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.
The fuxianhuiids — of which there are eight species described so far —are a clade of extinct soft-bodied euarthropods (colloquially: arthropods) that are known from early Cambrian marine deposits in South China (Figure 1). They are widely regarded as distant relatives of extant euarthropod groups — namely arachnids, myriapods, crustaceans and insects. The exceptional preservation of their fossils has provided unique information on their morphology (Figure 2). Fuxianhuiids have contributed towards a better understanding of the early evolutionary radiation of euarthropods —the most successful animal phylum in the history of life on Earth — and led to a detailed reconstruction of the origin of the euarthropod body plan.
As part of a comprehensive examination of all radiodontans from Cambrian localities in the USA, Pates et al. (2017a, b) and Pates and Daley (2017) revised the taxonomic affinities of several described specimens. This included the reinterpretation of two putative lobopodians, one from the Wheeler Formation (Utah, USA) and one from the Valdemiedes Formation (Spain), as frontal appendages of the radiodontan genera Stanleycaris and Caryosyntrips respectively. In their comment, Gámez Vintaned and Zhuravlev (2018) disagree with these conclusions and raise three topics for discussion: (i) anatomical features they suggest support a lobopodian affinity for “Mureropodia”; (ii) the identity of Caryosyntrips as a radiodontan, and the assignment of certain specimens to this genus; and (iii) the nomenclatural status of Stanleycaris hirpex as an invalid taxon. For (i), we dispute that the anatomical features put forward by Gámez Vintaned and Zhuravlev (2018) are biological and conclude that a lobopodian affinity for Mureropodia is untenable. In response to (ii), we provide further evidence supporting a radiodontan affinity for Caryosyntrips, and those specimens ascribed to this genus. Finally, we concur with (iii) Stanleycaris as an invalid taxon according to the International Code on Zoological Nomenclature (ICZN), and have rectified the situation by providing a valid systematic description.
The Weeks Formation in Utah is the youngest (c. 499 Ma) and least studied Cambrian Lagerstätte of the western United States. It preserves a diverse exceptionally-preserved fauna that inhabited a relatively deep-water environment at the offshore margin of a carbonate platform, resembling the setting of the underlying Wheeler and Marjum formations. Yet, the Weeks fauna differs significantly in composition from the other remarkable biotas of Cambrian Series 3 of Utah, suggesting a significant Guzhangian faunal restructuring. This bioevent is regarded as the onset of a transitional episode in the history of life, separating the two primary diversifications of the Early Palaeozoic. The Weeks fossils have been strongly affected by late diagenetic processes, but some specimens still preserve exquisite anatomical details.
Aysheaia prolata, was described as the only lobopodian from the Drumian (Cambrian) Wheeler Formation in Utah, USA, and the sole representative of this genus besides the type species Aysheaia pedunculata, from the Cambrian (Stage 5) Stephen Formation, British Columbia. A redescription of Aysheaia prolata reveals previously overlooked morphological features, including segmental boundaries between putative lobopods, and curved terminal spines on the putative anterior end. These observations undermine lobopodian affinities of Aysheaia prolata, and instead we interpret this specimen as an isolated radiodontan frontal appendage. The presence of 11 podomeres, five of which possess elongate and anteriorly recurved ventral blades with auxiliary spines, together with shorter robust dorsal spines, identify the specimen as Stanleycaris. This represents the first report of Stanelycaris outside of the Cambrian Stage 5 thin Stephen Formation in British Columbia, expanding its palaeobiogeographic and stratigraphic range. Aysheaia is left as a monotypic genus endemic to the Burgess Shale. The Spence Shale luolishaniid Acinocrinus stichus is currently the only lobopodian known from the Cambrian of Utah.
The panarthropod head represents a complex body region that has evolved through the integration and functional specialization of the anterior appendage-bearing segments. Advances in the developmental biology of diverse extant organisms have led to a substantial clarity regarding the relationships of segmental homology between Onychophora (velvet worms), Tardigrada (water bears), and Euarthropoda (e.g. arachnids, myriapods, crustaceans, hexapods). The improved understanding of the segmental organization in panarthropods offers a novel perspective for interpreting the ubiquitous Cambrian fossil record of these successful animals. A combined palaeobiological and developmental approach to the study of the panarthropod head through deep time leads us to propose a consensus hypothesis for the intricate evolutionary history of this important tagma. The contribution of exceptionally preserved brains in Cambrian fossils - together with the recognition of segmentally informative morphological characters - illuminate the polarity for major anatomical features. The euarthropod stem-lineage provides a detailed view of the step-wise acquisition of critical characters, including the origin of a multiappendicular head formed by the fusion of several segments, and the transformation of the ancestral protocerebral limb pair into the labrum, following the postero-ventral migration of the mouth opening. Stem-group onychophorans demonstrate an independent ventral migration of the mouth and development of a multisegmented head, as well as the differentiation of the deutocerebral limbs as expressed in extant representatives. The anterior organization of crown-group Tardigrada retains several ancestral features, such as an anterior-facing mouth and one-segmented head. The proposed model aims to clarify contentious issues on the evolution of the panarthropod head, and lays the foundation from which to further address this complex subject in the future.
The somasteroids are Ordovician star-shaped animals widely regarded as ancestors of Asterozoa, the group of extant echinoderms that includes brittle stars and starfish. The phylogenetic position of somasteroids makes them critical for understanding the origin and early evolution of crown group Echinodermata. However, the early evolution of asterozoans, the origin of their distinctive body organization and their relationships with other Cambrian and Ordovician echinoderms, such as edrioasteroids, blastozoans, crinoids, and other asterozoans, remain problematic due to the difficulties of comparing the calcitic endoskeleton of these disparate groups. Here we describe the new somasteroid Cantabrigiasterfezouataensis from the Early Ordovician (Tremadocian) Fezouata Lagerstatte in Morocco. Cantabrigiaster shares with other somasteroids the presence of rod-like virgal ossicles that articulate with the ambulacrals, but differs from all other known asterozoans in the absence of adambulacral ossicles defining the arm margins. The unique arm construction evokes parallels with non-asterozoan echinoderms. Developmentally informed Bayesian and parsimony based phylogenetic analyses, which reflect the homology of the biserial ambulacral ossicles in Paleozoic echinoderms according to the Extraxial-Axial Theory, recover Cantabrigiaster as basal within stem group Asterozoa. Our results indicate that Cantabrigiaster is the earliest diverging stem group asterozoan, revealing the ancestral morphology of this major clade and clarifying the affinities of problematic Ordovician taxa. Somasteroids are resolved as a paraphyletic grade within stem and crown group Asterozoa (starfishes), whereas stenuroids are paraphyletic within stem group Ophiuroidea (brittle stars). Cantabrigiaster also illuminates the relationship between Ordovician crown group Echinodermata and its Cambrian stem lineage, which includes sessile forms with incipient radial symmetry such as edrioasteroids and blastozoans. The contentious Pelmatozoa hypothesis (i.e. monophyly of blastozoans and crinoids) is not supported; instead, blastozoans represent the most likely sister-taxon of crown group Echinodermata.
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-Lagerstatte 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.