Ronald L. Parsley

Are homalozoans echinoderms? An answer from the extraxial-axial theory

Abstract Homalozoans include four classes of non-pentamerous Paleozoic echinoderms: Homostelea (cinctans), Ctenocystoidea (ctenoid-bearing homalozoans), Homoiostelea (solutes), and Stylophora (cornutes and mitrates). Their atypical morphologies have historically made it difficult to relate them to other classes. Therefore, their systematic positions have been represented by two hypotheses (H): as stem taxa to echinoderms (H1) or as stem taxa to chordates (H2). These conclusions rest on previous inability to recognize synapomorphies with more crownward echinoderms, resulting in a forcing of the homalozoans down the phylogenetic tree that is more artifactual than evolutionary. The Extraxial-Axial Theory (EAT) identifies body-wall homologies, common ontogenetic patterns, and major events in bodyplan evolution. Therefore, the EAT can identify synapomorphies among even the most disparate of echinoderms. Application of the EAT undermines both H1 and H2 and strongly suggests that the bizarre asymmetry of homalozoans is a derived characteristic, and not indicative of plesiomorphic morphology for either chordates or echinoderms. Each of the four homalozoan clades and their major features are reexamined using the EAT. New findings are presented concerning homologies of thecal body wall, but we focus on stems, arms, and brachioles, which are recognized as very distinct products of independent evolutionary events. The results support a new interpretation (H3) of homalozoans as a polyphyletic assemblage that can be parsed out into other, clearly echinoderm clades. The Homoiostelea and Homostelea share the blastozoan synapomorphy of a brachiole. The enigmatic Ctenocystoidea also seem to have brachioles. The Stylophora have an arm as in crinoids. H3 is also more congruent with the known fossil record. Although they are stratigraphically early echinoderms, homalozoans are not indicative of the plesiomorphic morphology of the phylum.

Towering sponges in an Early Cambrian Lagerstätte: Disparity between nonbilaterian and bilaterian epifaunal tierers at the Neoproterozoic-Cambrian transition

Epifaunal, suspension-feeding bilaterian animals in the Cambrian lived close to the sediment-water interface, and hence their ecological tiering levels were low (<10 cm). Here we report an Early Cambrian (Diandongian or probably Tommotian-Atdabanian) Lagerstatte from the Hetang Formation in Anhui Province, south China. The Hetang biota is characterized by high-tiering (to 50 cm) sponges and small (<0.5 cm) bilaterians (including orthothecid hyoliths and bivalved arthropods). Nonbilaterian suspension feeders (sponges, cnidarians, and archaeocyathids) as high-tiering animals and bilaterian suspension feeders as low-tiering animals also characterize other Neoproterozoic-Cambrian assemblages, such as the Ediacaran, Chengjiang, Burgess Shale, and Sinsk biotas. These data are consistent with medium- to high-tiering levels in Neoproterozoic-Cambrian epifaunal communities, but suggest that nonbilaterians achieved such tiering levels long before bilaterian suspension feeders did so in the Early Ordovician. The disparity between bilaterian and nonbilaterian tierers during the Neoproterozoic-Cambrian transition and the delayed appearance of high-tiering bilaterians demand phylogenetic and ecological explanations. The Cambrian substrate revolution may have triggered a cascade of ecological evolution, including the rise of bilaterian animals in high-tiering levels during the Ordovician radiation of the Paleozoic fauna.

LONG STALKED EOCRINOIDS IN THE BASAL MIDDLE CAMBRIAN KAILI BIOTA, TAIJIANG COUNTY, GUIZHOU PROVINCE, CHINA

Abstract Long-stemmed eocrinoids are limited to two species in the basal Middle Cambrian Kaili Biota, which occupies the middle portion of the Kaili Formation, Taijiang County, Guizhou Province, China. The Kaili Biota contains preserved soft-bodied organisms shared with either the Chengjiang Fauna (Southwest China) or the Burgess Shale Fauna (British Columbia) or with both. Echinoderms are preserved as limonitic external molds that produce excellent latex casts. Sinoeocrinus lui Zhao et al., 1994 has a complex ontogenetic development, which is described in terms of morphology of holdfast, number of thecal plate circlets, addition and morphology of thecal pores, ambulacral arrangement, and number of brachioles relative to thecal height. Because of the complex ontogeny the following species are now seen to be synonymous with S. lui: S. curtobrachiolus Zhao et al.,1994; S. lepidus Zhao et al., 1994; S. longus Zhao et al., 1994; S. minus Zhao et al., 1994; Paragogia globosa Zhao et al., 1994; and Curtoeocrinus guizhouensis Zhao et al., 1994. A second and rare eocrinoid of undetermined familial and ordinal placement, Balangicystis rotundus n. gen. and sp., has an unusually long holdfast and poreless thecal plates with prominent radial ridges. Sinoeocrinus lui and Balangicystis rotundus inhabited the outer shelf in disaerobic fine-grained shales and mudstones. Megascopic infauna in their community is not present.

Kailidiscus, A New Plesiomorphic Edrioasteroid from the Basal Middle Cambrian Kaili Biota of Guizhou Province, China

Abstract A new genus and species of edrioasteroid grade echinoderm, Kailidiscus chinensis, is described from the Kaili Biota of the basal lower Middle Cambrian Kaili Formation from Guizhou Province, China. This echinoderm has a strong resemblance to isorophid edrioasteroids, bearing a well-developed peripheral rim, long curved ambulacra, and slightly imbricate interambulacral plating at maturity. However, the presence of pore-bearing, double biserial floor plates, tiered cover plates, lack of radially positioned oral frame plates, and unincorporated hydropore/gonopore are unknown in isorophids. Many of these features bear strong resemblance to eocrinoids and helicoplacoids, attesting to the plesiomorphic nature of this taxon. Despite the obvious anatomical differences, specimens of this species preserve a complete ontogeny that resembles that of isorophids. Juveniles show a discoidal theca with straight ambulacra that transitions to an inflated theca with strongly curved ambulacra with maturity.

Basal Middle Cambrian Short-Stalked Eocrinoids from the Kaili Biota: Guizhou Province, China

Abstract Gogiid eocrinoids (Echinodermata) are the most abundant coelomate invertebrates in the Middle Cambrian (Taijiangian) Kaili Biota, Taijiang County, Guizhou Province, China. Both long- and short-stalked taxa are represented. The short-stalked Globoeocrinus globulus n. gen. and sp. has a globular theca, with well-developed sutural pores on all thecal plates in older mature specimens, short, thick stalk composed of small polygonal to rounded platelets, a large flattened attachment disc, and up to ten long, thin, helically coiled brachioles with tall, pointed cover plates. Some younger mature specimens have varying patches of non-pored plates that usually occur on the upper (probably) posterior portion of the theca. This species is over three times more abundant than the other co-occurring long-stalked gogiid Sinoeocrinus lui Zhao, Huang and Gong, 1994. Most specimens of G. globulus (ca. 80%) attach to acrotretid brachiopods, trilobite exuviae, or other skeletal remains. They are also capable of attaching directly to the seafloor.

Revision of lyracystid eocrinoids from the Middle Cambrian of South China and Western Laurentia

Abstract The Middle Cambrian eocrinoid genera Lyracystis Sprinkle and Collins, 2006, from western Laurentia and Balangicystis Parsley and Zhao, 2006, from South China, described in the same year, have turned out to be closely related genera assigned to the Family Lyracystidae. Both have erect, lyre-shaped, arm-like, brachiole-bearing, feeding appendages, here termed exothecal ambulacra, that are not homologous to crinoid arms. They also have a long, multiplated stalk to elevate the theca and feeding appendages well above the sea floor, making them among the highest tiered echinoderm suspension feeders known from the Middle Cambrian. The long stalk was either inserted a short distance into the muddy sediment, or attached to rare skeletal fragments lying on the sea floor. Both genera seem well adapted to quiet-water or slow-current conditions in deeper water (150–200 m) on the outer shelf or upper slope of their respective continents.

Morphology and biomechanical implications of isolated discocystinid plates (Edrioasteroidea, Echinodermata) from the Carboniferous of North America

Detailed examination of isolated thecal plates belonging to three discocystinid edrioasteroids, Spiraclavus nacoensis Sumrall, Hypsiclavus kinsleyi Sumrall, and Giganticlavus bennisoni Sumrall and Bowsher, reveals striking similarity in morphology among these species. Stereom observed in the ambulacral floor plates indicates that ligamentous connective tissue opened the ambulacral cover plates and muscle tissue closed them. The ambulacral floor plates are interpreted as rigid supports for the oral surface with the interambulacral areas acting as flexible integuments of plates. The aboral surface is interpreted as flexible and highly contractile. All discocystinid thecal openings are consistent in morphology with adaptations for thecal pressurization. Extension and contraction of the theca was accomplished by pumping water in and out of an inflatable sac associated with the periproctial opening. The pedunculate zone is interpreted as passively expanding and contracting by relaxing of mutable collagenous tissue and stiffening when the theca was in the desired position. All of these features illustrate that discocystinid edrioasteroids have highly–evolved morphology and function.

Chapter 13 Cambrian echinoderm diversity and palaeobiogeography

Abstract The distribution of all known Cambrian echinoderm taxa, encompassing both articulated specimens and taxonomically diagnostic isolated ossicles, is documented for the first time. The database described by 2011 comprises 188 species recorded from 65 formations from around the world. Formations that have yielded articulated echinoderms are unequally distributed in space and time. Only Laurentia and West Gondwana provide reasonably complete records at the resolution of Stage. The review of the biogeographical distributions of the eight major echinoderm clades shows that faunas from Laurentia and Northeast Gondwana (China and Korea) are distinct from those of West Gondwana and Southeast Gondwana (Australia); other regions are too poorly sampled to make firm palaeobiogeographical statements. Analysis of alpha diversity (species per formation) shows that diversity rose initially to Cambrian Stage 5, declined into Guzhangian and Paibian before returning to Stage 5 levels by the end of the Cambrian. This pattern is replicated in Laurentia and West Gondwana. We show that taxonomically diagnostic ossicles found in isolation typically occur significantly earlier than the first articulated specimens of the same taxa and provide important information on the first occurrence and palaeobiogeographical distribution of key taxa, and of the phylum as a whole. Supplementary material: Articulated Cambrian echinoderms and Isolated plates of Cambrian echinoderms are provided at: http://www.geolsoc.org.uk/SUP18668

Mandalacystis, a new rhipidocystid eocrinoid from the Whiterockian Stage (Ordovician) in Oklahoma and Nevada

Mandalacystis dockeryi n. gen. and sp. is known from seven complete or nearly com- plete thecae, six partial thecae, and numerous separate plates from the Oil Creek Formation in south-central Oklahoma, and a single thecal plate from the Antelope Valley Limestone in Nevada. Details of skeletal microstructure (stereom) are well preserved, permitting scanning electron mi- croscopy for the first time in the Eocrinoidea, and demonstrating that plate thickness on thecal faces is only 60-70 clm. Mandalacystis is characterized by labyrinthine ridges on both faces of the theca, relatively short brachioles, and a normal stem with holdfast (at least in juvenile stages). Based on the normal biserial arrangement of brachiolar plates in the brachioles and the possession of a normal stem, Mandalacystis appears to be one of the most primitive rhipidocystids, closely related to Petalocystites. Mandalacystis dockeryi lived in a storm-dominated, shallow-water offshore habitat on patches of skeletal debris surrounded by soft, terrigenous mud. This setting may have been conducive to the transition from an erect, attached mode of life to the prone, unattached lifestyle of later rhipidocystids.

Ontogeny, Functional Morphology, and Comparative Morphology of Lower (Stage 4) and Basal Middle (Stage 5) Cambrian Gogiids, Guizhou Province, China

Abstract Three gogiid eocrinoids, numbering in the thousands of specimens, are well known from the lower Cambrian (Stage 4) Balang Formation (Guizhoueocrinus yui) and basal middle Cambrian (Stage 5) Kaili Formation (Kaili Biota) (Sinoeocrinus lui and Globoeocrinus globulus) that resided on the Yangtze-South China Plate (modern Guizhou Province, China). In each species a complete ontogenetic sequence, using thecal height (TH) as a scale, juvenile stage (early, middle, and late substages), mature stage (early, middle, and late substages), and a gerontic stage can be identified. Sutural pores appeared in an orderly sequence; below the ambulacrals, above the stalk and generally over the theca, in that order. In younger species their emplacement and growth was commonly precocious relative to G. yui. Also, their shape ranges from circular to oval to triangular in a single growth sequence, after the establishment of the 2-1-2 pattern; brachioles were added in series of five and were also precocious in time of occurrence in younger species. Thecae in younger species tend to retain juvenile aspects relative to G. yui. All of these species lived in outer shelf settings in fine-grained, organic-rich siliciclastics.