RECONSIDERATION OF THE PHYLOGENETIC AFFINITIES OF ZHIJINITIDS

Isolated sclerites are an important component of the Early Cambrian small shelly fos-sils(SSF) ,accounting for one-third of both the total number of taxa and the total numberof specimens.Estimating the phylogenetic affinities of these sclerites is one of the mostdifficult problems in the analysis of the SSF.Sclerites referred to such taxa as the halkieri-ids,siphogonuchitids,chancelloriids,tommotiids,zhijinitids and cambroclavids generallyhave been interpreted as Problematica.As recently a…     

New Cambrian lobopods and chaetognaths of the Siberian Platform

The association of isolated remains of lobopods with a netlike structure, including plates of Microdictyon rozanovi sp. nov. and sclerites with spinelike process of Onychomicrodictyon spiniferum gen. et sp. nov., as well as saberlike sclerites of Protohertzina yudomica sp. nov. interpreted as grasping spines of chaetognaths, is described from the middle part of the Inikan Formation of the Toyonian Stage of the Lower Cambrian (Lermontovia dzevanovskii Zone). The diagnoses and species composition of the genera Microdictyon and Protohertzina are refined.     

贵州剑河寒武系“清虚洞组”管状微体骨骼化石初步研究

华南寒武系地层中广泛富集微体骨骼化石,为解决某些疑难化石的亲缘关系及研究早期后生动物的演化提供了重要化石证据。在贵州剑河八郎"清虚洞组"中发现一些管状微体骨骼化石。经鉴定后主要有4属,分别为小钻孔螺Torellella、似软舌螺Hyolithellus、鞘状螺Coleoloides和表面具鳞片状管状化石Mongolitubulus squamifer。Mongolitubulus分布范围较为广泛,本文结合前人对该化石亲缘关系的探讨及剑河寒武系"清虚洞组"化石的特征,推测M.squamifer可能是高肌虫的装饰刺。     

A comparative study of Lower Cambrian Halkieria and Middle Cambrian Wiwaxia

Two Cambrian lepidote metazoans known from different respective types of preservation have been compared in order to elucidate their biology and affinities. The widely distributed Lower Cambrian Halkieria is represented by isolated hollow sclerites, probably of originally calcareous composition. The Middle Cambrian Wiwaxia is known from the Burgess Shale as isolated sclerites (scales and spines) and as more or less complete individuals. Although Halkieria sclerites were mineralized and those of Wiwaxia were probably not, there are fundamental structural and morphological similarities between the two. Both bad an imbricating scaly and spiny armour consisting of hollow sclerites with a longitudinally fibrous structure. The sclerites did not grow, but were probably moulted during the course of ontogenetic growth. Halkieria and Wiwaxia are regarded as closely related. Both are referred to the Order Sachitida He 1980. The sclerite armour of Halkieria is reconstructed on the template provided by Wiwaxia. The interpretation of sachitid sclerites as protective armour is an alternative to the interpretation by Jell (1981, Alcheringa 5 )that sachitid sclerites were respiratory organs in an animal of probable annelid affinities. Sachitids are interpreted as sluggish, benthic deposit feeders that do not belong to any recognized phylum.     

<Emphasis Type="Italic">Diplectanocotyla</Emphasis> Yamaguti, 1953 (Monogenea: Diplectanoidea) from <Emphasis Type="Italic">Megalops cyprinoides</Emphasis> (Broussonet) (Teleostei: Megalopidae) off Peninsular Malaysia

Two known and two new species of Diplectanocotyla Yamaguti, 1953 (D. gracilis Yamaguti, 1953, D. megalopis Rakotofiringa & Oliver, 1987, D. langkawiensis n. sp. and D. parva n. sp.) were collected from Megalops cyprinoides (Megalopidae) off Langkawi, Kedah and Matang, Perak, Peninsular Malaysia. All four species possess similar types of sclerotised male and female reproductive structures and similar soft anatomical features. The squamodisc sclerites of all four species have spine-like projections with varying degrees of visibility and shapes (sharp-pointed to triangular). In D. megalopis and D. langkawiensis n. sp. the spines are sharp-pointed and distinct on sclerites from rows 5–6 onwards. In D. gracilis and D. parva n. sp. the sclerite spines are triangular, lightly sclerotised and occur on almost all of the sclerites. D. parva n. sp. has comparatively the smallest set of anchors, bars, squamodiscs and squamodisc suckers. The anchors and bars of the other three species are almost similar in overall size, and the main distinguishing feature is the relative lengths of the inner and outer roots of the ventral anchors. In D. gracilis the outer root is very much smaller than the inner root and they are disposed almost at a right angle to each other. In D. megalopis the outer root is usually about half the length of the inner root and the roots are inclined at c.60° to each other. In D. langkawiensis n. sp. the roots are inclined at c.40° degrees and the outer root is of a similar length or only slightly shorter than the inner root. The openings of the two squamodisc suckers of all four Diplectanocotyla species are surrounded by tiny scale-like spines. Bifid tegumental spines are found in the posterior region of all four species, differing only in their extent: in D. parva n. sp. the tegumental spines are only distributed in the peduncular region and not beyond, whilst in the other three species the tegumental spines extend from the posterior level of the testis to the end of the peduncle. An amended diagnosis of Diplectanocotyla and a key to its species are appended.     

Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian ‘explosion’. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution—but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils—including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum—as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the ‘middle’ Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.     

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。     

DEVELOPMENT AND DIVERSIFICATION OF TRUNK PLATES OF THE LOWER CAMBRIAN LOBOPODIANS

Abstract:  Isolated lobopodian plates are reported from Early Cambrian strata at five localities in southern China. A wide variety of morphologies is represented, reflecting a considerable diversification within the phylum at this time. The new taxon Microdictyon jinshaense is erected and new observations are recorded on established taxa, based on examination of more than 600 well-preserved plates; irregular patterns of node distribution and the presence of large spines are documented on some taxa for the first time. Rare specimens in which two plates are conjoined, with a larger plate underlying a smaller one, are interpreted as showing a new sclerite emerging underneath its predecessor, which has not yet been moulted. These specimens confirm the process of ecdysis in the lobopodians and contribute to a fuller understanding of the ontogeny of these organisms. A functional hypothesis that suggests that the plates were complex visual structures is refuted; it is possible that they were sites of muscle attachment, but a protective role is regarded as more plausible.     

New species of Tereancistrum Kritsky,Thatcher & Kayton, 1980 (Monogenea: Dactylogyridae: Ancyrocephalinae) from the gills of Prochilodus lineatus (Osteichthyes: Prochilodontidae) from the upper Paraná River floodplain,Brazil

Two new species of Tereancistrum Kritsky, Thatcher & Kayton, 1980 are described from the gills of Prochilodus lineatus (Prochilodontidae) collected from the upper Paraná River floodplain, Brazil. T. toksonum n. sp. is characterised by a slim ventral bar with a sclerotised membrane along the anterior margin. Another characteristic is the dorsal anchor with an elongated deep root. T. curimba n. sp. is similar to T. ornatus Kritsky, Thatcher & Kayton, 1980 but possesses a sclerotised structure between the accessory sclerites of the ventral anchors. Moreover, the accessory anchor sclerites of this new species are longer than those described for T. ornatus.     

Acanthotrema felis n. sp. (Digenea: Heterophyidae) from the small intestine of stray cats in the Republic of Korea

Acanthotrema felis n. sp. (Digenea: Heterophyidae) is described from adult flukes recovered in the small intestine of stray cats caught in the southwestern part of the Republic of Korea. The worms were characterized by minute body size, the presence of a bipartite seminal vesicle, and a ventral sucker associated with a ventrogenital sac, which enclosed 3 sclerites (2 long and pointed, and I short and thumblike), and an unarmed gonotyl. They were assigned to Acanthotrema Travassos, 1928, which currently contains 4 species. The new species resembles A. acanthotrema, the type species, because both have 3 sclerites on the ventrogenital sac. However, all 3 sclerites in the new species are armed with minute spines on their base, whereas only 2 sclerites in A. acanthotrema have minute spines near their tips. Three species of Stictodora Looss, 1899, namely,Stictodora tridactyla Martin and Kuntz, 1955, S. cursitans Kinsella and Heard, 1974, and S. tanayensis Velasquez, 1973, are transferred to Acanthotrema, and the new species is distinguished from them particularly in terms of the number and shape of its sclerites. The generic diagnosis of Acanthotrema is emended.     

A cryptic record of Burgess Shale‐type diversity from the early Cambrian of Baltica

Exceptionally preserved ‘Burgess Shale‐type’ fossil assemblages from the Cambrian of Laurentia, South China and Australia record a diverse array of non‐biomineralizing organisms. During this time, the palaeocontinent Baltica was geographically isolated from these regions, and is conspicuously lacking in terms of comparable accessible early Cambrian Lagerstätten. Here we report a diverse assemblage of small carbonaceous fossils (SCFs) from the early Cambrian (Stage 4) File Haidar Formation of southeast Sweden and surrounding areas of the Baltoscandian Basin, including exceptionally preserved remains of Burgess Shale‐type metazoans and other organisms. Recovered SCFs include taxonomically resolvable ecdysozoan elements (priapulid and palaeoscolecid worms), lophotrochozoan elements (annelid chaetae and wiwaxiid sclerites), as well as ‘protoconodonts’, denticulate feeding structures, and a background of filamentous and spheroidal microbes. The annelids, wiwaxiids and priapulids are the first recorded from the Cambrian of Baltica. The File Haidar SCF assemblage is broadly comparable to those recovered from Cambrian basins in Laurentia and South China, though differences at lower taxonomic levels point to possible environmental or palaeogeographical controls on taxon ranges. These data reveal a fundamentally expanded picture of early Cambrian diversity on Baltica, and provide key insights into high‐latitude Cambrian faunas and patterns of SCF preservation. We establish three new taxa based on large populations of distinctive SCFs: Baltiscalida njorda gen. et sp. nov. (a priapulid), Baltichaeta jormunganda gen. et sp. nov. (an annelid) and Baltinema rana gen. et sp. nov. (a filamentous problematicum).     

Chancelloriid sclerites from the Lower Cambrian (Meishucunian) of eastern Yunnan,China, and the early history of the group

Chancelloriids are a group of enigmatic sessile animals that are covered with sclerites shaped as rosettes of spines, producing an appearance like that of a barrel cactus. They are known only from Cambrian rocks. Isolated sclerites of chancelloriids are widespread in small shelly faunas, but they have proven difficult to treat taxonomically due to the variation within and between individual animals. We report on large samples of chancelloriid sclerites from the Meishucunian (pre‐trilobitic Lower Cambrian) of eastern Yunnan Province, China, including material from the Dahai Member of the Zhujiaqing Formation (the Zhujiaqing section, Dahai, Huize County) and the Shiyantou Formation (the Xiaotan section, Yongshan County). The material from the Dahai Member appears to come from a single species, Chancelloriella irregularis. Statistical analysis of morphotype co‐occurrences in different samples suggests that several species are represented in the material from the Shiyantou Formation, which we herein tentatively place in four species, Allonnia erromenosa, Allonnia tetrathallis, Archiasterella charma sp. nov. and Archiasterella cf. pentactina. We suggest therefore that careful study of large collections of isolated chancelloriid sclerites permits the identification of different species, opening the possibility of their use in evolutionary or stratigraphical studies. In particular, in South China, it seems that Chancelloriella irregularis is characteristic of rocks of the Paragloborilus subglobosus – Purella squamulosa Assemblage Zone and the Watsonella crosbyi Assemblage Zone, whilst Archiasterella and Allonnia first appear in the overlying Sinosachites flabelliformis – Tannuolina zhangwentangi Assemblage Zone. C. irregularis is thus amongst the oldest well‐characterized chancelloriids, and the irregularity and poor organization of its sclerites relative to those of younger forms support the hypothesis that chancelloriid sclerites are compound structures arising from the fusion of originally separate elements, perhaps homologous with sclerites of halkieriids and other coeloscleritophorans. The similarity between the arrangement of rays in a chancelloriid sclerite and the arrangement of bubbles in small bubble clusters suggests that in many cases, aspects of the form of chancelloriid sclerites are the result of simple physical processes.     

The macro‐ and microfossil record of the Cambrian priapulid Ottoia

The stem‐group priapulid Ottoia Walcott, 1911, is the most abundant worm in the mid‐Cambrian Burgess Shale, but has not been unambiguously demonstrated elsewhere. High‐resolution electron and optical microscopy of macroscopic Burgess Shale specimens reveals the detailed anatomy of its robust hooks, spines and pharyngeal teeth, establishing the presence of two species: Ottoia prolifica Walcott, 1911, and Ottoia tricuspida sp. nov. Direct comparison of these sclerotized elements with a suite of shale‐hosted mid‐to‐late Cambrian microfossils extends the range of ottoiid priapulids throughout the middle to upper Cambrian strata of the Western Canada Sedimentary Basin. Ottoiid priapulids represented an important component of Cambrian ecosystems: they occur in a range of lithologies and thrived in shallow water as well as in the deep‐water setting of the Burgess Shale. A wider survey of Burgess Shale macrofossils reveals specific characters that diagnose priapulid sclerites more generally, establishing the affinity of a wide range of Small Carbonaceous Fossils and demonstrating the prominent role of priapulids in Cambrian seas.     

Oikozetetes from the early Cambrian of South Australia: implications for halkieriid affinities and functional morphology

Shells of Oikozetetes and isolated halkieriid sclerites from a section of the lower Cambrian Mernmerna Formation in the Flinders Ranges, South Australia, are tentatively considered as being derived from the same scleritome. Details of shell morphology and the possible combination of biomineralized shell and sclerites suggest that Oikozetetes , if interpreted correctly, is closely related to Halkieria . A new interpretation of Oikozetetes shell morphology, in addition to the first report of paired muscle scars on the interior surface, sheds new light on the possible functional morphology of halkieriid shells and the means of attaching the shell to the body. The occurrence of Oikozetetes in South Australia extends its biostratigraphic range to the lower Cambrian and biogeographic range to East Gondwana.     

Antipredator defense and phenotypic plasticity of sclerites from Renilla muelleri, a tropical sea pansy

Calcified sclerites are common in many benthic marine invertebrates, and despite their widespread occurrence, little is known about their ecological roles. Previous studies suggested that the sclerite composition of coral colonies may be altered in response to environmental cues such as predation and water motion. Furthermore, larger sclerites are thought to be more effective than small ones in deterring predators, while small sclerites may provide greater stiffness and resistance to deformation. The present study compared the length of the sclerites of the sea pansy Renilla muelleri from three depths in Guanabara Bay in southeastern Brazil. Our results show that sclerites are larger in deep-water specimens than in those from shallow water. Field assays were conducted in which sclerites from sea pansies at three depths were incorporated into artificial foods and offered to a natural assemblage of fish. These assays demonstrate that sclerites from R. muelleri from all three depths significantly reduced consumption by generalist carnivorous fishes. We conclude that R. muelleri uses skeletal elements not only to give the body its form but also as a defense against biotic threats.     

A redescription of Heterocotyle pastinacae Scott, 1904 (Monogenea: Monocotylidae) from Dasyatis pastinaca (Dasyatididae), with a neotype designation

Heterocotyle pastinacae Scott, 1904 is redescribed and a neotype is designated. H. pastinacae is most easily distinguished from other members of the genus by the elaborate arrangement of sclerotised spines in the vagina. The male copulatory organ also serves to differentiate it from other species in the genus. A possible function for the vaginal spines and unique form of the male copulatory organ is discussed. The use of head organ number as a generic character is considered questionable.     

贵州松桃寒武系清虚洞组Mongolitubulus squamifer 壳体显微结构初步研究

本文主要对贵州松桃寒武系清虚洞组灰岩中酸泡获得的管状化石Mongolitubulus squamifer壳体显微及亚显微结构进行研究。研究结果显示, M. squamifer不仅内层广泛发育纵向纤维结构,具鳞片的外层外壁也发育有微弱的纵向纤维结构,可能解释了管体外壁广泛发育的纵向开裂现象。在一枚保存有圆卵形鳞片以下部位的标本中发现,鳞片排列形式是由基部密集的小瘤点状颗粒然后过渡为不规则的圆卵形鳞片,到管体中上部逐渐形成规则排列的三角形鳞片。通过测量部分已报道的M.squamifer管体宽度与鳞片宽度发现正常发育的棘刺,鳞片大小与管体宽度有一定相关性。贵州松桃的M.squamifer壳层原始结构为2层,包括致密的具鳞片的外层和具明显纵向纤维结构的内层,管体内外层之间发育空隙导致管体容易破损。一些标本近基部断口处的层间空隙容易被次生矿物充填,导致内层增厚致密纤维结构消失或产生中间填充层。该研究揭示了M.squamifer的鳞片排列特征和壳体微观结构,为解释其亲缘关系提供新的形态学证据。     

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.     

The Collins’ monster,a spinous suspension-feeding lobopodian from the Cambrian Burgess Shale of British Columbia

Lobopodians, a paraphyletic group of rare but morphologically diverse Palaeozoic vermiform animals bearing metameric appendages, are key to the origin of extant panarthropods. First discovered in 1983 on Mount Stephen (Yoho National Park, British Columbia), the Cambrian (Wuliuan) Burgess Shale lobopodian nicknamed ‘Collins’ monster’ is formally described as Collinsovermis monstruosus gen. et sp. nov. A formal systematic treatment of the comparable and poorly known lobopodian Acinocricus stichus from Utah is also provided. The body of Collinsovermis is plump and compact but shows the diagnostic suspension-feeding characters of luolishaniid lobopodians. It possesses 14 contiguous pairs of lobopods, lacking space between them. The 6 anterior pairs are elongate, adorned with about 20 pairs of long and slightly curved ventral spinules arranged in a chevron-like pattern. These appendages terminate in a pair of thin claws and their dorsal surfaces are covered in minute spines or setae. The 8 posterior lobopod pairs, which attach to a truncated body termination, are stout and smooth, each terminated by a single strong recurved claw. Each somite bears a pair of dorsal spines; somites 4 and posteriad bear an additional median spine. The spines on somites 1–3 are much shorter than the spines on the remaining somites. The head is short, bears a terminal mouth and a pair of antenniform outgrowths, and is covered by an oblong sclerite. Collinsovermis, plus Collinsium and Acinocricus, are found to comprise a sub-group of stout luolishaniid lobopodians with remarkably long spinules on the front lobopods, interpreted here as a clade (Teratopodidae fam. nov.) This clade is distinct from both the comparatively slenderer Luolishania and a sub-group composed of Facivermis and Ovatiovermis lacking body sclerites. Luolishaniids were mostly sessile forerunners of arthropods that had coupled efficient suspension-feeding devices and, as in Collinsovermis, strong defensive or deterrent features.     

SKELETAL MICROSTRUCTURE INDICATES CHANCELLORIIDS AND HALKIERIIDS ARE CLOSELY RELATED

Abstract: Chancelloriids are problematic, sac‐like animals whose sclerites are common in Cambrian fossil assemblages. They look like sponges, but were united with the slug‐like halkieriids in the group Coeloscleritophora Bengtson and Missarzhevsky, 1981 based on a unique mode of sclerite construction. Because their body plans are so different, this proposal has never been well accepted, but detailed study of their sclerite microstructure presented here provides additional support for this grouping. Both taxa possess walls composed of a thin, probably organic, sheet overlying a single layer of aragonite fibres orientated parallel to the long axis of the sclerite. In all halkieriids and in the chancelloriid genus Archiasterella Sdzuy, 1969 , bundles of these fibres form inclined projections on the upper surface of the sclerite giving it a scaly appearance. On the lower surface of the sclerite, the projections are absent. This microstructure appears to be unique to chancelloriids, halkieriids, and their relatives, siphogonuchitids and sachitids. (The sclerites of another putative halkieriid relative, Wiwaxia Walcott, 1911 , are unmineralized, making direct comparisons impossible.) Thus, similarity both at the level of sclerite construction and the level of sclerite microstructure suggests that chancelloriid, halkieriid, sachitid, and siphogonuchitid sclerites are homologous. The difference in chancelloriid and halkieriid body plans can be resolved in two ways. Chancelloriids either represent a derived condition exhibiting complete loss of bilaterian characters or they represent the ancestral condition from which the bilaterally symmetric halkieriids, and the Bilateria as a whole, derived. The latter interpretation, proposed by Bengtson (2005) , implies that coeloscleritophoran sclerites (‘coelosclerites’) are a plesiomorphy of the Bilateria, lost or transformed in descendent lineages. Given that mineralized coelosclerites appear in the fossil record no earlier than c. 542 Ma, this in turn implies either that the Ediacaran record of bilaterians has been misinterpreted or that coelosclerite preservability increased at the beginning of the Cambrian Period. The former is difficult to reconcile with Ediacaran trace and body fossil evidence, but the latter may be possible, reflecting either independent mineralization of organic‐walled sclerites in chancelloriids and halkieriids or the opening of a taphonomic window that favours coelosclerite preservation.