Convergent adaptation of true crabs (Decapoda: Brachyura) to a gradient of terrestrial environments.

For much of terrestrial biodiversity, the evolutionary pathways of adaptation from marine ancestors are poorly understood, and have usually been viewed as a binary trait. True crabs, the decapod crustacean infraorder Brachyura, comprise over 7,600 species representing a striking diversity of morphology and ecology, including repeated adaptation to non-marine habitats. Here, we reconstruct the evolutionary history of Brachyura using new and published sequences of 10 genes for 344 tips spanning 88 of 109 brachyuran families. Using 36 newly vetted fossil calibrations, we infer that brachyurans most likely diverged in the Triassic, with family-level splits in the late Cretaceous and early Paleogene. By contrast, the root age is underestimated with automated sampling of 328 fossil occurrences explicitly incorporated into the tree prior, suggesting such models are a poor fit under heterogeneous fossil preservation. We apply recently defined trait-by-environment associations to classify a gradient of transitions from marine to terrestrial lifestyles. We estimate that crabs left the marine environment at least seven and up to 17 times convergently, and returned to the sea from non-marine environments at least twice. Although the most highly terrestrial- and many freshwater-adapted crabs are concentrated in Thoracotremata, Bayesian threshold models of ancestral state reconstruction fail to identify shifts to higher terrestrial grades due to the degree of underlying change required. Lineages throughout our tree inhabit intertidal and marginal marine environments, corroborating the inference that the early stages of terrestrial adaptation have a lower threshold to evolve. Our framework and extensive new fossil and natural history datasets will enable future comparisons of non-marine adaptation at the morphological and molecular level. Crabs provide an important window into the early processes of adaptation to novel environments, and different degrees of evolutionary constraint that might help predict these pathways.

Post-Cambrian survival of the tubicolous scalidophoran Selkirkia.

Scalidophoran worms represent common infaunal components of early and middle Cambrian Burgess Shale-type fossil biotas. Early scalidophorans resemble extant priapulids based on overall morphology, but the genus Selkirkia represents the earliest record of tube dwelling for the group. Despite its ubiquitous presence in exceptional marine deposits, whether the exclusively Cambrian occurrence of Selkirkia reflects its entire evolutionary history or is affected by taphonomic biases remains unresolved. Here, we demonstrate the post-Cambrian survival of Selkirkia based on new material from the Lower Ordovician Fezouata Shale biota of Morocco. The discovery of Selkirkia in the Fezouata Shale extends the biostratigraphic range of the genus by 25 million years and its palaeobiogeographic occurrence to the high latitudes of Gondwana, strengthens the evolutionary links between Cambrian and Ordovician Burgess Shale-type biotas and increases scalidophoran diversity for the Fezouata Shale biota otherwise consisting exclusively of the palaeoscolecid Palaeoscolex? tenensis. The tube of Selkirkia underwent negligible external change for over 40 million years, indicating a high degree of morphological stasis during the Early Palaeozoic. A tubicolous mode of life is rare among extant priapulids and expressed only in Maccabeus, which forms a delicate tube from agglutinated plant debris, unlike the macroscopic secreted cuticular tube of Selkirkia.

Convergent evolution of ventral adaptations for enrolment in trilobites and extant euarthropods.

The ability to enrol for protection is an effective defensive strategy that has convergently evolved multiple times in disparate animal groups ranging from euarthropods to mammals. Enrolment is a staple habit of trilobites, and their biomineralized dorsal exoskeleton offered a versatile substrate for the evolution of interlocking devices. However, it is unknown whether trilobites also featured ventral adaptations for enrolment. Here, we report ventral exoskeletal adaptations that facilitate enrolment in exceptionally preserved trilobites from the Middle Ordovician Walcott-Rust Quarry in New York State, USA. Walcott-Rust trilobites reveal the intricate three-dimensional organization of the non-biomineralized ventral anatomy preserved as calcite casts, including the spatial relationship between the articulated sternites (i.e. ventral exoskeletal plates) and the wedge-shaped protopodites. Enrolment in trilobites is achieved by ventrally dipping the anterior margin of the sternites during trunk flexure, facilitated by the presence of flexible membranes, and with the close coupling of the wedge-shaped protopodites. Comparisons with the ventral morphology of extant glomerid millipedes and terrestrial isopods reveal similar mechanisms used for enrolment. The wedge-shaped protopodites of trilobites closely resemble the gnathobasic coxa/protopodite of extant horseshoe crabs. We propose that the trilobites' wedge-shaped protopodite simultaneously facilitated tight enrolment and gnathobasic feeding with the trunk appendages.

Three-dimensional morphology of the biramous appendages in Isoxys from the early Cambrian of South China, and its implications for early euarthropod evolution.

Early euarthropod evolution involved a major transition from lobopodian-like taxa to organisms featuring a segmented, well-sclerotized trunk (arthrodization) and limbs (arthropodization). However, the precise origin of a completely arthrodized trunk and arthropodized ventral biramous appendages remain controversial, as well as the early onset of anterior-posterior limb differentiation in stem-group euarthropods. New fossil material and micro-computed tomography inform the detailed morphology of the arthropodized biramous appendages in the carapace-bearing euarthropod Isoxys curvirostratus from the early Cambrian Chengjiang biota. In addition to well-developed grasping frontal appendages, I. curvirostratus possesses two batches of morphologically and functionally distinct biramous limbs. The first batch consists of four pairs of short cephalic appendages with robust endites with a feeding function, whereas the second batch has more elongate trunk appendages for locomotion. Critically, our new material shows that the trunk of I. curvirostratus was not arthrodized. The results of our phylogenetic analyses recover isoxyids as some of the earliest branching sclerotized euarthropods, and strengthens the hypothesis that arthropodized biramous appendages evolved before full body arthrodization.

New opabiniid diversifies the weirdest wonders of the euarthropod stem group.

Once considered 'weird wonders' of the Cambrian, the emblematic Burgess Shale animals Anomalocaris and Opabinia are now recognized as lower stem-group euarthropods and have provided crucial data for constraining the polarity of key morphological characters in the group. Anomalocaris and its relatives (radiodonts) had worldwide distribution and survived until at least the Devonian. However, despite intense study, Opabinia remains the only formally described opabiniid to date. Here we reinterpret a fossil from the Wheeler Formation of Utah as a new opabiniid, Utaurora comosa nov. gen. et sp. By visualizing the sample of phylogenetic topologies in treespace, our results fortify support for the position of U. comosa beyond the nodal support traditionally applied. Our phylogenetic evidence expands opabiniids to multiple Cambrian stages. Our results underscore the power of treespace visualization for resolving imperfectly preserved fossils and expanding the known diversity and spatio-temporal ranges within the euarthropod lower stem group.

A tardigrade in Dominican amber.

Tardigrades are a diverse group of charismatic microscopic invertebrates that are best known for their ability to survive extreme conditions. Despite their long evolutionary history and global distribution in both aquatic and terrestrial environments, the tardigrade fossil record is exceedingly sparse. Molecular clocks estimate that tardigrades diverged from other panarthropod lineages before the Cambrian, but only two definitive crown-group representatives have been described to date, both from Cretaceous fossil deposits in North America. Here, we report a third fossil tardigrade from Miocene age Dominican amber. Paradoryphoribius chronocaribbeus gen. et sp. nov. is the first unambiguous fossil representative of the diverse superfamily Isohypsibioidea, as well as the first tardigrade fossil described from the Cenozoic. We propose that the patchy tardigrade fossil record can be explained by the preferential preservation of these microinvertebrates as amber inclusions, coupled with the scarcity of fossiliferous amber deposits before the Cretaceous.

Biomechanical analyses of Cambrian euarthropod limbs reveal their effectiveness in mastication and durophagy.

Durophagy arose in the Cambrian and greatly influenced the diversification of biomineralized defensive structures throughout the Phanerozoic. Spinose gnathobases on protopodites of Cambrian euarthropod limbs are considered key innovations for shell-crushing, yet few studies have demonstrated their effectiveness with biomechanical models. Here we present finite-element analysis models of two Cambrian trilobites with prominent gnathobases-Redlichia rex and Olenoides serratus-and compare these to the protopodites of the Cambrian euarthropod Sidneyia inexpectans and the modern American horseshoe crab, Limulus polyphemus. Results show that L. polyphemus, S. inexpectans and R. rex have broadly similar microstrain patterns, reflecting effective durophagous abilities. Conversely, low microstrain values across the O. serratus protopodite suggest that the elongate gnathobasic spines transferred minimal strain, implying that this species was less well-adapted to masticate hard prey. These results confirm that Cambrian euarthropods with transversely elongate protopodites bearing short, robust gnathobasic spines were likely durophages. Comparatively, taxa with shorter protopodites armed with long spines, such as O. serratus, were more likely restricted to a soft food diet. The prevalence of Cambrian gnathobase-bearing euarthropods and their various feeding specializations may have accelerated the development of complex trophic relationships within early animal ecosystems, especially the 'arms race' between predators and biomineralized prey.

A new somasteroid from the Fezouata Lagerstätte in Morocco and the Early Ordovician origin of Asterozoa.

The somasteroids are Lower Palaeozoic star-shaped animals widely regarded as ancestors of Asterozoa, the group of echinoderms that includes brittle stars and starfish. However, the origin of asterozoans, the assembly of their distinctive body organization, and their relationships with other Cambrian and Ordovician echinoderms remain problematic owing to the difficulties of comparing the endoskeleton between disparate groups. Here, we describe the new somasteroid Cantabrigiaster fezouataensis, a primitive asterozoan from the Early Ordovician Fezouata Lagerstätte in Morocco. Cantabrigiaster shares with other somasteroids a unique endoskeletal arm organization and 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, evoking parallels with non-asterozoan echinoderms. Developmentally informed Bayesian and parsimony phylogenetic analyses, which reflect the homology of the biserial ambulacral ossicles in Palaeozoic echinoderms according to the extraxial-axial theory, recover Cantabrigiaster as the earliest divergent stem-group asterozoan. Our results illuminate the ancestral morphology of Asterozoa, and clarify the affinities of problematic Ordovician Asterozoa. Bayesian inference and parsimony demonstrate that somasteroids represent a paraphyletic grade within stem- and crown-group Asterozoa, whereas stenuroids are paraphyletic within stem-group Ophiuroidea. Our results also offer potential insights on the evolutionary relationships between asterozoans, crinoids and potential Cambrian stem-group representatives.

Computed tomography sheds new light on the affinities of the enigmatic euarthropod Jianshania furcatus from the early Cambrian Chengjiang biota.

BACKGROUND: The Chengjiang biota is one of the most species-rich Cambrian Konservat-Lagerstätten, and preserves a community dominated by non-biomineralized euarthropods. However, several Chengjiang euarthropods have an unfamiliar morphology, are extremely rare, or incompletely preserved. RESULTS: We employed micro-computed tomography to restudy the enigmatic euarthropod Jianshania furcatus. We reveal new morphological details, and demonstrate that the specimens assigned to this species represent two different taxa. The holotype of J. furcatus features a head shield with paired anterolateral notches, stalked lateral eyes, and an articulated tailspine with a bifurcate termination. The other specimen is formally redescribed as Xiaocaris luoi gen. et sp. nov., and is characterized by stalked eyes connected to an anterior sclerite, a subtrapezoidal head shield covering three small segments with reduced tergites, a trunk with 15 overlapping tergites with a well-developed dorsal keel, and paired tail flukes. CONCLUSIONS: The presence of antennae, biramous appendages with endopods composed of 15 articles, and multiple appendage pairs associated with the trunk tergites identify X. luoi nov. as a representative of Fuxianhuiida, an early branching group of stem-group euarthropods endemic to the early Cambrian of Southwest China. X. luoi nov. represents the fifth fuxianhuiid species described from the Chengjiang biota, and its functional morphology illuminates the ecological diversity of this important clade for understanding the early evolutionary history of euarthropods.

Hallucigenia's onychophoran-like claws and the case for Tactopoda.

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 appendicular morphology of Sinoburius lunaris and the evolution of the artiopodan clade Xandarellida (Euarthropoda, early Cambrian) from South China.

BACKGROUND: 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. RESULTS: 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. CONCLUSIONS: 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.

Proclivity of nervous system preservation in Cambrian Burgess Shale-type deposits.

Recent investigations on neurological tissues preserved in Cambrian fossils have clarified the phylogenetic affinities and head segmentation in pivotal members of stem-group Euarthropoda. However, palaeoneuroanatomical features are often incomplete or described from single exceptional specimens, raising concerns about the morphological interpretation of fossilized neurological structures and their significance for early euarthropod evolution. Here, we describe the central nervous system (CNS) of the short great-appendage euarthropod Alalcomenaeus based on material from two Cambrian Burgess Shale-type deposits of the American Great Basin, the Pioche Formation (Stage 4) and the Marjum Formation (Drumian). The specimens reveal complementary ventral and lateral views of the CNS, preserved as a dark carbonaceous compression throughout the body. The head features a dorsal brain connected to four stalked ventral eyes, and four pairs of segmental nerves. The first to seventh trunk tergites overlie a ventral nerve cord with seven ganglia, each associated with paired sets of segmental nerve bundles. Posteriorly, the nerve cord features elongate thread-like connectives. The Great Basin fossils strengthen the original description-and broader evolutionary implications-of the CNS in Alalcomenaeus from the early Cambrian (Stage 3) Chengjiang deposit of South China. The spatio-temporal recurrence of fossilized neural tissues in Cambrian Konservat-Lagerstätten across North America (Pioche, Burgess Shale, Marjum) and South China (Chengjiang, Xiaoshiba) indicates that their preservation is consistent with the mechanism of Burgess Shale-type fossilization, without the need to invoke alternative taphonomic pathways or the presence of microbial biofilms.

A soft-bodied euarthropod from the early Cambrian Xiaoshiba Lagerstätte of China supports a new clade of basal artiopodans with dorsal ecdysial sutures.

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.

Specialized appendages in fuxianhuiids and the head organization of early euarthropods.

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 Lagerstätte 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.

Fuxianhuiid ventral nerve cord and early nervous system evolution in Panarthropoda.

Panarthropods are typified by disparate grades of neurological organization reflecting a complex evolutionary history. The fossil record offers a unique opportunity to reconstruct early character evolution of the nervous system via exceptional preservation in extinct representatives. Here we describe the neurological architecture of the ventral nerve cord (VNC) in the upper-stem group euarthropod Chengjiangocaris kunmingensis from the early Cambrian Xiaoshiba Lagerstätte (South China). The VNC of C. kunmingensis comprises a homonymous series of condensed ganglia that extend throughout the body, each associated with a pair of biramous limbs. Submillimetric preservation reveals numerous segmental and intersegmental nerve roots emerging from both sides of the VNC, which correspond topologically to the peripheral nerves of extant Priapulida and Onychophora. The fuxianhuiid VNC indicates that ancestral neurological features of Ecdysozoa persisted into derived members of stem-group Euarthropoda but were later lost in crown-group representatives. These findings illuminate the VNC ground pattern in Panarthropoda and suggest the independent secondary loss of cycloneuralian-like neurological characters in Tardigrada and Euarthropoda.

Anamorphic development and extended parental care in a 520 million-year-old stem-group euarthropod from China.

BACKGROUND: Extended parental care 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 extended parental care 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. RESULTS: Here, we describe the post-embryonic growth of Fuxianhuia protensa from the early Cambrian Chengjiang Lagerstätte in South China. Our data demonstrate anamorphic post-embryonic development for F. protensa, in which new tergites were sequentially added from a posterior growth zone, the number of tergites varies from eight to 30. The growth of F. protensa is typified by the alternation between segment addition, followed by the depletion of the anteriormost abdominal segment into the thoracic region. The transformation of abdominal into thoracic tergite is demarcated by the development of laterally tergopleurae, and biramous walking legs. The new ontogeny data leads to the recognition of the rare Chengjiang euarthropod Pisinnocaris subconigera as a junior synonym of Fuxianhuia. Comparisons between different species of Fuxianhuia and with other genera within Fuxianhuiida suggest that heterochrony played a prominent role in the morphological diversification of fuxianhuiids. Functional analogy with the flexible trunk ontogeny of Cambrian and Silurian olenimorphic trilobites suggests an adaptation to sporadic low oxygen conditions in Chengjiang deposits for F. protensa. Finally, understanding the growth of F. protensa allows for the interpretation of an exceptional life assemblage consisting of a sexually mature adult alongside four ontogenetically coeval juveniles, which constitutes the oldest occurrence of extended parental care by prolonged cohabitation in the panarthropod fossil record. CONCLUSIONS: Our findings constitute the most detailed characterization of the post-embryonic development in a soft-bodied upper stem-group euarthropod available to date. The new ontogeny data illuminates the systematics, trunk segmentation and palaeoecology of F. protensa, offers insights on the macroevolutionary processes involved in the diversification of this clade, and contributes towards an improved understanding of complex post-embryonic reproductive ecology in Cambrian euarthropods.

A superarmored lobopodian from the Cambrian of China and early disparity in the evolution of Onychophora.

We describe Collinsium ciliosum from the early Cambrian Xiaoshiba Lagerstätte in South China, an armored lobopodian with a remarkable degree of limb differentiation including a pair of antenna-like appendages, six pairs of elongate setiferous limbs for suspension feeding, and nine pairs of clawed annulated legs with an anchoring function. Collinsium belongs to a highly derived clade of lobopodians within stem group Onychophora, distinguished by a substantial dorsal armature of supernumerary and biomineralized spines (Family Luolishaniidae). As demonstrated here, luolishaniids display the highest degree of limb specialization among Paleozoic lobopodians, constitute more than one-third of the overall morphological disparity of stem group Onychophora, and are substantially more disparate than crown group representatives. Despite having higher disparity and appendage complexity than other lobopodians and extant velvet worms, the specialized mode of life embodied by luolishaniids became extinct during the Early Paleozoic. Collinsium and other superarmored lobopodians exploited a unique paleoecological niche during the Cambrian explosion.

The phylogeny of aglaspidid arthropods and the internal relationships within Artiopoda.

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.

Humble origins for a successful strategy: complete enrolment in early Cambrian olenellid trilobites.

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.

A mid-Cambrian tunicate and the deep origin of the ascidiacean body plan.

Tunicates are an evolutionarily significant subphylum of marine chordates, with their phylogenetic position as the sister-group to Vertebrata making them key to unraveling our own deep time origin. Tunicates greatly vary with regards to morphology, ecology, and life cycle, but little is known about the early evolution of the group, e.g. whether their last common ancestor lived freely in the water column or attached to the seafloor. Additionally, tunicates have a poor fossil record, which includes only one taxon with preserved soft-tissues. Here we describe Megasiphon thylakos nov., a 500-million-year-old tunicate from the Marjum Formation of Utah, which features a barrel-shaped body with two long siphons and prominent longitudinal muscles. The ascidiacean-like body of this new species suggests two alternative hypotheses for early tunicate evolution. The most likely scenario posits M. thylakos belongs to stem-group Tunicata, suggesting that a biphasic life cycle, with a planktonic larva and a sessile epibenthic adult, is ancestral for this entire subphylum. Alternatively, a position within the crown-group indicates that the divergence between appendicularians and all other tunicates occurred 50 million years earlier than currently estimated based on molecular clocks. Ultimately, M. thylakos demonstrates that fundamental components of the modern tunicate body plan were already established shortly after the Cambrian Explosion.