First record of a bivalved larval shell in Early Cambrian tommotiids and its phylogenetic significance

Abstract: Brachiopods are marine Lophotrochozoa whose soft parts are enclosed in a bivalved shell. Although brachiopods are represented by a rich record from the Early Cambrian to the present, the origin of their bivalved body plan remains controversial. The Early Cambrian organophosphatic tommotiids Micrina and Paterimitra from Australia have been proposed as stem brachiopods. Here, we describe their earliest ontogeny, indicating that tommotiids possessed bivalved planktotrophic larvae. The curious combinations of characters in Micrina and Paterimitra indicate that they may belong to the stems of the Linguliformea and Rhynchonelliformea, respectively. The bivalved shell of adult living brachiopods may represent a plesiomorphic character retained from planktic tommotiid larvae; the crown group body plan of the Brachiopoda may have evolved through the paedomorphic retention of a bivalved larval state.     

Shell Structure And Inferred Growth, Functions And Affinities Of The Sclerites Of The Problematic Micrina

The stratiform laminae of Micrina sclerites originally consisted of rheomorphic successions of monolayers of micrometric–sized, apatitic tablets, presumably interleaved with chitin and glycosaminoglycans (GAGs). Paired laminae enclose slot–like chambers swelling into lobes distally that originally contained GAGs and deposits of spherulitic and prismatic apatite. The laminae are pervaded by apatitic tubes, apparently secreted by microvillous setoblasts and containing, at the surface, chitinous setae. Internal markings suggest that the triangular (sellate) sclerite supported a pair of muscles and the planospiral (mitral) sclerite, a medial muscle and gonadal sacs flanked by a pair of crescentic muscle bases. Both sclerites were secreted by a mantle with a circumferential fold. The sellate and mitral sclerites are homologized with the anterior and posterior shells of Halkieria and could have become the dorsal and ventral valves of the ancestral brachiopod by a sequence of transformations. These include: the folding of the halkieriid body axis; accelerated mixoperipheral growth of the anterior (dorsal) shell to enclose, with the posterior (ventral) shell, a mantle cavity lined with modified ciliated epithelium of the foot; reduction of sclerite–secreting epithelium to the locus of the brachiopod pedicle epithelium; and the anterior (dorsal) spread of gonadal lamellae.     

Paterimitra pyramidalis from South Australia: scleritome,shell structure and evolution of a lower Cambrian stem group brachiopod

The tommotiid Paterimitra pyramidalis Laurie, 1986, is redescribed based on well‐preserved material from the lower Cambrian Wilkawillina, Wirrapowie and Ajax limestones of the Flinders Ranges, South Australia. The material shows that the scleritome of Paterimitra pyramidalis includes three sclerite morphotypes (S1, S2 and L). Detailed shell microstructure studies show striking similarities with both the paterinid brachiopod Askepasma toddense and the tommotiid Eccentrotheca helenia, which strengthens the suggested evolutionary link between tommotiids and brachiopods. Based on the partly articulated specimens and similarities in shell microstructure and sclerite morphology with Eccentrotheca, Paterimitra pyramidalis is reconstructed as a tube‐dwelling, epifaunal, sessile, filter‐feeder with an organic pedicle‐like attachment structure. The proposed reconstruction of the scleritome comprises a basal unit composed of one S1 and one S2 sclerite, as well as an unresolved number of L sclerites lining a coniform tubular structure.     

A Stem Group Brachiopod From The Lower Cambrian: Support For A Micrina (Halkieriid) Ancestry

The shell structure of the Lower Cambrian Mickwitzia , a bilaterally symmetrical bivalve hitherto doubtfully assigned to the Brachiopoda, confirms that the genus shares characters with linguliform brachiopods. The columnar lamination of its organophosphatic shell is homologous with that characterizing acrotretides. The shell, however, is also pervaded by striated apatitic tubes indistinguishable from those permeating the sclerites of the problematic organophosphatic, laminar–shelled Micrina which is close to Halkieria . No crown group brachiopods have such tubes that are presumed to have contained setae. The presence of both these features in the Mickwitzia shell suggests that the stock is a stem group brachiopod with a halkieriid ancestry.     

Tommotiids from the early Cambrian (Series 2, Stage 3) of Morocco and the evolution of the tannuolinid scleritome and setigerous shell structures in stem group brachiopods

An assemblage of tannuolinid sclerites is described from the Amouslek Formation (Souss Basin) of the Anti‐Atlas Mountains in Morocco. The assemblage contains two species, Tannuolina maroccana n. sp., which is represented by a small number of mitral and sellate sclerites, and Micrina sp., represented by a single mitral sclerite. Tannuolina maroccana differs from other species of the genus in the presence of both bilaterally symmetrical and strongly asymmetrical sellate sclerites. This observation suggests that the scleritome of Tannuolina was more complex than previously thought and that this tommotiid may have held a more basal position in the brachiopod stem group than previously assumed. The shell structure of both T. maroccana and Micrina sp. is well preserved and exhibits two fundamentally different sets of tubular structures, only one of which was likely to contain shell‐penetrating setae. Based on these observations, the structure of the tannuolinid shell is discussed and its implications for the evolution of tubular microstructures in stem and crown group brachiopods are analysed.     

The tommotiid Kelanella and associated fauna from the early Cambrian of southern Montagne Noire (France): implications for camenellan phylogeny

A carbonate bed of the Pardailhan Formation, early Cambrian, southern Montagne Noire (southern France), provided microfossils such as Hyolithellus sp., Torellella cf. mutila and Torellella sp. along with numerous disarticulated pieces of composite skeletons such as valves of the brachiopod Eoobolus priscus and of the bradoriid Monceretia erisylvia, and chancelloriid sclerites (Chancelloria sp.). The assemblage also furnished a rich set of sclerites from the tommotiid Kelanella altaica. Five morphological variations of the latter have been identified. The presence of concentric ribs formed by distal inflation of selected shell laminae in Kelanella supports its assignment to the camenellans. More particularly, the presence of transverse structures within the internal cavity (septa) of Kelanella suggests a close relationship with the Lapworthellidae. However, the latter differ from Kelanella by the continuous morphological variation along their scleritome which is also composed of simple conical elements with uniform ornamentation. Several forms of Kelanella are similar to mitral and sellate sclerites of Camenella, whereas some other forms are comparable to Kennardia. The new material suggests that Kelanella occupies a transitional position between Lapworthellidae and the grouping of Tommotiidae and Kennardiidae. Such a phylogenetic position also implies that the number of sclerite morphotypes tends to decrease within the camenellan scleritome during evolution.     

云南永善肖滩剖面早寒武世Lapworthella (托莫特壳类)骨片化石

描述了采自云南永善肖滩剖面下寒武统石岩头组的托莫特壳类(tommotiids)骨片化石Lnpworthella rete Yue,1987,其骨片锥状或塔状,横向环脊明显,环脊尖棱状;骨片外表面具多边形网状纹饰,但无齿、瘤或纵脊等装饰;骨片可分为两种类型,即弯曲型和近直立型,前者数量明显多于后者。简要地讨论了托莫特壳类化石的分类及生物亲缘关系,认为目前为止,其仍为亲缘关系不明的化石类别。尚难断定所有的托莫特壳类骨片化石皆属于同一生物门类,因为难以证明将它们归人同类生物所依据的骨片性状是共同衍征(synapomorphy)还是由于趋同演化(convergent evolution)的结果;在缺乏躯干化石记录的情况下,离散骨片的生物亲缘关系研究较为困难。     

A new mongolian species of the lower cambrian genus camenella and the problems of scleritome-based taxonomy of the tommotiidae

The tommotiidCamenella parilobata n. sp. is described on the basis of a collection of mitral and sellate sclerites from the Lower Cambrian Khairkhan »Formation« of western Mongolia. The detailed documentation of co-occurrence of sclerite types is essential for the taxonomic analysis of the problematic Tommotiidae and related groups; most »tommotiid« (sensu lato) taxa are still known only from a few isolated sclerites representing an unknown spectrum of variability. Until a detailed analysis of the scleritome composition and sclerite microstructure in representatives of all genera has been made, any suprageneric classification can only be speculative.     

Molecular phylogenetic evidence supports a new family of octocorals and a new genus of Alcyoniidae (Octocorallia,Alcyonacea)

Molecular phylogenetic evidence indicates that the octocoral family Alcyoniidae is highly polyphyletic, with genera distributed across Octocorallia in more than 10 separate clades. Most alcyoniid taxa belong to the large and poorly resolved Holaxonia–Alcyoniina clade of octocorals, but members of at least four genera of Alcyoniidae fall outside of that group. As a first step towards revision of the family, we describe a new genus, Parasphaerasclera gen. n., and family, Parasphaerascleridae fam. n., of Alcyonacea to accommodate species of Eleutherobia Pütter, 1900 and Alcyonium Linnaeus, 1758 that have digitiform to digitate or lobate growth forms, completely lack sclerites in the polyps, and have radiates or spheroidal sclerites in the colony surface and interior. Parasphaerascleridae fam. n. constitutes a well-supported clade that is phylogenetically distinct from all other octocoral taxa. We also describe a new genus of Alcyoniidae, Sphaerasclera gen. n., for a species of Eleutherobia with a unique capitate growth form. Sphaerasclera gen. n. is a member of the Anthomastus–Corallium clade of octocorals, but is morphologically and genetically distinct from Anthomastus Verrill, 1878 and Paraminabea Williams & Alderslade, 1999, two similar but dimorphic genera of Alcyoniidae that are its sister taxa. In addition, we have re-assigned two species of Eleutherobia that have clavate to capitate growth forms, polyp sclerites arranged to form a collaret and points, and spindles in the colony interior to Alcyonium, a move that is supported by both morphological and molecular phylogenetic evidence.     

A starfish with three-dimensionally preserved soft parts from the Silurian of England

Palaeozoic asteroids represent a stem-group to the monophyletic post-Palaeozoic Neoasteroidea, but many aspects of their anatomy are poorly known. Using serial grinding and computer reconstruction, we describe fully articulated Silurian (ca 425 Myr) specimens from the Herefordshire Lagerst?tte, preserved in three dimensions complete with soft tissues. The material belongs to a species of Bdellacoma, a genus previously assigned to the ophiuroids, but has characters that suggest an asteroid affinity. These include a pyloric system in the gut, and the presence of large bivalved pedicellariae, the latter originally described under the name Bursulella from isolated valves. Ampullae are external and occur within podial basins; the radial canal is also external. Podia are elongate and lack terminal suckers. The peristome is large relative to the mouth. Aspects of the morphology are comparable to that of the extant Paxillosida, supporting phylogenetic schemes that place this order at the base of the asteroid crown group.     

Scleritome construction,biofacies, biostratigraphy and systematics of the tommotiid Eccentrotheca helenia sp. nov. from the Early Cambrian of South Australia

Abstract: Large collections of Eccentrotheca helenia sp. nov. from the lower Cambrian Wilkawillina and Ajax limestones in the Arrowie Basin, South Australia, contain abundant low, cap‐shaped and high, laterally compressed isolated sclerites in addition to partially articulated tubular specimens. The scleritome of Eccentrotheca helenia sp. nov. is fully described for the first time and shown to be formed by ontogenetic fusion of sclerites into successively stacked sclerite rings, forming a larger, tubular structure. The apical termination of the tube is highly variable, but is primarily constructed by low, cap‐shaped sclerites and characterised by a central aperture of variable inclination. The adapical portion of the tube is predominantly constructed by high, laterally compressed sclerites, but individual sclerite rings can contain both cap‐shaped and laterally compressed sclerites along with sclerites of intermediate morphology. The apical aperture presumably housed organic structures for attachment to a hard substrate, but the scleritome also occasionally preserves small lateral perforations between fused sclerites, which may have served to stabilise the scleritome by providing additional points of anchorage. In the Arrowie Basin, E. helenia is found in association with archaeocyath‐microbial‐spongiomorph‐dominated bioherms and most likely inhabited pendant or cryptic habitats within these bioherms. Eccentrotheca‐like sclerites form an integral part of the scleritomes of many tommotiids which may confuse taxonomic analysis. Sclerites previously assigned to ‘E.’guano, consistently occur together with sclerites of Kulparina rostrata in stratigraphic intervals consistently older than strata hosting E. helenia. Rare fused specimens indicate that the sclerites of K. rostrata and ‘E.’guano belong to the same scleritome.     

Halkieria系统进化及其化石分类学研究进展

Halkieria是寒武纪一类躯体上披有骨质鳞片的疑难后生动物,前后各有一个大壳板,两侧对称,其钙质骨片在软躯体上排列形成几条规律的纵向带。半个世纪以来,学术界对Halkieria生物系统确切位置的讨论以及Halkieriidae不同属种亲缘关系的探索从未停止。本文总结了Halkieria化石在骨片属种分类、古地理和地层分布、生物系统分类学的研究进展,对当前的生物系统分类归纳出以下几种观点:1) Halkieria为腕足动物的祖先,并与托莫特壳类有一定的亲缘关系;2)Halkieria与Wiwaxia等骨片化石具有单源性,组成一种新的分类单元Halwaxiidae,属于软体动物干群或腕足动物和环节动物的干群;3)Halkieria为软体动物有刺亚门干群,与无板纲、多板纲更为亲近; 4) Halkieria为环节动物的祖先。目前,虽然多数人根据Halkieria与现生多板纲的骨片排列一致将Halkieria归属软体动物有刺亚门的干群,但近年来世界各地新化石材料的陆续发现及壳体发育研究使得Halkieria的系统分类位置出现新的争议。通过对化石分类学的系统厘定和生物分类研究进展的总结梳理,...     

A new long-necked ‘sauropod-mimic’ stegosaur and the evolution of the plated dinosaurs

Stegosaurian dinosaurs have a quadrupedal stance, short forelimbs, short necks, and are generally considered to be low browsers. A new stegosaur, Miragaia longicollum gen. et sp. nov., from the Late Jurassic of Portugal, has a neck comprising at least 17 cervical vertebrae. This is eight additional cervical vertebrae when compared with the ancestral condition seen in basal ornithischians such as Scutellosaurus. Miragaia has a higher cervical count than most of the iconically long-necked sauropod dinosaurs. Long neck length has been achieved by ‘cervicalization’ of anterior dorsal vertebrae and probable lengthening of centra. All these anatomical features are evolutionarily convergent with those exhibited in the necks of sauropod dinosaurs. Miragaia longicollum is based upon a partial articulated skeleton, and includes the only known cranial remains from any European stegosaur. A well-resolved phylogeny supports a new clade that unites Miragaia and Dacentrurus as the sister group to Stegosaurus; this new topology challenges the common view of Dacentrurus as a basal stegosaur.     

寒武纪叶足动物链状心网虫躯干骨板功能形态学研究*

链状心网虫(Cardiodictyon catenulum)是寒武纪澄江生物群中较为常见的叶足动物, 其躯干骨板的形态一直存在较大争议。本项研究显示该叶足动物的躯干具有一纵列马鞍状骨板, 每个体节具有单一的骨板, 每个骨板从背部延伸至躯干两侧。因此该叶足动物的骨板与其余具有成对或环状分布的躯干骨板的叶足动物存在显著的区别。骨板中央存在一个圆锥形凸起, 骨板边缘显著增厚且呈锯齿状。链状心网虫的躯干骨板边缘增厚和鞍点的凸起提升了整体的刚性; 锯齿状边缘也指示了更优的保护及防御功能。本项研究还发现链状心网虫骨板表面具有显微筛状结构, 筛孔被充填并保存为众多等距离分布的黄铁矿化瘤突, 与微网虫、怪诞虫及爪网虫躯干骨板的显微网状结构类似, 可能为纤细乳突的出口。本文认为: 寒武纪叶足动物的躯干骨板普遍具有显微筛状结构; 这些骨板除了保护与防御, 可能还有感觉的功能。每个躯干体节对应骨板为寒武纪叶足动物的独有衍征, 可能为背骨板的早期形态, 也可能由成对骨板愈合而来。     

湖北房县下寒武统西蒿坪段钉形骨片化石

本文对采自湖北房县三座庵剖面下寒武统西蒿坪段的钉形骨片(nail-shaped)化石进行了研究,系统厘定和描述了2属2种,分别为Parazhijinites guizhouensis和Cambroclavus angxianensis;其中以C.fangxianensis骨片化石最为丰富,由三种类型的骨片组成,它们可能...     

Phylogenetic reconsideration of Myrmekiaphila systematics with a description of the new trapdoor spider species Myrmekiaphila tigris (Araneae, Mygalomorphae, Cyrtaucheniidae, Euctenizinae) from Auburn, Alabama

The trapdoor spider genus Myrmekiaphila currently comprises 11 nominal species. A recent molecular phylogenetic evaluation of the group identified a number of problems with respect to how species and species groups were delineated by Bond and Platnick in their 2007 taxonomic revision of the genus. We report herein the discovery of a new species, Myrmekiaphila tigrissp. n. The phylogenetic position of the species is evaluated using a molecular phylogenetic approach based on a set of mtDNA markers. Our preferred phylogenetic hypothesis supports the recognition of a new species and further highlights the need to more carefully investigate species boundaries within the genus. These results further indicate that palpal bulb morphology is rapidly evolving and has likely been a contributing factor in rendering a number of species paraphyletic with respect to the molecular data.     

Two new species of gorgonian octocorals from the Tropical Eastern Pacific Biogeographic Region (Cnidaria,Anthozoa, Gorgoniidae)

The gorgoniid Eugorgia is exclusively an eastern Pacific genus. It has a wide geographic and bathymetric range of distribution, found from California to Perú and extends down to 65 m deep. Two new species are herein described. The morphological characters were analyzed and illustrated by light and scanning electron microscopy. Eugorgia beebei sp. n. can be distinguished by its white, ascending, sparse colony growth. Eugorgia mutabilis sp. n. can be distinguished by its white colony that changes color after collection, and the conspicuous sharp-crested disc sclerites. From a morphological point of view the new species are related to the daniana-group, the rubens-group and the siedenburgae-group of Eugorgia; their affiliations, and the proposal of a new group are discussed. These new species increases the number of species in the genus to 15, and contribute to the knowledge of the eastern Pacific octocoral biodiversity.