The Palaeozoic form-taxon Lobopodia encompasses a diverse range of soft-bodied 'legged worms' known from exceptional fossil deposits. Although lobopodians occupy a deep phylogenetic position within Panarthropoda, a shortage of derived characters obscures their evolutionary relationships with extant phyla (Onychophora, Tardigrada and Euarthropoda). Here we describe a complex feature in the terminal claws of the mid-Cambrian lobopodian Hallucigenia sparsa--their construction from a stack of constituent elements--and demonstrate that equivalent elements make up the jaws and claws of extant Onychophora. A cladistic analysis, informed by developmental data on panarthropod head segmentation, indicates that the stacked sclerite components in these two taxa are homologous-resolving hallucigeniid lobopodians as stem-group onychophorans. The results indicate a sister-group relationship between Tardigrada and Euarthropoda, adding palaeontological support to the neurological and musculoskeletal evidence uniting these disparate clades. These findings elucidate the evolutionary transformations that gave rise to the panarthropod phyla, and expound the lobopodian-like morphology of the ancestral panarthropod.
The present contribution is the first instalment of a systematic revision of the Early Cretaceous (Barremian to Early Albian) corals of Puebla, Mexico. The coral fauna preserved in the Tehuacán region represents one of the most species rich associations of its kind from the Early Cretaceous of the New World. This article provides a brief overview of the research history, geology, lithostratigraphy and biostratigraphy, and reports the presence of corals of the family Stylinidae (suborder Stylinina). The family is represented by the genera Acanthocoenia (one species), Enallhelia (two species), Stylangia (one species) and Stylina (six species).
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 phylogenetic position of aglaspidids, a problematic group of Lower Palaeozoic arthropods of undetermined affinities, is re-examined in the context of the major Cambrian and Ordovician lamellipedian arthropod groups. A cladistic analysis of ten genera of aglaspidids sensu stricto, six aglaspidid-like arthropods and 42 Palaeozoic arthropod taxa indicates that Xenopoda, Cheloniellida, Aglaspidida sensu lato and Trilobitomorpha form a clade (Artiopoda Hou and Bergström, 1997) nested within the mandibulate stem-lineage, thus discarding previous interpretations of these taxa as part 'of the chelicerate stem-group (Arachnomorpha Heider, 1913). The results confirm an aglaspidid identity for several recently described arthropods, including Quasimodaspis brentsae, Tremaglaspis unite, Chlupacaris dubia, Australaglaspis stonyensis and an unnamed Ordovician Chinese arthropod. The problematic Bohemian arthropod Kodymirus vagans was recovered as sister taxon to Beckwithia typa, and both form a small clade that falls outside Aglaspidida sensu stricto, thus discarding eurypterid affinities for the former. The analysis does not support the phylogenetic position of Kwanyinaspis maotianshanensis at the base of Conciliterga as proposed in recent studies, but rather occupies a basal position within Aglaspidida sensu lato. The results indicate a close association of aglaspidid arthropods with xenopods (i.e. Emeraldella and Sidneyia) and cheloniellids (e.g. Cheloniellon, Duslia); the new clade “Vicissicaudata” is proposed to encompass these arthropods, which are characterized by a differentiated posterior region. The phylogenetic position of aglaspidid arthropods makes them good outgroup candidates for analysing the internal relationships within the groups that form Trilobitomorpha. This work provides a much clearer picture of the phylogenetic relationships among Lower Palaeozoic lamellipedians.
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, Hesselboniagen. nov.
Trilobites are typified by the behavioural and morphological ability to enrol their bodies, most probably as a defence mechanism against adverse environmental conditions or predators. Although most trilobites could enrol at least partially, there is uncertainty about whether olenellids-among the most phylogenetically and stratigraphically basal representatives-could perform this behaviour because of their poorly caudalized trunk and scarcity of coaptative devices. Here, we report complete-but not encapsulating-enrolment for the olenellid genus Mummaspis from the early Cambrian Mural Formation in Alberta, the earliest direct evidence of this strategy in the fossil record of polymerid trilobites. Complete enrolment in olenellids was achieved through a combination of ancestral morphological features, and thus provides new information on the character polarity associated with this key trilobite adaptation.
The enigmatic marrellomorph arthropod Furca bohemica from the Upper Ordovician Letná Formation, is redescribed. Based on existing museum specimens and new material collected from the southern slope of Ostrý Hill (Beroun, Czech Republic), the morphology and taphonomy of F. bohemica is reappraised and expanded to produce a new anatomical interpretation. The previously distinct taxa F. pilosa and Furca sp., are synonymised with F. bohemica, the latter being represented by a tapho-series in which decay has obscured some of the diagnostic features. A cladistic analysis indicates close affinities between F. bohemica and the Hunsrück Slate marrellomorph Mimetaster hexagonalis, together forming the Family Mimetasteridae, contrary to previous models for marrellomorph internal relationships. As with other representatives of the group, the overall anatomy of F. bohemica is consistent with a benthic, or possibly nektobenthic, mode of life. The depositional setting of the Letná Formation indicates that F. bohemica inhabited a shallow marine environment, distinguishing it palaeoecologically from all other known marrellomorphs, which have been reported from the continental shelf.
The organization of the head provides critical data for resolving the phylogenetic relationships and evolutionary history of extinct and extant euarthropods. The early Cambrian-period fuxianhuiids are regarded as basal representatives of stem-group Euarthropoda, and their anterior morphology therefore offers key insights for reconstructing the ancestral condition of the euarthropod head. However, the paired post-antennal structures in Fuxianhuia protensa remain controversial; they have been interpreted as both 'great appendages' and as gut diverticulae. Here we describe Chengjiangocaris kunmingensis sp. nov. and Fuxianhuia xiaoshibaensis sp. nov. from a new early Cambrian (Stage 3) fossil Lagerstatte in Yunnan, China. Numerous specimens of both species show a unique 'taphonomic dissection' of the anterodorsal head shield, revealing the cephalic organization in detail. We demonstrate the presence of a pair of specialized post-antennal appendages (SPAs) in the fuxianhuiid head, which attach at either side of the posteriorly directed mouth, behind the hypostome. Preserved functional articulations indicate a well-defined but restricted range of limb movement, suggestive of a simple type of sweep feeding. The organization of the SPAs in fuxianhuiids is incompatible with the (deutocerebral) anterior raptorial appendages of megacheirans, and argue against the presence of protocerebral limbs in the fuxianhuiids. The positions of the fuxianhuiid antennae and SPAs indicate that they are segmentally homologous to the deutocerebral and tritocerebral appendages of crown-group Euarthropoda respectively. These findings indicate that antenniform deutocerebral appendages with many podomeres are a plesiomorphic feature of the ancestral euarthropod head.
Diverse carbonaceous microfossils, including exceptionally preserved remains of non-biomineralizing metazoans, are reported from a basal middle Cambrian interval of the Kaili Formation (Guizhou Province, China). The application of a gentle acid maceration technique complements previous palynological studies by revealing a larger size-class of acritarchs, a richer assemblage of filamentous microfossils, and a variety of previously unrecovered forms. Metazoan fossils include Wiwaxia sclerites and elements derived from biomineralizing taxa, including chancelloriids, brachiopods and hyolithids, in common with previously studied assemblages from the early and middle Cambrian of Canada. In addition, the Kaili Formation has yielded pterobranch remains and an assemblage of cuticle fragments representing “soft-bodied” worms, including a priapulid-like scalidophoran. Our results demonstrate the wide distribution and palaeobiological importance of microscopic “Burgess Shale-type” fossils, and provide insights into the limitations and potential of this largely untapped preservational mode.
Trilobites have a rich and abundant fossil record, but little is known about the intrinsic mechanisms that orchestrate their body organization. To date, there is disagreement regarding the correspondence, or lack thereof, of the segmental units that constitute the trilobite trunk and their associated exoskeletal elements. The phylogenetic position of trilobites within total-group Euarthropoda, however, allows inferences about the underlying organization in these extinct taxa to be made, as some of the fundamental genetic processes for constructing the trunk segments are remarkably conserved among living arthropods. One example is the expression of the segment polarity gene engrailed, which at embryonic and early postembryonic stages is expressed in extant panarthropods (i.e. tardigrades, onychophorans, euarthropods) as transverse stripes that define the posteriormost region of each trunk segment. Due to its conservative morphology and allegedly primitive trunk tagmosis, we have utilized the centipede Strigamia maritima to study the correspondence between the expression of engrailed during late embryonic to postembryonic stages, and the development of the dorsal exoskeletal plates (i.e. tergites). The results corroborate the close correlation between the formation of the tergite borders and the dorsal expression of engrailed, and suggest that this association represents a symplesiomorphy within Euarthropoda. This correspondence between the genetic and phenetic levels enables making accurate inferences about the dorsoventral expression domains of engrailed in the trunk of exceptionally preserved trilobites and their close relatives, and is suggestive of the widespread occurrence of a distinct type of genetic segmental mismatch in these extinct arthropods. The metameric organization of the digestive tract in trilobites provides further support to this new interpretation. The wider evolutionary implications of these findings suggest the presence of a derived morphogenetic patterning mechanism responsible for the reiterated occurrence of different types of trunk dorsoventral segmental mismatch in several phylogenetically distant, extinct and extant, arthropod groups.
The aglaspidid arthropod, Australaglaspis stonyensis gen. et sp. nov., is described from the Idamean (Upper Cambrian) of Stony Point, Montagu, north-western Tasmania. The dorsal exoskeleton comprises a semicircular cephalon with a well-defined marginal rim, acute genal angles, oval eyes, subtriangular glabella area and a subtrapezoidal hypostome. The trunk bears eleven somites with well-developed pleural spines that progressively curve backwards. Paired postventral plates cover the last trunk tergites and the base of the tailspine, which is long and characterized by a medial cleft. Bilobed ventral impressions indicate the presence of homopodous appendages, forming a food groove along the trunk. The fossil assemblage and its mode of preservation suggest that Australaglaspis originally possessed a phosphatic cuticle, but chemical traces of it have been lost because of regional metamorphism and weathering. Australaglaspis appears to be closely related to the North American aglaspidid Chraspedops modesta Raasch, 1939, based on its medially cleft tailspine.