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.     

Cambrian lobopodians–ancestors of extant onychophorans?

This paper is a review of published information on fossil lobopodians, with addition of observations and ideas based on new material. It is also an analysis of the phytogeny of the group, and presents a new classification. A character shared by three or four families, yet not seen before, is a pair of enlarged sclerites covering the head. This forms an argument for re-orienting Hallucigenia once again. This genus no longer being enigmatic, a corner-stone in Stephen Jay Gould's evidence for extinct phyla is therefore gone. It is suggested that the lobopodians, phylum Lobopodia, are arranged in two classes, the extinct Xenusia for marine forms, and the Onychophora for terrestrial forms. The marine lobopodians are morphologically much more diverse than the extant onychophorans, a condition expressed in the classification. New taxa are: Hallucigenia fortis sp. nov., the families Luolishaniidae, Cardiodictyidae and Onychodictyidae, and the new orders Archonychophora, Scleronychophora and Paronychophora.     

A new basal ornithuromorph bird (Aves: Ornithothoraces) from the Early Cretaceous of China with implication for morphology of early Ornithuromorpha

Ornithuromorpha is the most derived avian group in the Early Cretaceous, advanced members of which encompass all living birds (Neornithes). Here we report on a new basal ornithuromorph bird, Bellulia rectusunguis gen. et sp. nov., represented by a nearly complete skeleton from the Early Cretaceous Jehol Biota in northeastern China. A comprehensive phylogenetic analysis resolved the new taxon in a basal position that is only more derived than Archaeorhynchus and Jianchangornis among ornithuromorphs, increasing the morphological diversity of basal ornithuromorphs. The new specimen has a V‐shaped furcula with a short hypocleidium, a feature otherwise known only in Schizooura among Cretaceous ornithuromorphs. We discuss the implications of the new taxon on the evolution of morphology of primitive ornithuromorphs, particularly of pectoral girdle, sternum and limb proportion pertaining to powered flight. The preserved gastroliths and pedal morphology indicate herbivory and lakeshore adaption for this new species. © 2015 The Linnean Society of London     

Middle Triassic conodont apparatus architecture revealed by synchrotron X-ray microtomography

The composition of conodont apparatuses is crucial for understanding the feeding mechanisms of these early vertebrates. However, the multielement apparatus reconstructions of most species remain equivocal because they have been inferred from loose element collections, guided by knowledge from rare articulated ‘bedding plane assemblages’ and fused clusters, often from distantly related taxa. Even these natural assemblages can be difficult to interpret because the component elements can be closely juxtaposed or embedded in matrix, making it hard to discern the morphology of each element and their relative positions within the architecture of the feeding apparatus. Here we report five exceptionally preserved conodont clusters from the Middle Triassic Luoping Biota, Yunnan Province, Southwest China. These materials were scanned using synchrotron radiation X-ray tomographic microscopy (SRXTM), revealing the morphology and positional homology of the component elements in the fused clusters. We confirm that the apparatus of Nicoraella was composed of eight types of elements, comprising a total of 15 elements. SRXTM reveals the positional homologies of the component elements, viz. a single alate element is located in the S₀ position, flanked successively abaxially by pairs of breviform digyrate S₁ and S₂ elements, bipennate S₃ and S₄ elements, and a pair of inwardly curved breviform digyrate M elements. Carminate elements occupy the P₁ and P₂ positions. The apparatus of Nicoraella is among the most completely characterised of all conodonts and serves as a template for the reconstruction of gondollellids.     

New data on the palaeobiology of the enigmatic yunnanozoans from the Chengjiang Biota,Lower Cambrian,China

Interpretation of the enigmatic soft‐bodied yunnanozoans from the Lower Cambrian (Stage 3, Series 2) Chengjiang Biota has remained controversial for decades because of their strange body plan and their variable taphonomic alteration. Proposed affinities have ranged from stem bilaterian to stem vertebrate. A study of over seven hundred slabs, many newly collected, from five sections at two localities demonstrates that yunnanozoans have sclerotized dorsal and axial segments, a body cavity, a unique feeding region and a coiled alimentary canal. Although two genera have been previously discriminated, all specimens have the same number of filamentous arches (seven pairs) and of ventral circular structures (four pairs); this indicates that Haikouella Chen, Huang and Li, 1999 is a junior synonym of Yunnanozoon Hou, Ramsköld and Bergström, 1991. Our analysis reveals new details of the putative pharyngeal pores of Yunnanozoon lividum, and although there are similarities with the gill slits of deuterostomes, the question of their homology remains moot. The filamentous arches and their supporting rods were probably sclerotized and represent a specialized feeding structure. Yunnanozoans exhibit characters that are open to a number of alternative interpretations of putative homology, encompassing comparisons with deuterostomes and protostomes (including members of the Platyzoa). Given their unique features, and the current lack of secure homologies, the affinities of yunnanozoans should continue to be considered in a wider bilaterian context.     

The origins and key innovations of vertebrates and arthropods

The Lower Cambrian Maotianshan Shale near Kunming (Southern China) not only retains beautifully preserved and diverse organisms but also documents a missing evolutionary history between living vertebrates and their amphioxus-like ancestor. Presented here are the key novelties both for the evolutionary origins of the chordates and for the evolutionary transition from amphioxus-like ancestor toward the vertebrates. The adaptation for a burrowing life style is considered a key adaptive pressure for generating novel chordate-only anatomical characters including the first axial skeleton, e.g., notochord and myotomes. The transition from amphioxus-like ancestor toward the vertebrates was presumably triggered by the way toward more active life style, resulted in the origination of numerous novel structures including neural crests, a more complex head with upper and lower lips, an active gilled pharyngeal system, a large brain, image-forming paired eyes, and a bony axial skeleton (vertebrae). The diverse limb-bearing organisms from the Lower Cambrian Maotianshan Shale arthropods representing different evolutionary stages shed light on the mysterious field of the evolutionary origins of the arthropods and reveal a grand scenario of how the arthropods paved their way through step-wise evolution from worm-like ancestor toward living crown-lineage arthropods.     

Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

Liaoxia Cao et S.Q. Wu (Gnetales): ephedroids from the Early Cretaceous Yixian Formation in Liaoning, northeastern China

Gnetalean compression-impression fossils are described from the Early Cretaceous Yixian Formation, Liaoning Province, north-eastern China, and assigned to six species of Liaoxia Cao et S.Q. Wu. The fossils have opposite-decussate phyllotaxis and cones comprising 2–12 pairs of bracts. Ovulate cones have seeds typically in a distal position. The species differ from each other and from previously described fossils in the absence or presence of leaves, shape of cones and seeds, and shape and position of cone bracts. The species of Liaoxia are probably close relatives of extant species of Ephedra L., but diagnostic reproductive details that could confirm this hypothesis are not preserved. The restricted information in the fossils and the poorly understood morphological diversity of extant Ephedra, prevent assignment of the fossils to any particular subgroup of Ephedra, as well as an explicit exclusion of them from the extant genus.