Euryhalinity of Palaeozoic articulate brachiopods

OMonotypic and very low diversity virgianid shell beds from the Upper Ordovician to Lower Silurian dolomites of North Greenland were formed in marginal marine quiet-water hypersaline environments. In the light of this evidence the salinity tolerances of other Palaeozoic articulate brachiopods is evaluated. There are only a small number of species apparently invading hypersaline or brackish environments, but it is significant considering that previously all articulate brachiopods were thought to be fully marine. Two types of occurrence are noted, those species specifically related to marginally marine environments, disappearing with the introduction of fully marine faunas, and the majority of species which extend their normal marine range into marginal conditions. No brachiopod species appears to have invaded very hypersaline or truly brackish conditions. No single group of articulate brachiopods specifically specialised in colonising marginal marine environments, apart from possibly the virgianid pentamerids. Palaeozoic, Upper Ordovician, Lower Silurian, Brachiopoda. Pentamerida, Virgianidae, Greenland, palaeoecology. hypersaline environments, brackish environments .     

Revision of the Lower Cambrian brachiopod Heliomedusa Sun & Hou, 1987

Jin Yugan & Wang Huayu 1992 01 15: Revision of the Lower Cambrian brachiopod Heliomedusa Sun & Hou, 1987. Lethaia , voi. 25. pp. 35–49. Oslo. ISSN 0024–1164.
Fossils referred to Heliomedusa , from the Chiungchussu Formation of Early Cambrian age in Yunnan. are extraordinarily preserved. They were previously considered to represent jellyfish. and have been suspected more recently of being an obolellid brachiopod. A study of the morphology of this genus indicates that it is related closely lo the non-pediculate craniopsids. which occurred from Early Ordovician to Carboniferous time. The soft parts of this brachiopod. such as setae and nerves, which usually are not preserved in fossils also are described. Brachiopoda, Inarticulata, Craniapsidue, anatomy, setae, nervous system, mantle canals, Early Cambrian .     

BILLENGSELLIDE AND ORTHIDE BRACHIOPODS: NEW INSIGHTS INTO EARLIEST ORDOVICIAN EVOLUTION AND BIOGEOGRAPHY FROM NORTHERN IRAN

Abstract:  The eastern Alborz Mountains of Iran comprise a significant peri-Gondwanan terrane relevant to the early evolution of late Cambrian – early Ordovician brachiopods incorporated into the emerging benthic biota of the Paleozoic Evolutionary Fauna. A low diversity brachiopod assemblage from the late Tremadocian unit of the Lashkarak Formation contains six new species including the polytoechioideans Polytoechia and Protambonites and the orthoideans Paralenorthis, Ranorthis, Tarfaya and Xianorthis . The fauna preserves the earliest records of Polytoechia , unknown previously outside Laurentia and the Uralian margin of Baltica, and of Paralenorthis and Ranorthis , which were widespread along Gondwanan margins and in Baltica from the Floian (Arenig), plus Xianorthis , known hitherto only from the Floian of South China. The enigmatic Tarfaya has an impunctate shell fabric and setigerous perforations along the posterior margin, indicating placement within the Orthoidea in a new Family Tarfayidae. New species of Polytoechia , Protambonites , Paralenorthis, Ranorthis , Tarfaya , Xianorthis are described.     

THE ENIGMATIC EARLY CAMBRIAN SALANYGOLINA– A STEM GROUP OF RHYNCHONELLIFORM CHILEATE BRACHIOPODS?

Abstract:  New material of the enigmatic brachiopod Salanygolina obliqua Ushatinskaya from the Early Cambrian of Mongolia shows that it has a colleplax – a triangular plate – in the umbonal perforation, which is enlarged by resorption. This structure is otherwise only known from the equally enigmatic Palaeozoic orders Chileida and Dictyonellida (Rhynchonelliformea, Chileata). The colleplax in Salanygolina is here considered to be homologous with that of the chileates. Salanygolina is also provided with a ridge-like pseudodeltidium, which is another chileate feature. Other characters of Salanygolina , like the radial arrangement of adductor muscle scars and postero-medially placed internal oblique muscles are characteristic of chileates, but also found in the paterinates. In contrast, mixoperipheral dorsal valves with low rudimentary interareas are well known in paterinates, but not yet recorded from chileates. Thus, Salanygolina shows a mosaic combination of morphologic characters, known both from the paterinates and chileates, indicating that it may represent a stem group of the rhynchonelliform chileate brachiopods. The laminar phosphatic secondary shell of Salanygolina is composed of closely packed and nearly identical hexagonal prisms, oriented with their long axis normal to the laminae in a honeycomb pattern. The prism walls appear to have originally been composed of organic membranes and might represent precursors of the organic sheaths of calcite fibers that are typical of calcitic shells with a fibrous microstructure.     

The elkaniide brachiopodVolborthia from the lower ordovician of Baltoscandia

The rare, enigmatic brachiopodVolborthia recurva (Kutorga) is redescribed based on new, well preserved collections from the Lower Ordovician (lower Llanvirn; Oeland Series; Kunda Stage) of Ingria (St. Petersburg district); it is also recorded for the first time from Sweden, where it occurs in the Kunda Stage on the Island of Oland.V. recurva has usually been classified questionably within the order Paterinida, but the shell structure, micro-ornamentation, muscle scars, and presence of a lingulid-type pedicle groove now indicate that it is an aberrant member of the lingulid family Elkaniidae; it is the only known lingulid with a high conical dorsal valve characterised by mixoperipheral growth.     

A NOTE ON THE SHELL STRUCTURE OF THE TRIPLESIACEAN BRACHIOPODS

The triplesiacean shell has long been accepted as being impunctate, and has recently (Williams 1968) been shown to be of the laminar type which characterises the non-plectambonitacean strophomenids. Examination of a specimen of the triplesiid Oxoplecia , however, shows a clear development of pseudopuncta; and what appears to be a differentiated primary layer external to the laminar shell in this specimen is confirmed from the shell of another Oxoplecia species.
The presence of rarely developed pseudopuncta within the superfamily supports the association of the Triplesiacea with the Davidsoniacea by Williams (1970), while the establishment of a primary layer in the Triplesiacea confirms his expectation based on its presence in the presumed billingsellacean ancestors.     

First report of brachiopod–brachiopod endoparasitism

Schemm-Gregory, M. & Sutton, M. 2010: First report of brachiopod–brachiopod endoparasitism. Lethaia , Vol. 43, pp. 111–115.
The first example of brachiopod–brachiopod endoparasitism is reported from the Lower Devonian of China. Three-dimensional reconstructions following serial sectioning show a specimen of a strophomenide ( Dicoelostrophia sp.) within an articulated shell of the spiriferide Rostrospirifer tonkinensis ; morphological modifications of the host and the positioning of the strophomenide with respect to the inhalant current demonstrate the in vivo nature of the association. The symbiosis is interpreted as parasitic; it appears to be facultative but demonstrates the viability of this mode of life, expanding the ecological range known to be exploitable by the Brachiopoda. □ 3-D reconstruction , Brachiopoda , Dicoelostrophia , endoparasitism , Lower Devonian , Rostrospirifer .     

Size, shape and the distribution of organic matter in the Recent Antarctic brachiopod Liothyrella uva

The living terebratulid articulate brachiopod Liothyrella uva (Jackson 1912) was sampled from a shallow water population at Signy Island, South Orkney Islands, Antarctica. Neither shell height nor shell breadth were directly proportional to length and as a result there was a change in shell shape with size (and hence age); this change was small but statistically significant. The proportion of the total organic matter found in the shell and internal (mantle) tissues also changed with size. In small (5–7 mm length) brachiopods 70–80% of the total organic matter was located within the shell; this fraction decreased with increasing size until above about 25 mm length the proportion of organic matter in the shell was constant at 30–45%. Variability in this measure was influenced by infection with endolithic red algae. Punctal density was independent of size with a mean value of 95.7 per mm² [SE (standard error) ± 2.2], which was greater than in populations sampled from higher latitudes. With increasing shell length there was a slight increase in the size of puncta close to the shell edge. In all morphometric measures the range of variation observed was similar to that described from populations of Liothyrella from other areas of the Southern Ocean.     

Structure of the first-formed shell of the Middle Ordovician orthid-like brachiopods from the Leningrad Region

Shell structure of the first-formed shell of the Middle Ordovician orthid-like brachiopods from the Leningrad Region is described. The 190-μm-wide first-formed shell is composed of finely granular layer while 700-μm-wide first-formed shell is fibrous. Thus the order Orthida in the Early Paleozoic included brachiopods with both planktotrophic and lecithotrophic larvae in the ontogeny.     

The oldest Lingulids of the Siberian Platform: Microornamentation and shell structure

New brachiopod genus of the order Lingulida Sukharilingula with new type species S. luchininae is described from the Lower Botomian (Lower Cambrian) of northwestern Siberian Platform, basin of the Yenisei River, section on the Sukharikha River. The study of the microornamentation and shell structure of the new genus revealed that the initial shell structure of lingulid shells could transform in the course of early diagenesis. The importance of reconstruction of burial conditions of the valves for the reconstruction of posthumous transformations is discussed.     

Molecular phylogeny of brachiopods and phoronids based on nuclear-encoded small subunit ribosomal RNA gene sequences

Brachiopod and phoronid phylogeny is inferred from SSU rDNA sequences of 28 articulate and nine inarticulate brachiopods, three phoronids, two ectoprocts and various outgroups, using gene trees reconstructed by weighted parsimony, distance and maximum likelihood methods. Of these sequences, 33 from brachiopods, two from phoronids and one each from an ectoproct and a priapulan are newly determined. The brachiopod sequences belong to 31 different genera and thus survey about 10% of extant genus-level diversity. Sequences determined in different laboratories and those from closely related taxa agree well, but evidence is presented suggesting that one published phoronid sequence (GenBank accession UO12648) is a brachiopod-phoronid chimaera, and this sequence is excluded from the analyses. The chiton, Acanthopleura, is identified as the phenetically proximal outgroup; other selected outgroups were chosen to allow comparison with recent, non-molecular analyses of brachiopod phylogeny. The different outgroups and methods of phylogenetic reconstruction lead to similar results, with differences mainly in the resolution of weakly supported ancient and recent nodes, including the divergence of inarticulate brachiopod sub-phyla, the position of the rhynchonellids in relation to long- and short-looped articulate brachiopod clades and the relationships of some articulate brachiopod genera and species. Attention is drawn to the problem presented by nodes that are strongly supported by non-molecular evidence but receive only low bootstrap resampling support. Overall, the gene trees agree with morphology-based brachiopod taxonomy, but novel relationships are tentatively suggested for thecideidine and megathyrid brachiopods. Articulate brachiopods are found to be monophyletic in all reconstructions, but monophyly of inarticulate brachiopods and the possible inclusion of phoronids in the inarticulate brachiopod clade are less strongly established. Phoronids are clearly excluded from a sister-group relationship with articulate brachiopods, this proposed relationship being due to the rejected, chimaeric sequence (GenBank UO12648). Lineage relative rate tests show no heterogeneity of evolutionary rate among articulate brachiopod sequences, but indicate that inarticulate brachiopod plus phoronid sequences evolve somewhat more slowly. Both brachiopods and phoronids evolve slowly by comparison with other invertebrates. A number of palaeontologically dated times of earliest appearance are used to make upper and lower estimates of the global rate of brachiopod SSU rDNA evolution, and these estimates are used to infer the likely divergence times of other nodes in the gene tree. There is reasonable agreement between most inferred molecular and palaeontological ages. The estimated rates of SSU rDNA sequence evolution suggest that the last common ancestor of brachiopods, chitons and other protostome invertebrates (Lophotrochozoa and Ecdysozoa) lived deep in Precambrian time. Results of this first DNA-based, taxonomically representative analysis of brachiopod phylogeny are in broad agreement with current morphology-based classification and systematics and are largely consistent with the hypothesis that brachiopod shell ontogeny and morphology are a good guide to phylogeny.     

Shell structure of <Emphasis Type="Italic">Kutorgina</Emphasis> Billings (Brachiopoda,Kutorginida)

Shell structure in members of the genus Kutorgina, Lower Cambrian brachiopods with calcareous shells and primitive articulation, is described for the first time. This type of shell structure resembles that of the Lower Cambrian mollusks and hyoliths rather than of brachiopods.     

Brachiopod shell thickness links environment and evolution

While it is well established that the shapes and sizes of shells are strongly phylogenetically controlled, little is known about the phylogenetic constraints on shell thickness. Yet, shell thickness is likely to be sensitive to environmental fluctuations and has the potential to illuminate environmental perturbations through deep time. Here we systematically quantify the thickness of the anterior brachiopod shell which protects the filtration chamber and is thus considered functionally homologous across higher taxa of brachiopods. Our data come from 66 genera and 10 different orders and shows well-defined upper and lower boundaries of anterior shell thickness. For Ordovician and Silurian brachiopods we find significant order-level differences and a trend of increasing shell thickness with water depth. Modern (Cenozoic) brachiopods, by comparison, fall into the lower half of observed shell thicknesses. Among Ordovician–Silurian brachiopods, older stocks commonly have thicker shells, and thick-shelled taxa contributed more prominently to the Great Ordovician Biodiversification but suffered more severely during the Late Ordovician Mass Extinction. Our data highlight a significant reduction in maximum and minimum shell thickness following the Late Ordovician mass extinction. This points towards stronger selection pressure for energy-efficient shell secretion during times of crisis.     

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.     

The mesolophe, a new lophophore type for chonetacean brachiopods

Rachebaeuf, P. R. & Copper, P. 1990 10 15: The mesolophe, a new lophophore type for chonetacean brachiopods. Lethaia , Val. 23, pp. 341–346. Oslo. ISSN 0024–1164.
Following a summary of previous lophophore reconstructions for the chonetaceans, we describe an unusual pyritized structure within the calcite infill of an exceptionally preserved shell of Archeochonetes primigenius (Twenhofel) from the Late Ordovician (Ashgill) of Anticosti Island, Quebec, Canada. The brachial valve interior of most Lower Devonian to Permian chonetaceans shows the development of three depressed deepened areas (gutters) in the valve floor. The disposition of these gutters coincides remarkably with the shape of the pyritized structure, which we postulate as a new type of lophophore, the mesolophe. ▭ Brachiopoda, Chonetacea, functional morphology, lophophore .     

The brachiopod fold: a neglected body plan hypothesis

Attention is drawn to Nielsen's radical body plan concept, here named the 'brachiopod fold hypothesis', under which brachiopods and phoronids are recognized to be transversely folded across the ontogenetic anterior–posterior axis so that, to make useful comparisons with other phyla, these organisms must be conceptually unfolded. Under the hypothesis brachiopod brachial and pedicle shell valves are respectively 'anterior' and 'posterior' rather than 'dorsal' and 'ventral' as traditionally described. The hypothesis makes sense of the symmetry axes of the brachiopod shell, is consistent with various indications from fossil and Recent brachiopods, and gives rise to predicted patterns of axis–determining gene expression that differ from those obtaining under the traditional view of the body plan, whilst the variety of folding movements in different lineages implies that superficially dissimilar morphogenetic folds may be fundamentally homologous. Convergent folding patterns are noted in some other organisms. A previous conjecture that inarticulate linguloid brachiopods were derived from halkieriid–like ancestors is elaborated with proposals that recognize possible functional continuities of coelomic and marginal sclerite functions, and it is noted that an ancestrally facultative fold could have become incorporated by genetic assimilation into the brachiopod developmental program. An experimental approach is outlined to test the possibility that some members of the 'small shelly fauna' may have been members of the halkieriid–like brachiopod stem lineage and it is also suggested that buoyancy modification may have been an important function of mineralization amongst Lower Cambrian floaters and swimmers, since negative buoyancy would facilitate access to the benthic niche.     

ORIGIN OF LAMINAR-SHELLED ARTICULATE BRACHIOPODS

The calcareous shell of Billingsella is composed of a primary layer of crystallites commonly disposed vertically or at high angles to the shell surface, and a laminar secondary layer of flat-lying blades which usually amalgamate laterally to form a succession of plates. The succession is unlike that of other billingsellaceans such as Nisusia or contemporaneous orthaceans like Oligomys and Orusia which have a secondary layer composed of fibres. It is, however, closely comparable with the impunctate, laminar secondary layer of the Triplesiacea and early Davidson-iacea. The other pene-contemporaneous laminar-shelled articulates, the Stropho-menacea and Plectambonitacea, differ in being pseudopunctate. These differences in shell structure suggest that the Strophomenida were polyphyletically derived, mainly from the Nisusiidae but also from the Billingsellidae which gave rise to the Davidsoniacea (and Triplesiacea).     

Middle Ordovician Aporthophyla brachiopod fauna from the roof of the World,southern Tibet

A Darriwilian (late Middle Ordovician) brachiopod fauna from the Lower Formation of the Chiatsun Group at Jiacun, northern Nyalam, southern Tibet, consists of ten brachiopod species, forming a distinct Aporthophyla–Paralenorthis Association. Its taxonomic composition is typical of the Aporthophyla Fauna that occupied lower BA2 to upper BA3 benthic environments on sandy lime mud substrates. The occurrence of Paralenorthis in southern Tibet is confirmed for the first time, represented by P. costata sp. nov. Numerical analyses (PCA and CA) of 18 Darriwilian brachiopod faunas from ten palaeoplates or terranes indicate that: (1) the Aporthophyla Fauna was confined to a specific latitudinal belt although it had a wide lateral distribution from the large palaeocontinents of Gondwana to Laurentia; (2) the Saucrorthis Fauna, a typical late Middle Ordovician regional fauna, is limited to a much smaller area, marginal to the Gondwana supercontinent; (3) the strong provincialism persistent in the late Middle Ordovician contributed to increased gamma biodiversity during the Great Ordovician Biodiversification Event.