Anatomy and lifestyle of Kunmingella (Arthropoda, Bradoriida) from the Chengjiang fossil Lagerstätte (lower Cambrian; Southwest China)

An updated reconstruction of the body plan, functional anatomy and life attitude of the bradoriid arthropod Kunmingella is proposed, based on new fossil specimens with preserved soft parts found in the lower Cambrian of Chengjiang and Haikou (Yunnan, SW China) and on previous evidence. The animal has a single pair of short antennae pointing towards the front (a setal pattern indicates a possible sensory function). The following set of seven appendages (each composed of a 5-segmented endopod and a leaf-like exopod fringed with setae) is poorly differentiated, except the first three pairs (with possible rake-like endopodial outgrowths, smaller exopods) and the last pair of appendages (endopod with longer and more slender podomeres). The endopods are interpreted as walking legs with a possible role in handling food particles (marginal outgrowth with setae). The leaf-like exopods may have had a respiratory function. The trunk end is short, pointed, flanked with furcal-like rami and projects beyond the posterior margin of the carapace. The attachment of the body to the exoskeleton is probably cephalic and apparently lacks any well-developed adductor muscle system. The inferred life attitude of Kunmingella (e.g. crawling on the surface of the sediment) was that of a dorsoventrally flattened arthropod capped by a folded dorsal shield (ventral gape at least 120°), thus resembling the living ostracode Manawa. The animal was also probably able to close its carapace as a response to environmental stress or to survive unfavourable conditions (e.g. buried in sediment). The anterior lobes of the valves are likely to have accommodated visual organs (possibly lensless receptors perceiving ambient light through the translucent head shield). Preserved eggs or embryos suggest a possible ventral brood care. The presence of Kunmingella in coprolites and its numerical abundance in Chengjiang sediment indicate that bradoriids constituted an important source of food for larger predators. Kunmingella differs markedly from the representatives of the crown group Crustacea (extant and Cambrian taxa) and from the stem group derivatives of Crustacea (exemplified by phosphatocopids and some ‘Orsten’ taxa) in showing no major sign of limb specialization (e.g. related to feeding strategies). Although it resembles other Chengjiang euarthropods in important aspects of its body plan (e.g. uniramous antennae, endopod/exopod configuration), Kunmingella possesses several features (e.g. antennal morphology, post-antennular appendages with 5-segmented endopods) which support the view that bradoriids may be very early derivatives of the stem line Crustacea.     

Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China)

Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstätten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Luo & Hu, 1999 from the Lower Cambrian Maotianshan Shale biota brings new information concerning the anatomical complexity, functional morphology and lifestyles of the Early Cambrian priapulids. Comparisons are made with Recent priapulids from Sweden (live observations, SEM). The cuspidate pharyngeal teeth of Anningvermis (circumoral pentagons) and the most peculiar radiating oral crown of Corynetis added to the very elongate pharynx of these two forms are interpreted as two different types of grasping apparatus possibly involved in the capture of small prey. Corynetis and Anningvermis are two representative examples of the Early Cambrian endobenthic communities largely dominated by priapulid worms (more than ten species in the Maotianshan Shale biota) and to a much lesser extent by brachiopods. Corynetis and Anningvermis were probably active mud-burrowers and predators of small meiobenthic animals. Likewise predator priapulid worms exploited the interface layer between the seawater and bottom sediment, where meiobenthic organisms were abundant and functioned as prey. This implies that complex prey-predator relationship between communities already existed in the Early Cambrian. This study also shows that the circumoral pentagonal teeth and caudal appendage were present in the early stages of the evolutionary history of the group and were important features of the priapulid body plan already in the Early Cambrian. Two new families, one new genus and new species are introduced and described in the appendix.     

Digestive system and feeding mode in Cambrian naraoiid arthropods

The function of the digestive system of naraoiid arthropods is interpreted in the light of new observations on Early Cambrian specimens from China and detailed comparisons with Recent crustaceans and other arthropods. In naraoiids, paired tubular diverticulae ending as blind caeca are present along the entire midgut, and are interpreted as sites for the secretion of digestive enzymes. Naraoia bears one pair of long, ramifying, distensible diverticulae, possibly used for both food storage and digestion as suggested by Recent analogues (e.g. branchiuran and isopod crustaceans and limulids). Naraoiids were probably epibenthic scavengers/predators rather than mud-eaters. They were either opportunistic intermittent feeders ( Naraoia ) or more regular feeders ( Misszhouia ). The mud-fills of the alimentary canals are likely to be artefacts due to taphonomic and weathering processes or, less likely, to sediment ingestion by animals trapped alive in turbiditic flows. The case study of naraoiid arthropods adds to other fossil evidence supporting the idea that predation played a key role in the Early Cambrian food-webs and that organs adapted for this purpose had already reached a high level of diversity and anatomical sophistication.     

Arthropod visual predators in the early pelagic ecosystem: evidence from the Burgess Shale and Chengjiang biotas

Exceptional fossil specimens with preserved soft parts from the Maotianshan Shale (ca 520 Myr ago) and the Burgess Shale (505 Myr ago) biotas indicate that the worldwide distributed bivalved arthropod Isoxys was probably a non-benthic visual predator. New lines of evidence come from the functional morphology of its powerful prehensile frontal appendages that, combined with large spherical eyes, are thought to have played a key role in the recognition and capture of swimming or epibenthic prey. The swimming and steering of this arthropod was achieved by the beating of multiple setose exopods and a flap-like telson. The appendage morphology of Isoxys indicates possible phylogenetical relationships with the megacheirans, a widespread group of assumed predator arthropods characterized by a pre-oral ‘great appendage’. Evidence from functional morphology and taphonomy suggests that Isoxys was able to migrate through the water column and was possibly exploiting hyperbenthic niches for food. Although certainly not unique, the case of Isoxys supports the idea that off-bottom animal interactions such as predation, associated with complex feeding strategies and behaviours (e.g. vertical migration and hunting) were established by the Early Cambrian. It also suggests that a prototype of a pelagic food chain had already started to build-up at least in the lower levels of the water column.     

Reconsideration of the supposed naraoiid larva from the Early Cambrian Chengjiang Lagerstätte, South China

The Chengjiang Lagerstätte in the Lower Cambrian of South China yields a small, larva‐like arthropod, which was considered to be a protaspis of naraoiids by many authors. The discovery of a large number of well‐preserved specimens from many new localities has allowed the original study to be revised. The relatively large size, stable morphology and unusual structure of the appendages indicate that these specimens represent adults of a new arthropod, Primicaris larvaformis. The larva‐like outline is considered to have arisen by the heterochronic process of progenesis. In addition, this animal displays primitive aspects of bodyplan and limb morphology that suggest a basal position within arachnomorphs, or perhaps even arthropods, and the similarities to the Vendian arthropod‐like animal Parvancorina probably provide an evolutionary link between Vendian forms and Cambrian arthropods.     

Unlocking the early fossil record of the arthropod central nervous system

Extant panarthropods (euarthropods, onychophorans and tardigrades) are hallmarked by stunning morphological and taxonomic diversity, but their central nervous systems (CNS) are relatively conserved. The timing of divergences of the ground pattern CNS organization of the major panarthropod clades has been poorly constrained because of a scarcity of data from their early fossil record. Although the CNS has been documented in three-dimensional detail in insects from Cenozoic ambers, it is widely assumed that these tissues are too prone to decay to withstand other styles of fossilization or geologically older preservation. However, Cambrian Burgess Shale-type compressions have emerged as sources of fossilized brains and nerve cords. CNS in these Cambrian fossils are preserved as carbon films or as iron oxides/hydroxides after pyrite in association with carbon. Experiments with carcasses compacted in fine-grained sediment depict preservation of neural tissue for a more prolonged temporal window than anticipated by decay experiments in other media. CNS and compound eye characters in exceptionally preserved Cambrian fossils predict divergences of the mandibulate and chelicerate ground patterns by Cambrian Stage 3 (ca 518 Ma), a dating that is compatible with molecular estimates for these splits.     

Recent Priapulidae and their Early Cambrian ancestors: comparisons and evolutionary significance

Although priapulid worms form a relatively small phylum in present-day marine environments, they were important animals in Cambrian endobenthic communities. Two Early Cambrian priapulids, namely Xiaoheiqingella peculiaris and Yunnanpriapulus halteroformis nov. gen., nov. sp. from the Maotianshan Shale Lagerstätte of SW China are revised and described. Several key-features of the body plan of Recent Priapulidae are recognized in these two forms: 1) the four-fold body division (introvert, neck, trunk, and caudal appendage); 2) the well-developed introvert armed with ca. 25 longitudinal rows of scalids; 3) the caudal appendage; 4) the pharyngeal teeth arranged in a pentagonal disposition (Xiaoheiqingella); 5) the ventral nerve cord present in Yunnanpriapulus. This morphology indicates close evolutionary relationships with modern priapulids. Xiaoheiqingella and Yunnanpriapulus nov. gen. are tentatively placed within the recent family Priapulidae. The Priapulidae lineage may therefore have a remote origin (Early Cambrian) much older than was previously assumed (Priapulites; Late Carboniferous). The functional morphology of Xiaoheiqingella and Yunnanpriapulus nov. gen. suggests that these two worms were chiefly carnivorous with possible occasional mud-eating habits.     

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.     

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.     

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.     

New occurrence of the family Hipponicharionidae (Bradoriida, Arthropoda), in the lower and middle Cambrian of the Cadenas Ibéricas, Spain

The bradoriids Hipponicharion aff. hispanicum and Wimanicharion aff. matthewi are reported from the lower and middle Cambrian strata of the Cadenas Ibéricas, Spain. The genus Hipponicharion seems to be restricted to the Acadobaltic Province. Wimanicharion has been recorded from Sweden and Canada (Nova Scotia). The new discovery of Wimanicharion in Spain indicates its similar palaeobiogeographical distribution to Hipponicharion.     

The soft anatomy of Isoxys minor from the Guanshan fauna,lower Cambrian of Southwest China

Isoxys is a very common Cambrian bivalved arthropod, specimens of which are normally preserved only as valves. The discovery of the soft anatomy of Isoxys may greatly assist understanding affinities and functional morphology. Isoxys minor Luo and Hu in Luo et al., 2008 is the most common representative of all animal species known from the lower Cambrian Guanshan fauna (Cambrian Series 2, Stage 4) at the Shitangshan Section, near Kunming, Yunnan Province, Southwest China. Here we describe and reconstruct the morphology of I. minor on the basis of newly illustrated fossils and a few new specimens that bear soft-parts including new discovery of frontal grasping appendages. Like the soft anatomy of other known Isoxys, it bears a pair of large stalked eyes, a pair of specialized frontal grasping appendages, approximately 12–14 paired biramous limbs, and a helm-like tail exposed outside the valves.     

昆明海口早寒武世澄江生物群中叶状红藻化石

主要报道了釆自云南省昆明市海口马房村鞍山早寒武世筇竹寺组玉案山段澄江生物群中的叶状红藻-似红叶藻(新属、新种)(Paradelesseria sanguinea Xu,gen.et sp.Nov.)。通过比较形态学研究,探讨了该宏观化石藻类的分类归属与亲缘关系,进一步丰富了澄江生物群生物物种多样性的认识,并为研究早寒武世生物演化及其古环境提供了新的化石证据。同时,通过对现生红藻的比较研究,进一步证明了澄江生物群在云南海口地区发生于一个水深在30m以上的亚潮带和下潮间带的海水环境。     

昆明海口早寒武世澄江生物群中叶状红藻化石

主要报道了采自云南省昆明市海口马房村鞍山早寒武世筇竹寺组玉案山段澄江生物群中的叶状红藻-似红叶藻(新属、新种)(Paradelesseria sanguinea Xu,gen.et sp.Nov.).通过比较形态学研究,探讨了该宏观化石藻类的分类归属与亲缘关系,进一步丰富了澄江生物群生物物种多样性的认识,并为研究早寒武世生物演化及其古环境提供了新的化石证据.同时,通过对现生红藻的比较研究,进一步证明了澄江生物群在云南海口地区发生于一个水深在30 m以上的亚潮带和下潮间带的海水环境.     

Unique lowermost Cambrian small shelly fossils (SSFs) from eastern Yunnan,South China: implications for the early diversification history of animals

With a particular focus on the earliest Cambrian diversification of small shelly fossils (SSFs), stratigraphic analysis was conducted on the lower Cambrian Zhongyicun Member at the Hongjiachong section in the Chengjiang area, Yunnan, China. From ca. 3-m-thick bedded phosphorites (Unit A) in the lower part of the member, we recovered unique SSFs. This interval with 4 SSF-bearing horizons is characterized by the dominance in blade-shaped SSFs, including Halkieria spp., Brushenodus prionodes, Sinosachites delicatus, and Pteronus sp., and the absence of typical tube/cap-shape SSFs common in the Fortunian. This interval tentatively named “Halkieria-dominant interval” is stratigraphically positioned ca. 3 m below the previously known level of the oldest mollusks in the continuous bedded phosphorite sequence between the Anabarites trisulcatus-Protohertzina anabarica Assemblage Zone and the overlying Paragloborilus subglobosa-Purella squamulosa Assemblage Zone of the Fortunian (the earliest Cambrian). The “Halkieria-dominant interval” yields some SSFs likely affiliated with ostracods in view of size and morphology, suggesting that arthropod body fossils appeared in the Fortunian, considerably earlier than previously believed.     

Cambrian anemones with preserved soft tissue from the Chengjiang biota, China

The group Cnidaria includes 'jellyfish', soft-bodied anemone and anemone-like forms and calcified corals. These diploblastic organisms have a fossil record extending back to the earliest metazoans of the Neoproterozoic; however certain cnidarians of the subclass Zoantharia, characterized by soft-bodied anemone-like forms, are absent or poorly represented in the fossil record. Despite the paucity of fossils, it is thought that calcification by soft anemone-like animals was responsible for producing the skeleton that allowed the preservation of the first corals. We report discovery of an abundant assemblage of in situ soft-bodied polyps with tissues. They are preserved in exquisite detail and come from the well-known Lower Cambrian Chengjiang biota of Yunnan, China. The soft-bodied polyps display a simple anatomy that is comparable to some extant anemones of the order Actinaria. The new fossils are assigned to Archisaccophyllia kunmingensis n. gen. et n. sp. Their simple and conservative form suggests that these fossils may represent some kind of ancestral rootstock. The preserved life assemblage provides a unique snapshot of Lower Cambrian anemone life and provides clues for relationships with extant actiniarians as well as calcified corals.     

辽东寒武纪三叶虫Redlichia (Pteroredlichia) murakamii Resser et Endo的蜕壳

华北辽东本溪寒武纪三叶虫Redlichia(Pteroredlichia)murakamii蜕壳标本完好地保存了头盖向下翻转的蜕壳状态,为寒武纪Redlichia的蜕壳模式提供了新的视角。此标本自由颊未保留,胸尾相连,头盖与胸部分离,头盖向下翻转,基本保持原位,为原地埋藏的蜕壳标本。蜕壳过程如下:面线裂开,头盖耸起卷曲并翻转,与胸部分离,胸部扭动,虫体摆脱老壳向前爬出。此蜕壳方式在McNamara(1986)报道的澳大利亚寒武纪Redlichia的蜕壳类型中并未出现。     

Note on the gut preserved in the Lower Cambrian Lingulellotreta (Lingulata, Brachiopoda) from southern China

Lingulellotreta malongensis Rong is the earliest known taxon of the family Lingulellotretidae, which is characterized by the presence of a pedicle foramen as well as an internal pedicle tube. New material from the Early Cambrian Chengjiang Lagerstätte of southern China provides improved anatomical knowledge for lingulellotretid species especially for the digestive system. Additional gut fossils exhibit distinctly the anterior portion composed of esophagus and distended stomach, situated in the alleged visceral cavity, with the recurved intestine accommodated inside a hollow pseudointerarea. The frequency of occurrence of this intestinal layout suggests that this is not just an artefact of preservation. The gross configuration of the guts and the way they are preserved in the fossils suggest that they are in situ and, therefore, we can assume that Lingulellotreta had a hollow cavity presumably subtended by the pseuodelthyrium, which was invaded and occupied by the visceral organs. Hence, these fossils demonstrate the dangers of extrapolating crown‐group soft‐tissue configuration to the stem group.     

Morpho-anatomy of the lobopod Magadictyon cf. haikouensis from the Early Cambrian Chengjiang Lagerstätte, South China

Magadictyon haikouensis (Luo and Hu, 1999) from the Early Cambrian Chengjiang Lagerstätte, an incomplete specimen of a large lobopod with strong appendages, has been regarded as related to the lobopods Microdictyon and Onychodictyon. Newly discovered complete specimens of Magadictyon cf. haikouensis (found by the Early Life Institute field team) show that the taxon, in addition to its strong appendages with appendicules, also had a head bearing similar caecum‐like structures to those of the arthropod Naraoia and Chelicerate, ‘Peytoia’‐like mouthparts and frontal appendages. Because of their similarity, the caecum‐like structures of Magadictyon cf. haikouensis are considered to be homologous with those of stem‐group arthropods. The ‘Peytoia’‐like mouthparts and the frontal appendages are similar to those of the AOPK (Anomalocaris–Opabinia–Pambdelurion–Kerygmachela) group. In addition, the appendages with appendicules show that Magadictyon cf. haikouensis is closely related to Onychodictyon. Therefore, Magadictyon cf. haikouensis is regarded here as a rare transitional form between lobopods and arthropods. Besides, together with other lobopods, the morphology of Magadictyon cf. haikouensis demonstrates that the Cambrian lobopods appear to have been diverse and not particularly closely related to one another, and do not seem to represent a monophyletic clade.     

Early Cambrian Salterella and Volborthella (Phylum Agmata) re-evaluated

The genus Volborthella is removed from synonymy under Salterella and placed in the extinct phylum Agmata. Volborthella is more restricted geographically and occurs in older Lower Cambrian strata than Salterella, but distinguishing younger specimens of that genus from poorly preserved Salterella is difficult. Vologinella is not related to Volborthella or other Agmata. Other genera assigned to the phylum by various workers are discussed, but their assignment to this phylum is either enigmatic or rejected. The published concept that specimens of Volborthella and Salterella are agglutinated sclerites or spines on a larger soft-bodied organism is discussed. The material on which this notion is based is re-illustrated and the notion rejected in favor of the view that these conical forms are the hard parts of discrete organisms.