tergites were sequentially added from a posterior growth zone. A life assemblage consisting of a sexually mature F. protensa adult alongside four ontogenetically coeval juveniles, constitutes the oldest occurrence of XPC in the panarthropod fossil record. These findings provide the most phylogenetically basal evidence of anamorphosis in the evolutionary history of total-group Euarthropoda, and reveal a complex post-embryonic reproductive ecology for its early representatives.
Euarthropods owe their evolutionary and ecological success to the morphological plasticity of their appendages. Although this variability is partly expressed in the specialization of the protopodite for a feeding function in the post-deutocerebral limbs, the origin of the former structure among Cambrian representatives remains uncertain. Here, we describe Alacaris mirabilis gen. et sp. nov. from the early Cambrian Xiaoshiba Lagerstätte in China, which reveals the proximal organization of fuxianhuiid appendages in exceptional detail. Proximally, the post-deutocerebral limbs possess an antero-posteriorly compressed protopodite with robust spines. The protopodite is attached to an endopod with more than a dozen podomeres, and an oval flap-shaped exopod. The gnathal edges of the protopodites form an axial food groove along the ventral side of the body, indicating a predatory/scavenging autecology. A cladistic analysis indicates that the fuxianhuiid protopodite represents the phylogenetically earliest occurrence of substantial proximal differentiation within stem-group Euarthropoda illuminating the origin of gnathobasic feeding.