Settling strategy of stalked echinoderms from the Kaili Biota (middle Cambrian), Guizhou Province,South China

A total of 124 articulated echinoderms, including Sinoeocrinus lui (n = 30), “S. globus” (n = 92), one indeterminate gogiid, and one indeterminate edrioasteroid, from the Kaili Formation (Middle Cambrian), eastern Guizhou, China was examined. Among them, 73% of gogiid echinoderms are preserved attached to skeletal substrates. Suitable skeletal substrates for gogiid holdfasts in the Kaili Biota are organophosphatic brachiopods; large trilobite fragments, including cranidia, free cheeks, thoracic segments, and pygidia; hyoliths; and Scenella shells. The high frequency of “S. globus” attached to organophosphatic brachiopods is due to the high richness of organophosphatic brachiopods rather than a host-specific association in the Kaili Biota. However, the possible discrimination against attachment to small trilobites, such as Pagetia, requires further investigation. Based on 11 hand samples, echinoderm population density (EPD) varies from 0.3 to 4.5 echinoderms per 50 grind points (0.06 to 0.96 echinoderms per cm²), depending on the percent coverage of skeletal debris. Crowding effect is evident when EPD is approximately 0.32 echinoderms per cm² and the skeletal coverage is less than 20%.     

Crypto-helical body plan in partially disarticulated gogiids from the Cambrian of South China

All living echinoderms have a pentaradial symmetry that is unique within the Bilateria. However, the Cambrian origin of echinoderm radial/pentaradiate symmetry is a long-standing problem. During the Cambrian (542–488 Ma), gogiids were the most common stalked echinoderm characterized by an “irregularly” plated body. Based on recently discovered material from the Balang Formation (Cambrian Series 2), eastern Guizhou, China, three unusual, partially disarticulated specimens of Guizhoueocrinus have clear evidence for a helical body plan. This helical plating is only evident in partially disarticulated specimens, thus a crypto-helical body construction is present. Crypto-helical construction in a gogiid raises the possibility of a phylogenetic connection among helicoplacoids, gogiids, and Helicocystis. The crypto-helical body construction may be an important evolutionary innovation among pre-radiate echinoderms.     

Cavity-dwelling organisms in Lower Cambrian patch reefs from southern Labrador

An intact and well-preserved biota of cavity-dwelling organisms (coelobionts) of early Cambrian age is found within growth-framework cavities in archaeocyathid patch reefs of the lower Forteau Formation (upper Bonnia-Olenellus Zone) of southern Labrador. The biota was diverse, and consisted of encrusting and vagrant organisms. Renalcis , a Renalcis-like form, Epiphyton, Girvanella , and Serligia , all lived attached to walls and roofs of the cavities; other attached forms were Bija, Archaeotrypa , and two types of Wetheredella , an agglutinated foraminifer. Other organisms, including trilobites, probable calcareous ostracodes, brachiopods, echinoderms, and worms, as well as fungi, are also found preserved within the cavities. The structure of the coelobiontic community is surprisingly modern; it indicates that as early as the Lower Cambrian coelobiontic organisms were well-established in reefs, and were an important part of the reef community as a whole.     

Cambrian spiral-plated echinoderms from Gondwana reveal the earliest pentaradial body plan

Echinoderms are unique among animal phyla in having a pentaradial body plan, and their fossil record provides critical data on how this novel organization came about by revealing intermediate stages. Here, we report a spiral-plated animal from the early Cambrian of Morocco that is the most primitive pentaradial echinoderm yet discovered. It is intermediate between helicoplacoids (a bizarre group of spiral-bodied echinoderms) and crown-group pentaradiate echinoderms. By filling an important gap, this fossil reveals the common pattern that underpins the body plans of the two major echinoderm clades (pelmatozoans and eleutherozoans), showing that differential growth played an important role in their divergence. It also adds to the striking disparity of novel body plans appearing in the Cambrian explosion.     

Traces of predation in the Cambrian

Series two marks a revolution in Cambrian predation when new predators and new predation methods appeared, which led to general increase in predation intensities and in the diversity of prey groups. The number of bored taxa and taxa with the predation scars is similar in the Cambrian. Most of the borings are associated with brachiopods and most of the scars with trilobites. Brachiopods, arthropods, molluscs, cnidarians and echinoderms were the most common prey in the Cambrian. The Cambrian record of predation is dominated by damage inflicted on brachiopods and trilobites. The fossils with predation signs are known from a majority of paleocontinents and all the Cambrian series.     

First report of Gogia (Eocrinoidea, Echinodermata) from the Early-Middle Cambrian of Sonora (Mexico), with biostratigraphical and palaeoecological comments

The blastozoan echinoderm genus Gogia is reported for the first time in the Early and the Middle Cambrian of Mexico. Reports in different members of the section of San José de Gracia (Sonora State, northwestern Mexico) extend the palaeogeographical range of the genus to the South Laurentia, and the stratigraphic range of Gogia granulosa to the whole first half of middle Middle Cambrian. Isolated plates occur in rocks deposited in detrital inner platform and complete specimens, in carbonate outer platform, confirming their ability to live in diverse environments. Their presence in these different environments through the Early-Middle Cambrian on Laurentia agrees with the onshore-offshore expansion of echinoderms during Cambrian.     

Deciphering the early evolution of echinoderms with Cambrian fossils

Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister‐group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm‐type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan.     

Diseases of echinoderms

Diseases of echinoderms are poorly documented. Most reports concern biotic diseases caused by animal agents. While parasites on echinoderms have been described in increasing numbers of papers for more than one century, the host-parasite relationship and the effects of parasitism on echinoderm life-cycles were rarely considered. The parasitic fauna differs markedly according to the echinoderm group concerned, depending on various factors such as feeding-habits or symbiogenesis. Microorganismic diseases undoubtedly occur in echinoderms but they were not investigated until recently. Microorganisms have frequently been assumed to act as agents causing mass mortalities. As for stress-caused diseases, the only — and very preliminary — data available concern almost exclusively those induced by pollution. Since echinoderms are major components of benthic ecosystems, echinoderm diseases may be expected to exert prominent ecological effects.     

Middle Cambrian cystoid (sensu lato) stem columnals from Bornholm, Denmark

A rich material of echinoderm fragments from two Middle Cambrian stratigraphical levels on Bornholm are preserved due to phosphatization of the original calcitic stereom. Preservation of echinoderms in this way - not previously recorded from the Middle Cambrian - permits detailed analysis of the three-dimensional stereom structure. Identifiable are fragments of stylophorans and eocrinoids. Stem columnals, most likely from eocrinoids, show a wide and advanced morphological variation indicating articulation similar to that of crinoids. The material from the Exsulans Limestone/Kalby marl ( Ptychagnostus gibbus Zone) represents stem-bearing cystoids older than Akadocrinus from Bohemia. The Andrarum Limestone ( Sole-nopleura brachymetopa Zone) contains echinoderm fragments from a higher stratigraphical level, a level correlatable with that from which the oldest North American stem–bearing cystoid, Eustypocystis , has been recorded.     

Plated Cambrian bilaterians reveal the earliest stages of echinoderm evolution

Echinoderms are unique in being pentaradiate, having diverged from the ancestral bilaterian body plan more radically than any other animal phylum. This transformation arises during ontogeny, as echinoderm larvae are initially bilateral, then pass through an asymmetric phase, before giving rise to the pentaradiate adult. Many fossil echinoderms are radial and a few are asymmetric, but until now none have been described that show the original bilaterian stage in echinoderm evolution. Here we report new fossils from the early middle Cambrian of southern Europe that are the first echinoderms with a fully bilaterian body plan as adults. Morphologically they are intermediate between two of the most basal classes, the Ctenocystoidea and Cincta. This provides a root for all echinoderms and confirms that the earliest members were deposit feeders not suspension feeders.     

凯里生物群腕足动物与游泳动物共存现象研究

腕足动物是贵州凯里生物群中重要的化石门类,不仅化石数量丰富,分异度高,还具有丰富的生态现象。在寒武系的化石记录上,腕足动物常与海绵、藻类、棘皮动物、水母状生物、软舌螺、威瓦西亚虫、其他腕足动物等保存在一起,凯里生物群中的腕足动物也有类似的共存现象。本文就凯里生物群中腕足动物与游泳动物的共存现象进行了初步研究,认为腕足动物与贵州拟轮盘水母Pararotadiscus guizhouensis存在共栖、共埋两种关系,讨论了这两种关系的不同之处;分析了腕足动物与大型双瓣壳节肢动物加拿大虫Canadaspis的共存现象,推测一只腕足动物与Canadaspis共同保存的化石记录表明这两种生物存在共栖关系。本文的研究丰富了凯里生物群物种之间的生态关系,为凯里生物群的生态多样性提供了更多的证据。     

Cambrian stalked echinoderms show unexpected plasticity of arm construction

Feeding arms carrying coelomic extensions of the theca are thought to be unique to crinoids among stemmed echinoderms. However, a new two-armed echinoderm from the earliest Middle Cambrian of Spain displays a highly unexpected morphology. X-ray microtomographic analysis of its arms shows they are polyplated in their proximal part with a dorsal series of uniserial elements enclosing a large coelomic lumen. Distally, the arm transforms into the more standard biserial structure of a blastozoan brachiole. Phylogenetic analysis demonstrates that this taxon lies basal to rhombiferans as sister-group to pleurocystitid and glyptocystitid blastozoans, drawing those clades deep into the Cambrian. We demonstrate that Cambrian echinoderms show surprising variability in the way their appendages are constructed, and that the appendages of at least some blastozoans arose as direct outgrowths of the body in much the same way as the arms of crinoids.     

Early Cambrian coelobiontic communities in tectonically unstable crevices developed in Neoproterozoic andesites, Ossa-Morena, southern Spain

A well-preserved biota of Lower Cambrian cavity-dwelling organisms is recorded within fissures in Neoproterozoic andesites in Ossa-Morena (southern Spain). The cavities are unique among described Lower Cambrian coelobiontic communities due to the igneous character of the host rock. Coelobiontic habitat was episodically enlarged by synsedimentary tectonic fracturing reflecting polyphase infill of recurrent facies. The pioneer coelobiontic biota was diverse, and consisted of encrusting stromatolites and thromboids (dominated by Epiphyton and Renalcis), attached to walls and ceilings of the cavities, associated with archaeocyaths. Sponge spicules and chancelloriid sclerites occur as dense clusters indicating in situ growth, death and decay of spiculate sponges and coeloscleritophorans. Other organisms, such as echinoderms, trilobites and brachiopods, are also found within the cavities as reworked skeletons, indicating that they were washed in from the overlying, open seafloor. The main feature of the coelobiontic biota is the dominance of a sessile, chemosynthetic and filter-feeding epibenthos, composed of microbial communities, archaeocyaths, spiculate sponges (demosponges and rarer hexactinellides) and coeloscleritophorans.     

Quantification of echinoderms (Echinodermata) on Georges Bank,and the potential influence of marine protected areas on these populations

The spatiotemporal variation of the distribution of echinoderms in the Georges Bank ecosystem was examined from 2005 to 2012. Density and abundance of representatives from echinoderm classes (brittle stars, sand dollars, sea stars, and sea urchins) were estimated using a drop camera video survey of the benthos in areas open and closed to fish trawling. The influence of closed areas on these echinoderm populations relative to a suite of key environmental and biotic factors was evaluated using Canonical Correspondence Analysis (CCA). While marine protected areas appeared to influence the distribution of sand dollars and sea stars, the distribution of brittle stars and sea urchins seemed to be defined mainly by preferred habitat conditions. According to the CCA, depth, sediment stability, temperature, predator abundance, and management area were the most important factors explaining this echinoderm assemblage. On Georges Bank, echinoderms dominate the benthic biota and are present in a variety of habitats. They can alter marine communities and are preferred prey and main predators for several commercially targeted species. The detailed information presented here (on the scale of kilometers) on abundance and spatial distribution of these populations is thus valuable towards the implementation of ecosystem‐based fisheries management.     

Marine carbonate facies in response to climate and nutrient level: The upper carboniferous and permian of central spitsbergen (Svalbard)

Summary Carbonate-dominated successions of the Gipsdalen and Tempelfjorden Groups from Svalbard record a significant shift from Photozoan to Heterozoan particle associations in neritic settings during the late Palaeozoic. During the Bashkirian, benthic particle associations which included photoautotrophs such as phylloid algae (Chloroforam Association) characterised shallow subtidal environments. Most depositional settings which endured siliciclastic terrestrial input exhibited poorly diversified associations dominated by brachiopods, bryozoans and siliceous sponges (Bryonoderm Association). During the Moscovian to Asselian, highly diversified associations typified by various calcareous green algae,Palaeoaplysina, Tubiphytes, fusulinids, smaller and encrusting foraminifers (Chloroforam Association) prevailed in carbonate sediments from supratidal to shallow subtidal environments. During the Sakmarian and Early Artinskian, oolitic carbonate sands (Chloroforam Association) typified intertidal flats, whereas shallow subtidal environments were occupied by moderately diversified associations with fusulinids, smaller foraminifers, echinoderms and bryozoans (Bryonoderm-extended Association) and poorly diversified associations with echinoderms, brachiopods and bryozoans (Bryonoderm Association). During the Late Artinskian to Kazanian, poorly diversified associations characterised by brachiopods, echinoderms and bryozoans (Bryonoderm Association), and sponge-dominated associations (Hyalosponge Association) reigned within siliceous carbonates of intertidal and shallow subtidal environments. This trend is interpreted as a result of climatic cooling and fluctuations of prevailing levels of trophic resources within shallow-water settings during the studied time period. While raised nutrient levels were restricted to near-shore settings during the Bashkirian, steady mesotrophic conditions arose from the Sakmarian onward and increased to late Permian times.     

THE EARLY RADIATION AND PHYLOGENY OF ECHINODERMS

1. Living echinoderms are characterized by an extensive water vascular system developed from the larval left hydrocoel, a complex, multi-plated endoskeleton with stereom structure, and pentamery. Fossil evidence shows that stereom evolved before pentamery, but both were acquired during the Lower Cambrian. 2. Cladistic analysis of Lower Cambrian genera reveals very few characters in common between carpoids and true echinoderms, and that the split between them was the first fundamental evolutionary dichotomy within the Dexiothetica. 3. Helicoplacoids are stem group echinoderms with spiral plating and three ambulacra arranged radially around a lateral mouth. They are the most primitive echinoderms and the first to show a radial arrangement of the water vascular and ambulacral systems. Unlike later echinoderms, their skeleton shows no dorsal/ventral (aboral/oral) differentiation. They were probably sedentary suspension feeders. 4. Camptostroma is the most primitive known pentaradiate echinoderm and, in our view, possibly a common ancestor of all living groups. It had a short conical dorsal (aboral) surface with imbricate plating, a ridged lateral wall and a slightly domed ventral (oral) surface with five curved ambulacra in a 2-1-2 arrangement inherited from the triradiate pattern of the helicoplacoids. Interambulacral areas bore epispires and the CD interambulacrum contained the anus, hydropore and/or gonopore. All parts of the theca had plates in at least two layers. 5. All other echinoderms belong to one of two monophyletic subphyla, the Pelmatozoa and the Eleutherozoa. 6. Stromatocystites is the earliest known eleutherozoan and differs from Camptostroma in having a test with only one layer of plates and having lost the dorsal elongation. In Stromatocystites the dorsal surface is flat and the plating tesselate. Stromatocystites was an unattached, low-level suspension feeder. 7. The lepidocystoids are the earliest known pelmatozoans. They differ from Camptostroma in having an attached dorsal stalk which retained the primitive imbricate plating, and by developing erect feeding structures along the ambulacra. In Kinzercystis, the ambulacra are confined to the thecal surface and erect, biserial brachioles arise alternately on either side. Lepidocystis has a similar arrangement except that, the distal part of each ambulacrum extends beyond the edge of the theca as a free arm. 8. Pelmatozoans diverged more or less immediately into crinoids, with multiple free arms composed of uniserial plates, and cystoids sensu lato, which retained brachioles. Gogia (Lower to Middle Cambrian) is the most primitive known cystoid and differs from Kinzercystis principally in having all plating tesselate, while Echmatocrinus (Middle Cambrian) is the most primitive known crinoid and differs from Lepidocystis in lacking brachioles and in having more than five free arms with uniserial plates. 9. Post Lower Cambrian differentiation of pelmatozoan groups proceeded rapidly, exploiting the primitive suspension-feeding mode of life. Maximum morphological diversity was reached in the Ordovician, but thereafter crinoids progressively displaced cystoid groups and reached their peak diversity during the Carboniferous. The eleutherozoans were slower to diversify, but by the Arenig the earliest ‘sea-stars’ (in reality, advanced members of the eleutherozoan stem group) had reversed their living orientation and had begun to exploit a deposit-feeding mode of life. These in turn led to the ophiuroids, echinoids and holothuroids. 10. The basic echinoderm ambulacrum was already present in the helicoplacoids. It had biserial, alternate flooring plates and complexly plated sheets of cover plates on either side. The radial water vessel lay in the floor of the ambulacrum, external to the body cavity, and gave rise ventrally to short, lateral branches (fore-runners of tube feet) that were used to open the cover plate sheets, and dorsally was connected to internal compensation sacs which acted as fluid reservoirs (and were preadapted for a role in gaseous exchange). Plating on the cover plate sheets was organized and reflected the positions of the lateral branches from the radial water vessel. In Camptostroma, the cover plate sheets had biserially aligned rows of cover plates associated with the lateral branches. 11. Brachioles arose by extension of the lateral branches of the radial water vessel and associated serially aligned cover plates found in Camptostroma. They bear a single alternate series of cover plates. In Lepidocystis the ambulacra extended beyond the edge of the oral surface as true arms. Brachial plates of arms are homologues of primary ambulacral flooring plates, and arms bear multiple series of cover plates. Uniserial ambulacral plating is a derived condition and evolved independently in crinoids, paracrinoids and isorophid edrioasteroids. Pinnules in crinoids arose independently in inadunates and camerates by a progressively more unequal branching of the arms. Thus all parts of the subvective system in crinoids are internally homologous, whereas in cystoids, brachioles and arms (or ambulacra) are not homologous structures. 12. The position of the hydropore is the best reference point in orientating echinoderms. Carpenter's system of identifying ambulacra by letters, arranged clock-wise in oral view with the A ambulacrum opposite the hydropore, is consistent in all echinoderm classes. In all Lower Cambrian pentaradiate echinoderms the anus, gonopore and hydropore lie in the CD interambulacrum and this is accepted as the primitive arrangement. In helicoplacoids we tentatively suggest that the A ambulacrum spiralled down from the mouth while the two ambulacra that spiralled up represent the B + C and D + E ambulacra combined. 13. The pelmatozoan stem arose from a polyplated stalk, via a meric stem to a true column with holomeric (single piece) columnals. This happened independently in the crinoids and the cystoids. 14. Our analysis of echinoderm phylogeny leads us to recommend the following changes to the higher level classification of echinoderms: The phylum Echinodermata includes only those groups with radial symmetry superimposed upon a fundamental larval asymmetry. It has a stem group that contains the triradiate helicoplacoids and a crown group to which all other (pentaradiate) echinoderms belong. The crown group contains two monophyletic subphyla, the Pelmatozoa and the Eleutherozoa, and the Pelmatozoa contains two superclasses, the Crinoidea which are extant and the Cystoidea, which are extinct.     

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.     

A spinose stem group brachiopod with pedicle from the Middle Cambrian Burgess Shale

A new genus and species of a Middle Cambrian stem group brachiopod, Acanthotretella spinosa n. gen. and n. sp., is described from the Burgess Shale Formation. Most of the 42 specimens studied came from the Greater Phyllopod bed (Walcott Quarry) and were collected from five bed assemblages, each representing a single obrution event. Specimens are probably preserved within their original habitat. In contrast to all brachiopods known from the Burgess Shale, the shells of the new stem group brachiopod are often deformed and do not show signs of brittle breakage, which suggests that the valves were originally either entirely organic in composition or, more likely, had just a minor mineral component. Acanthotretella spinosa differs from all the other described Cambrian brachiopods in that it is covered by long, slender and possibly partly mineralized spines that are posteriorly inclined at an oblique angle away from the anterior margin. The spines penetrate the shell and are mainly comparable with the thorn‐like organic objects that have been inferred from early siphonotretoid brachiopods. The pedicle was slender and was composed of a central coelomic region and emerged from an apical foramen at the end of an internal pedicle tube. The finding of a pedicle attached to the macrobenthic algae Dictyophycus and other epibenthos implies that A. spinosa did not have an infaunal mode of life. The visceral region and interior characters are poorly preserved.     

Sea urchins, sea stars and brittle stars from Terra Nova Bay (Ross Sea, Antarctica)

Although echinoderms constitute some of the most conspicuous taxa of the Antarctic benthic communities, the echinoderm fauna of Terra Nova has not been described yet. The present study provides the first species list of echinoids, ophiuroids and asteroids from Terra Nova Bay (30–500 m depth) and describes the depth distribution of these species. Preliminary observations of the summer reproductive condition of some of the species are also included.