Jisuo Jin

Chapter 11 Biodiversity, biogeography and phylogeography of Ordovician rhynchonelliform brachiopods

Abstract The phylogeographical evolution and the consequent changing distribution and diversity of rhynchonelliform brachiopods through the Ordovician are linked to the dynamic palaeogeography of the period. The Early Ordovician (Tremadocian and Floian) is characterized by globally low-diversity faunas with local biodiversity epicentres, notably on the South China Palaeoplate; low-latitude porambonitoid-dominated faunas with early plectambonitoid and clitambonitoid representatives, as well as high-latitude assemblages mostly dominated by orthoids, can be recognized, but many taxa are rooted in Late Cambrian stocks. The Early Ordovician displays a steady increase in rhynchonelliformean biodiversity, which was mostly driven by the increasing success of the Porambonitoidea and Orthoidea, but the billingsellids and early plectambonitoids also contributed to this expansion. During the Early to Mid Ordovician (Dapingian–Darriwilian), marine life experienced an unprecedented hike in diversity at the species, genus and family levels that firmly installed the suspension-feeding benthos as the main component of the Palaeozoic fauna. However, this may have occurred in response to an early Darriwilian annihilation of existing clades, some of which had been most successful during the Early Ordovician. New clades rapidly expanded. The continents were widely dispersed together with a large number of microcontinents and volcanic arcs related to intense magmatic and tectonic activity. Climates were warm and sea-levels were high. Pivotal to the entire diversification is the role of gamma (inter-provincial) diversity and by implication the spread of the continents and frequency of island arcs and microcontinents. The phylogeographical analysis demonstrates that this new palaeogeographical configuration was particularly well explored and utilized by the strophomenides, especially the Plectambonitoidea, which radiated rapidly during this interval. The porambonitoids, on the other hand, were still in recovery following the early Darriwilian extinctions. Orthides remained dominant, particularly at high latitudes. Biodiversity epicentres were located on most of the larger palaeoplates, as well as within the Iapetus Ocean. Provincial patterns were disrupted during the Sandbian and early Katian with the migration of many elements of the benthos into deeper-water regimes, enjoying a more cosmopolitan distribution. Later Katian faunas exhibit a partition between carbonate and clastic environments. During the latest Katian, biogeographical patterns were disrupted by polewards migrations of warm-water taxa in response to the changing climate; possibly as a consequence of low-latitude cradles being developed in, for instance, carbonate reef settings. Many clades were well established with especially the strophomenides beginning to outnumber the previously successful orthides, although this process had already begun, regionally, in the mid to late Darriwilian. At the same time, atrypoid and pentameroid clades also began to radiate in low-latitude faunas, anticipating their dominance in Silurian faunas. The Hirnantian was marked by severe extinctions particularly across orthide-strophomenide clades within the context of few, but well-defined, climatically controlled provincial belts. Supplementary material: The individual localities and a reference list for the data sources are provided at: http://www.geolsoc.org.uk/SUP18667

Precisely locating the Ordovician equator in Laurentia

The Late Ordovician equatorial zone, like the zone today, had few hurricane-grade storms within 10o of the equator, as emphasized by the preservation of massive-bedded Thalassinoides ichnofacies in a trans-Laurentian belt more than 6000 km long, from the southwestern United States to North Greenland. That belt also includes nonamalgamated shell beds dominated by the brachiopod Proconchidium , which would not have been preserved after hurricane-grade storms. The belt lacks such storm-related sedimentary features as rip-up clasts, hummocky cross-stratification, or large channels. In contrast, other contemporaneous Laurentian Thalassinoides facies and shell beds on either side of the belt have been disturbed by severe storms below fair-weather wave base. The position of the biofacies-defined equatorial belt coincides with the Late Ordovician equator deduced from paleomagnetic data from Laurentia, thus providing both a high-precision equatorial location and an independent test of the geocentric axial dipole hypothesis for that time.

Relic aragonite from Ordovician-Silurian brachiopods: Implications for the evolution of calcification

Understanding the influence of aragonite/calcite sea conditions on the evolution of biocalcification relies strongly on the correct interpretation of the original composition of calcareous taxa. Aragonite dissolves or inverts into calcite over geological time, and its preservation is currently unknown to predate the Pennsylvanian. Here we present direct evidence for the common occurrence of relic aragonite in Ordovician and Silurian trimerellid brachiopods, thereby extending the known range of aragonite preservation by more than 130 million years. Together with associated hypercalcifying taxa of putatively original aragonite or high-magnesium calcite composition and considerations of the temperature dependence of aragonite and calcite precipitation, our results suggest that the evolution of aragonite biomineralization might have presented an adaptive advantage in shallow marine tropical waters of calcite seas. A targeted search for Paleozoic aragonite should both resolve the original composition of consistently recrystallized taxa and enable the reassessment of the aragonite/calcite sea paradigm in a paleoenvironmental context.

EPIPUNCTAE AND PHOSPHATIZED SETAE IN LATE ORDOVICIAN PLAESIOMYID BRACHIOPODS FROM ANTICOSTI ISLAND, EASTERN CANADA

Abstract Epipunctae, a new type of shell perforation, are well developed in typical taxa of the family Plaesiomyidae, a group of common orthide brachiopods from Laurentia and some other tropically located tectonic plates of Late Ordovician age. These minute, prominently elongate, tubular structures are similar to endopunctae in size and density, but differ in being oblique, intersecting the shell surface at a relatively low angle, and being confined largely to the outer portion of the shell wall. The tubules are similar in orientation to aditicules within the same shells but are much smaller and denser, usually aligned along fine growth lines and arranged in crude longitudinal columns. Exceptionally preserved phosphatic molds of bundled setal canals inside epipunctae and aditicules, described for the first time in this paper, are direct evidence that these two types of tubular structures of different sizes had the same function of housing sensory setae along the shell margin, but both the setae and the tubules became abandoned in the outer portion of the shell wall through burial by the secondary shell layer when the shell margin migrated forward. Epipunctae have been found so far only in plaesiomyid shells, but aditicules are common in many groups of the order Orthida. The taxonomic value of epipunctae is shown by a reassessment of Pionorthis Schuchert and Cooper, 1932. The hypotype previously regarded widely, but erroneously, as the archetype of ‘Orthis sola’ Billings, 1866, the type species of Pionorthis, is allied to Plaesiomys Hall and Clarke, 1892. It bears the characteristic epipunctae. The holotype of Orthis sola is a dalmanelloid shell with true punctae, assignable to Mendacella Cooper, 1930. This warrants rejection of the genus Pionorthis.

The deep-water brachiopod Dicoelosia King, 1850, from the Early Silurian tropical carbonate shelf of Anticosti Island, eastern Canada

Dicoelosia occurs in two deep water benthic shelly assemblages on an Early Silurian (uppermost Aeronian, Stimulograptus sedgwickii Zone) carbonate ramp to shelf, within the 25 m thick bluish-grey mudstone of the Richardson Member in the middle Jupiter Formation, Anticosti Island, Quebec. Dicoelosia dauphinensis new species is erected on the basis of its relatively large, elongate shell, with a concavo-convex lateral profile, moderately wide, planoconvex lobes and subparallel lateral margins. Dicoelosia dauphinensis first occurs in a Gotatrypa-Dicoelosia Community, in which it constitutes about 7 percent of the individuals within sampled populations. This is replaced about 2 m up section by a Resserella-Dicoelosia Community, where D. dauphinensis makes up 22 percent of the individuals preserved on single bedding planes. Subsequently, Dicoelosia becomes a rare component of the Stegerhynchus-Triplesia Community in a shallowing-upward succession of the upper Richardson Member. It is in this upper Richardson shelly community that Stimulograptus sedgwickii is locally common, together with in situ brachiopod nests of Eocoelia, Triplesia, and Lissatrypa at the upper Jupiter Cliff section. The water depth estimated for the Resserella-Dicoelosia Community is between 100-120 m, below the normal depth range of the Clorinda Community (BA5) on Anticosti, in a distal shelf setting about 80-100 km offshore from the Laurentia paleocontinent, on the west side of the Iapetus Ocean. The interpretation of water depth is based on the facts that, 1) Dicoelosia-rich communities are confined to strata, which lack shallow water sedimentological evidence such as thick calcarenites and hummocky cross stratification, 2) Dicoelosia does not occur with shallower water taxa seen in the overlying Clorinda and Stricklandia communities associated with cyclocrinitid algae, corals and stromatoporoids in the Cybele Member, and 3) Dicoelosia occurs only in the deeper water sections of the Richardson Member, some 10-15 km basinwards from mid-shelf shallower Richardson facies of the Anticosti Basin. Thus, the Dicoelosia-rich communities are interpreted to mark a maximum flooding surface within the distal shelf to ramp Llandovery succession of Anticosti Island.

NEW DATA ON THE FOLIOMENA FAUNA (BRACHIOPODA) FROM THE UPPER ORDOVICIAN OF SOUTH CHINA

Abstract A newly discovered Late Ordovician (early Ashgill) brachiopod assemblage from the Linhsiang Formation in the middle part of the Yangtze Platform, South China, bridges the paleobiogeographical gap between the early Ashgill Foliomena-bearing associations known previously from the upper and the lower parts of the Yangtze Platform. Characterized by minute shells in calcareous to siliciclastic mudstones, the fauna contains 13 brachiopod genera, of which two plectambonitoids are new: Hadroskolos and Jingshanella. Cluster and principal component analyses, based on 29 global occurrences of the Foliomena fauna in Laurentia, Avalonia, Kazakhstan, Baltica, Sardinia, Bohemia, Sibumasu, North China, and South China, revealed broad trends of spatial and temporal faunal differentiation in terms of taxonomic compositions. The analyses demonstrate for the first time that early Foliomena-bearing associations of Caradoc age occupied primarily deepwater (distal shelf) environments with a siliciclastic or calcareous mud substrate. The fauna attained its widest paleogeographical distribution and paleoecological range (midshelf to shelf margin settings) during the early Ashgill.

Parastrophinella (Brachiopoda): Its paleogeographic significance at the Ordovician/Silurian boundary

The type species of Parastrophinella, P. reversa, a pentamerid brachiopod from the uppermost Ordovician (Hirnantian) Ellis Bay Formation of Anticosti Island, Quebec, shows that the genus is characterized by three features: 1) a ventral median septum apically buried in a thickened valve floor, but anteriorly rising above valve floor; 2) prominent alate plates, which are homologous to brachial processes; and, 3) a pseudocruralium consisting of a dorsal median septum, which is largely buried in the valve floor posteriorly, and outer plates that are connected to the median septum via prismatic substance (with poorly developed lamellar layer at the junctions) at, or slightly above, the valve floor. These constitute criteria by which many species previously assigned to Parastrophinella are excluded from the genus. Late Ordovician species that fit the redefinition of Parastrophinella are now confined to eastern North America. The genus crosses the Ordovician/Silurian boundary without major morphologic change, and Early Silurian species of Parastrophinella occur in both eastern North America and Great Britain. In the Late Ordovician, the pentamerid fauna of North America (Laurentia) contrasted sharply with that of Baltica and Kazakhstan, where the Holorhynchus fauna was dominant. Typical elements of the Holorhynchus fauna, such as the large-shelled Holorhynchus and Proconchidium, are known only in the northern parts of Laurentia (Baffin Island, Greenland, and Kolyma), and are absent in the Hudson Bay and Williston basins and southwards.

True Dalmanella and taxonomic implications for some Late Ordovician dalmanellid brachiopods from North America

Examination of topotype material of the type species of Dalmanella, D. testudinaria, from the Hirnantian strata of Borenshult, southern Sweden, revealed several diagnostic characters that distinguish the type species from some common North American species assigned previously to the genus. The typical Dalmanella testudinaria has a consistently developed, primary medial interspace in the dorsal valve, a cardinal process with an invariably bilobed myophore, punctae of two distinctly different sizes, and a lack of aditicules. In contrast, many Late Ordovician species from North America, such as ‘Dalmanella’ meeki and ‘Dalmanella’ multisecta from the Cincinnati type area, consistently have a dorsal medial costa, with a strong tendency to develop aditicules and a trilobate cardinal process. This study confirms the previous notion of many authors that true Dalmanella testudinaria, or the genus as a whole, is either rare or entirely absent in North America. Typical Dalmanella was predominant in cool-water depositional environments (such as the Hirnantia Fauna), whereas the North American dalmanellids (e.g. Cincinnati-type ‘Dalmanella’, Paucicrura, and Diceromyonia) were most abundant and diverse in tropical, epicontinental sea settings. A clear definition of the true Dalmanella, therefore, has important implications for the study of Late Ordovician–Early Silurian brachiopod evolution, paleoecology, and paleobiogeography.

Taxonomic reassessment of two virgianid brachiopod genera from the Upper Ordovician and Lower Silurian of South China

Abstract Reexamination of type and topotype material revealed the presence of well-developed ventral and dorsal interareas in two virgianid brachiopods from South China, Eoconchidium jiangshanensis Liang (in Liu et al., 1983) and Paraconchidium shiqianensis Rong, Xu, and Yang, 1974, of Late Ordovician and Early Silurian ages, respectively. A cladistic analysis of the common virgianid taxa, incorporating new data on the development of interareas, confirms Paraconchidium Rong, Xu, and Yang, 1974 as a valid genus (not a junior synonym of Pseudoconchidium Nikiforova and Sapelnikov, 1971) and warrants E. jiangshanensis as the type species of Deloprosopus new genus (not allied to either Eoconchidium or Tcherskidium as previously believed). Our preliminary survey on the suborder Pentameridina, based on available material or illustrations of well-preserved, disarticulated (typically silicified) valves, indicates that 1) the interareas are more commonly developed in the superfamily Pentameroidea than was reported previously, especially in the families Virgianidae and Subrianidae; 2) the ventral and dorsal interareas commonly do not occur as paired planar surfaces in the Pentameroidea, as they do in the superfamily Stricklandioidea; and 3) despite the common absence of a ventral interarea, the development of a sharply delimited dorsal interarea appears to be ubiquitous in the Pentameroidea and possibly in the suborder Pentameridina. In the currently adopted classification, the presence of matching ventral and dorsal interareas is treated as one of the diagnostic characters that separate Stricklandioidea from other superfamilies of the Pentameridina. The new data presented herein on the development of interareas imply the need to reevaluate the taxonomic and evolutionary significance of the ventral and dorsal interareas, pending a thorough survey on their distribution in the suborder Pentameridina.

The Late Ordovician and Early Silurian pentameride brachiopod Holorhynchus Kiaer, 1902 from North China

Abstract Holorhynchus giganteus Kiaer, 1902, a common Late Ordovician (mid-Ashgill) pentameride brachiopod in the Baltic region, Kazakhstan, and southern Tien Shan, is documented for the first time from the Badanjilin Formation (mid-Ashgill) of western Inner Mongolia (Alxa block), North China. Serial sections of the Chinese material confirm the presence of a vestigial ventral median septum in the early growth stage of H. giganteus, but the septum becomes embedded in the secondary shell thickening at the adult growth stage. A survey of the type material from Norway and additional material from other regions indicates that the incipient ventral median septum is a much more commonly developed structure than was previously believed. The presence of a well-developed pseudodeltidium in the Tien Shan material of H. giganteus and the absence of such a structure in conspecific material from many other regions require a systematic revision of the generic group. Holorhynchus has rodlike crura (=brachial processes) that do not form flanges at their junctions with the inner hinge plates (=outer plates = crural plates) and outer hinge plates (inner plates). This, together with the development of a crude spondylial comb structure, points to its affinity to the Virgianidae rather than to the Stricklandiidae. Holorhynchus can be regarded as a Lazarus taxon because of its absence during the crisis (Hirnantian) and survival (early-middle Rhuddanian) intervals associated with the Late Ordovician mass extinction and its reappearance in Kazakhstan and North China during the Early Silurian (late Rhuddanian-early Aeronian). The mid-Ashgill Holorhynchus fauna, typified by a number of large-shelled pentamerides, was common in the Baltic region, the Urals, Kazakhstan, Tien Shan, Alxa, Qaidam, Kolyma, and east-central Alaska, but largely absent from Laurentia and Siberia (except for Taimyr) in the ancient tropical-subtropical regions. This paleobiogeographic pattern agrees with the general pattern of the Late Ordovician brachiopod provincialism.