The origin of crustaceans: new evidence from the Early Cambrian of China.

One of the smallest arthropods recently discovered in the Early Cambrian Maotianshan Shale Lagerstätte is described. Ercaia gen. nov. has an untagmatized trunk bearing serially repeated biramous appendages (long and segmented endopods and flap-like exopods), a head with an acron bearing stalked lateral eyes and a sclerite and two pairs of antennae. The position of this 520 million-year-old tiny arthropod within the Crustacea is supported by several anatomical features: (i) a head with five pairs of appendages including two pairs of antennae, (ii) highly specialized antennae (large setose fans with a possible function in feeding), and (iii) specialized last trunk appendages (segmented pediform structures fringed with setae). The segmentation pattern of Ercaia (5 head and 13 trunk) is close to that of Maxillopoda but lacks the trunk tagmosis of modern representatives of the group. Ercaia is interpreted as a possible derivative of the stem group Crustacea. Ercaia is likely to have occupied an ecological niche similar to those of some Recent meiobenthic organisms (e.g. copepods living in association with sediment). This new fossil evidence supports the remote ancestry of crustaceans well before the Late Cambrian and shows, along with other fossil data (mainly Early Cambrian in China), that a variety of body plans already coexisted among the primitive crustacean stock.     

The morphology of Opabinia regalis and the reconstruction of the arthropod stem-group

Opabinia regalis Walcott is an enigmatic fossil from the Middle Cambrian Burgess Shale of uncertain affinities. Recent suggestions place it in a clade with Anomalocaris Whiteaves from the Burgess Shale and Kerygmachela Budd from the Greenlandic Sirius Passet Fauna; these taxa have been interpreted as 'lobopods'. Consideration of available Opabinia specimens demonstrates that reflective extensions from the axial region, previously thought to be either gut diverticula or musculature, can be accommodated in neither the trunk nor the lateral lobes that arise from it. They must therefore be external structures independent of the lateral lobes. On the basis of their sub-triangular appearance, size and taphonomy, they are considered here to represent lobopod limbs. Some evidence for the existence of terminal claws is also presented. The question of whether Kerygmachela, Opabinia and Anomalocaris constitute a monophyletic or paraphyletic grouping is considered. While they share several characters, most of these are plesiomorphies. Further, Opabinia and Anomalocaris share several arthropod-like characters not possessed by Kerygmachela. It is concluded that these three taxa probably form a paraphyletic grouping at the base of the arthropods. Retention of lobopod-like characters within the group provides important documentation of the lobopod-arthropod transition. A proper understanding of Opabinia and its close relatives, which may include the tardigrades, opens the way for a reconstruction of the arthropod stem-group. This in turn allows the construction of a speculative but satisfying scenario for the evolution of major arthropod features, including the origin of the biramous limb, tergites and arthropod segmentation. 'Arthropodization' may thus be seen not to be a single event but a series of adaptive innovations. OPABINIA, ANOMALOCARIS, KERYGMACHELA, Burgess Shale, problematica, Lobopodia, Arthropoda.     

A pyritized polychaete from the Devonian of Ontario

A polychaete from the Middle Devonian Arkona Shale at Hungry Hollow, Arkona, Ontario is preserved in three dimensions in pyrite. The prostomium bears a single median antenna, a pair of lateral antennae and a pair of ventral palps. It is assumed to be fused to a reduced peristomium. The anteriormost three pairs of trunk appendages are modified as tentacular cirri, the third long and biramous. The remainder of the finely annulated trunk bears at least 21 similar biramous parapodia, some of which preserve evidence of chaetae. The postsegmental pygidium is very small and may bear up to two pairs of cirri. The polychaete, Arkonips topororum, falls within the Palpata, Aciculata, among the crown group Phyllodocida. Its remarkable preservation highlights the potential of the Arkona Shale to yield other examples of soft-tissue preservation.     

The bivalved arthropods Isoxys and Tuzoia with soft‐part preservation from the Lower Cambrian Emu Bay Shale Lagerstätte (Kangaroo Island,Australia)

Abstract: Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft‐bodied features previously unknown from this Burgess Shale‐type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better‐known Lagerstätten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada.     

A New Bivalved Arthropod from the Early Cambrian Sirius Passet Fauna, North Greenland

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five‐segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.     

Cambrian stem-group annelids and a metameric origin of the annelid head

The oldest fossil annelids come from the Early Cambrian Sirius Passet and Guanshan biotas and Middle Cambrian Burgess Shale. While these are among the best preserved polychaete fossils, their relationship to living taxa is contentious, having been interpreted either as members of extant clades or as a grade outside the crown group. New morphological observations from five Cambrian species include the oldest polychaete with head appendages, a new specimen of Pygocirrus from Sirius Passet, and an undescribed form from the Burgess Shale. We propose that the palps of Canadia are on an anterior segment bearing neuropodia and that the head of Phragmochaeta is formed of a segment bearing biramous parapodia and chaetae. The unusual anatomy of these taxa suggests that the head is not differentiated into a prostomium and peristomium, that palps are derived from a modified parapodium and that the annelid head was originally a parapodium-bearing segment. Canadia, Phragmochaeta and the Marble Canyon annelid share the presence of protective notochaetae, interpreted as a primitive character state subsequently lost in Pygocirrus and Burgessochaeta, in which the head is clearly differentiated from the trunk.     

The nature and significance of the appendages of Opabinia from the Middle Cambrian Burgess Shale

Opabinia regalis has long been regarded as a curious animal, with its five eyes, its long flexible anterior process, and gill lamellae carried on the outside of overlapping lateral lobes. More recently, Opabinia has been reconstructed with lobopod limbs lying adaxial but separate from the lateral lobes. This version of Opabinia represented a lobopod–arthropod transition and prompted a hypothesis for the origin of the biramous limb that involved uniting the lobopod limb with a lateral lobe. New evidence of elemental maps is consistent with previous interpretations of the triangular structures in Opabinia as lateral extensions of the gut; there is no convincing evidence for the presence of lobopod limbs. Re-examination of critical specimens reveals that the gill lamellae are not on the outside of the lateral lobes. The limbs of Opabinia resemble the phyllopodous exopod of arthropods; the posterior margin is fringed with blades. Opabinia remains on the stem of euarthropods but not as a part of a paraphyletic Lobopodia. The Lobopodia is a clade of Cambrian armoured lobopods and onychophorans. A new hypothesis for the origin of the arthropod biramous limb from an exopod like that in Opabinia is presented, which involves an endite-bearing phyllopodous limb as an intermediate stage.     

The Early Cambrian colonization of pelagic niches exemplified by Isoxys (Arthropoda)

The anatomy of the bivalved arthropod Isoxys (Early and Middle Cambrian) is reconstructed, based on new evidence from soft parts and exoskeletal design and on a critical review of previous work. Isoxys had a long segmented body flanked with a pair of short antennules, followed by a series of 14 biramous appendages provided with long paddle-like exopods concealed under a widely open bivalved carapace folded dorsally and bearing long cardinal spines. The close resemblance between Isoxys and Recent pelagic crustaceans (halocyprid ostracods, larval stages of malacostracans) indicates that Isoxys was probably an active epipelagic swimmer (evidence from soft parts, carapace design and distributional pattern). Some species (e.g. I. auritus and I. paradoxus from the Maotianshan Shale biota; Early Cambrian) may have lived in the vicinity of the bottom either permanently or temporarily, whereas others may have had ecological preferences for more open-marine settings. The spinosity of Isoxys had a possible role in predatorial deterrence rather than in buoyancy control or in retarding sinking within the water column. The presence of Isoxys in the Maotianshan Shale of S. China indicates that arthropods had already colonized midwater niches by the Early Cambrian. The midwater communities of the Maotianshan Shale comprised numerous other invertebrates, such as abundant medusiform eldonids, vetulicolids, chordates and possibly early vertebrates. This contradicts the opinion that pelagic communities remained poorly developed until late Cambrian/Ordovician times and that the occupation of the midwater niches largely post-dates the initial diversification of the benthic faunas.     

A Silurian 'marrellomorph' arthropod

Xylokorys chledophilia, a new arthropod with three-dimensionally preserved soft tissues, is described from the Herefordshire (Silurian) Lagerstätte of England. The head and trunk are covered by a relatively featureless ovoid carapace, which comprises a domed central part and a flange-like border. The head bears five pairs of appendages. The first is uniramous, with dorsal and ventral projections distally. Appendages two to four are biramous and each endopod terminates in two projections. Appendage five is possibly biramous. The hypostome is very long and subrectangular in outline. There are approximately 35 pairs of biramous trunk appendages. Each exopod comprises a long slender shaft bearing numerous fine filaments; each endopod comprises a ribbon-like shaft bearing paddle-like endites. Morphological comparisons and cladistic analyses of X. chledophilia indicate affinity with Vachonisia rogeri from the Lower Devonian Hunsrück Slate, within the marrellomorphs, but assignment to Marrellomorpha is provisional pending revision of other members of this clade. Xylokorys is the first ‘marrellomorph’ to be reported from the Silurian. It is interpreted as a benthic particle filter feeder, which may also have consumed prey items.     

Martinssonia elongata gen. et sp.n., a crustacean-like euarthropod from the Upper Cambrian 'Orsten' of Sweden

A tiny arthropod, with five growth stages, is described. Three of the instars are metanauplius-like larvae, having unsegmented bodies and four pairs of appendages. The largest stage, with a length of about 1.5 mm, may still be immature. Its body is divided into three tagmata. The cephalon, including five appendiculate segments, h a projecting forehead with a rostral spine and a small shield with a joint between fourth and fifth segments. Eyes are absent. The trunk is composed of seven annular segments, the anterior two with appendages. The caudal end is a long pleotelson-like segment with the anus on its ventral surface. There are seven pairs of appendages: uniramous antennulae, composed of few tubular podomeres; four pairs of biramous postantennular, almost homeomorphic cephalic appendages; two pairs on the trunk, the anterior pair being similar to the cephalic appendages except for the exopodite, the posterior being much smaller, uniramous and apparently rudimentary. Martinssonia was probably benthic, feeding on detritic particles which it stirred up from the bottom. Besides various crustacean-like features, the new form reveals structures different from Crustacea as well as from all other known arthropodan groups. Martinssonia presumably is a descendant of an euarthropodan group, originating from the crustacean branch long before reaching the eucrustacean level of evolution.     

Cephalic and Limb Anatomy of a New Isoxyid from the Burgess Shale and the Role of “Stem Bivalved Arthropods” in the Disparity of the Frontalmost Appendage

We herein describe Surusicaris elegans gen. et sp. nov. (in Isoxyidae, amended), a middle (Series 3, Stage 5) Cambrian bivalved arthropod from the new Burgess Shale deposit of Marble Canyon (Kootenay National Park, British Columbia). Surusicaris exhibits 12 simple, partly undivided biramous trunk limbs with long tripartite caeca, which may illustrate a plesiomorphic “fused” condition of exopod and endopod. We construe also that the head is made of five somites (= four segments), including two eyes, one pair of anomalocaridid-like frontalmost appendages, and three pairs of poorly sclerotized uniramous limbs. This fossil may therefore be a candidate for illustrating the origin of the plesiomorphic head condition in euarthropods, and questions the significance of the “two-segmented head” in, e.g., fuxianhuiids. The frontalmost appendage in isoxyids is intriguingly disparate, bearing similarities with both dinocaridids and euarthropods. In order to evaluate the relative importance of bivalved arthropods, such as Surusicaris, in the hypothetical structuro-functional transition between the dinocaridid frontal appendage and the pre-oral—arguably deutocerebral—appendage of euarthropods, we chose a phenetic approach and computed morphospace occupancy for the frontalmost appendages of 36 stem and crown taxa. Results show different levels of evolutionary decoupling between frontalmost appendage disparity and body plans. Variance is greatest in dinocaridids and “stem bivalved” arthropods, but these groups do not occupy the morphospace homogeneously. Rather, the diversity of frontalmost appendages in “stem bivalved” arthropods, distinct in its absence of clear clustering, is found to link the morphologies of “short great appendages,” chelicerae and antennules. This find fits the hypothesis of an increase in disparity of the deutocerebral appendage prior to its diversification in euarthropods, and possibly corresponds to its original time of development. The analysis of this pattern, however, is sensitive to the—still unclear—extent of polyphyly of the “stem bivalved” taxa.     

Skimming the surface with Burgess Shale arthropod locomotion

The first arthropod trackways are described from the Middle Cambrian Burgess Shale Formation of Canada. Trace fossils, including trackways, provide a rich source of biological and ecological information, including direct evidence of behaviour not commonly available from body fossils alone. The discovery of large arthropod trackways is unique for Burgess Shale-type deposits. Trackway dimensions and the requisite number of limbs are matched with the body plan of a tegopeltid arthropod. Tegopelte, one of the rarest Burgess Shale animals, is over twice the size of all other benthic arthropods known from this locality, and only its sister taxon, Saperion, from the Lower Cambrian Chengjiang biota of China, approaches a similar size. Biomechanical trackway analysis demonstrates that tegopeltids were capable of rapidly skimming across the seafloor and, in conjunction with the identification of gut diverticulae in Tegopelte, supports previous hypotheses on the locomotory capabilities and carnivorous mode of life of such arthropods. The trackways occur in the oldest part (Kicking Horse Shale Member) of the Burgess Shale Formation, which is also known for its scarce assemblage of soft-bodied organisms, and indicate at least intermittent oxygenated bottom waters and low sedimentation rates.     

An early Cambrian chelicerate from the Emu Bay Shale,South Australia

The Emu Bay Shale Lagerstätte (Cambrian Series 2, Stage 4) occurs on the north coast of Kangaroo Island, South Australia. Over 50 species are known from here, including trilobites and non‐biomineralized arthropods, palaeoscolecids, a lobopodian, a polychaete, vetulicolians, nectocaridids, hyoliths, brachiopods, sponges and chancelloriids. A new chelicerate, Wisangocaris barbarahardyae gen. et sp. nov., is described herein, based on a collection of some 270 specimens. It is up to 60 mm long, with the length of the cephalic shield comprising about 30% that of the exoskeleton. The cephalic margin has three pairs of bilaterally‐symmetrical small triangular spines. A pair of small eyes is placed well forwards on the ventral margin of the cephalic shield. The trunk comprises 11 segments that increase in length while narrowing posteriorly, each possibly bearing a pair of biramous appendages; the most posterior segment is almost square whereas the others are transversely elongated. The spatulate telson is proportionately longer than in taxa such as Sanctacaris, Utahcaris and Leanchoilia. Up to eight (?four pairs) of 3 mm‐long elements bearing alternating inward‐curving short and long spines beneath the cephalic shield are interpreted as basipodal gnathobases, part of a complex feeding apparatus. A well‐developed gut includes a stomach within the cephalic shield; it extends to the base of the telson. In a few specimens there are shell fragments within the gut, including those of the trilobite Estaingia bilobata (the most common species in the biota); these fragments have sharp margins and extend across the gut lumen. The species may have been a predator or a scavenger, ingesting material already broken up by a larger predator/scavenger. The morphology of this taxon shares many overall body features with Sanctacaris, and some with Sidneyia, particularly its gnathobasic complex. These chelicerate affinities are corroborated by phylogenetic analyses.     

New Middle Cambrian bivalved arthropods from the Burgess Shale (British Columbia,Canada)

The morphology of two new bivalved arthropods, Loricicaris spinocaudatus gen. et sp. nov. and Nereocaris briggsi sp. nov. from the middle Cambrian (Series 3, Stage 5) Burgess Shale Formation (Collins Quarry locality on Mount Stephen, Yoho National Park, British Columbia, Canada), is described. The material was originally assigned to the genus Branchiocaris, but exhibits distinctive character combinations meriting its assignment to other taxa. Loricicaris spinocaudatus possesses an elongate and spinose abdomen comparable to the contemporaneous Perspicaris and Canadaspis, as well as chelate second head appendages and subtriangular exopods, comparable to Branchiocaris. Nereocaris briggsi possesses a laterally compressed carapace, elongate and delicate appendages and a medial eye located between a pair of lateral eyes on a rhomboidal eye stalk. Although undoubtedly congeneric with Nereocaris exilis from a slightly younger horizon of the Burgess Shale Formation, N. briggsi differs in overall proportions and segment number, warranting assignment to a new species. The newly described taxa were coded into an extensive cladistic analysis of 755 characters, and 312 extinct and extant panarthropods, including a variety of Cambrian bivalved arthropods from both the Burgess Shale and the Chengjiang Lagerstätten. Cambrian bivalved arthropods consistently resolved as a paraphyletic assemblage at the base of Arthropoda. Important innovations in arthropod history such as the specialization of the deutocerebral head appendages and a shift from a nekton‐benthic deposit feeding habit to a benthic scavenging/predatory habit, the symplesiomorphic feeding condition of Euarthropoda (crown‐group arthropods), were found to have occurred among basal bivalved arthropods.     

Redescription of the Chengjiang arthropod Squamacula clypeata Hou and Bergström, from the Lower Cambrian, south-west China

The anatomy of the arthropod Squamacula clypeata Hou and Bergström, 1997 from the Lower Cambrian Chengjiang Lagersta¨tte is redescribed based on four newly excavated specimens. The new material was collected from localities recently discovered in the Kunming area, Yunnan Province, south-west China, and preserves remarkable details of the ventral morphology, revealed by preparation. Squamacula clypeata is dorsoventrally flattened and rounded in outline. The cephalon was covered by a wide, short shield, with a large doublure and a pair of uniramous antennae on the ventral side. The thorax consists of nine somites, each protected by a tergite and carrying at least one pair of biramous limbs. The pygidium is covered with a small rounded tergum. The endopod is segmented, equipped with short spines on the inner margin of the coxa and a claw-like structure distally, and the exopod flap-like, fringed with setae. The limbs in the pygidium are like those in the thorax in shape. Squamacula was most probably a nektobenthic predator. The spinose endopod could walk, grasp and grind. The large flap-like exopod was adapted for swimming and respiration. Its affinities lie with the Arachnomorpha, but the relationships with other known taxa remain ambiguous.     

REASSESSMENT OF THE GENUS LEANCHOILIA (ARTHROPODA, ARACHNOMORPHA) FROM THE MIDDLE CAMBRIAN BURGESS SHALE, BRITISH COLUMBIA, CANADA

Abstract:  The collection, since 1975, of over 1500 specimens of Leanchoilia Walcott by the Royal Ontario Museum has prompted reassessment of the genus and its species from the Middle Cambrian Burgess Shale. Among new characters in Leanchoilia superlata , the type species, are the presence of two pairs of eyes, a dorsal double carina bracketing the axis of the body segments, segmentation of the gill branch of the appendages, and serration along the body edges from the posterior third of the cephalic shield to the last body segment. Leanchoilia persephone Simonetta, previously synonymized with L. superlata , is also well represented in the Burgess Shale, and is re-established as a valid species, owing to conspicuous differences from the type species. These are the absence of the diagnostic up-curving snout of the cephalic shield, the absence of carina, the shorter 'great appendages', the smooth edges of the body, and its overall shape in dorsal aspect. Leanchoilia superlata and L. persephone may be sexual dimorphs of each other. The ROM collections extend considerably the geographical distribution and stratigraphic range of Leanchoilia in western Canada.     

Class Seticoronaria and Phylogeny of the Phylum Priapulida

The Recent class Seticoronaria (Priapulida) as represented by the genus Maccabeus is compared with the newly described Priapulida from the Middle Cambrian Burgess Shale. It is concluded that the Seticoronaria retained such plesiomorphic characters of the Cambrian Priapulida as sedentary life and food trapping tentacles, whereas the other Recent class, the Priapulomorpha, is the derived, apomorphic branch of this small phylum. The bearing of this hypothesis on an eventual phylogenetic sequence of the Metazoa is discussed.     

Phylogenetic Significance of the Burgess Shale Crustacean Canadaspis

The crustaceans, like the other major living groups of arthropods, have a long evolutionary history. The earliest examples occur in the Cambrian, and fossils of this age are a critical source of evidence of relationships both within the Crustacea, and between the Crustacea and other major arthropod groups. Canadaspis perfecta, from the Middle Cambrian Burgess Shale, is important as one of the oldest well-documented crustaceans. The evidence for reconstructing its remarkable combination of primitive and derived characters is reviewed, and its possible phylogenetic significance re-assessed.     

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.