Substrate specificity in the Devonian tabulate coral Pleurodictyum

The tabulate coral Pleurodictyum americanum Roemer has been cited as an example of a host-specific organism occurring exclusively on the shells of gastropods, particularly Palaeozygopieura hamiltoniae (Hall). Examination of over 1600 specimens of P. americanum, from the Middle Devonian Hamilton Group of western New York, reveals additional complexities which require reinterpretation. While substrate selectivity for Palaeozygopieura shells is evident in all 42 subsamples, a variety of other substrates were also utilized by Pleurodictyum including corals, brachiopods, other molluscs and pebbles. Recent scleractinian corals inhabiting soft bottoms show similar substrate preference, selecting for the tubes of live serpulids, or gastropod shells (invariably with a secondary sipunculid host), but also occasionally settling on unoccupied shells or pebbles. Shell surfaces of P. hamiltoniae, preserved as external molds on the Pleurodictyum epitheca, exhibit encrustation by worm tubes and bryozoans as well as borings and mechanical shell damage, suggesting that these were not the shells of live gastropods. However, the invariant aperture-downward orientation and the high degree of selectivity of P. americanum strongly suggest that the shells were occupied by secondary hosts. □ Substrate specificity, commen-salism, tabulate coral, gastropod, sipunculid, Devonian, Hamilton Group, New York.     

Substrate specific associations of epebionts on Middle Devonian brachiopods: Implications for paleoecology

A detailed study of over 2500 host brachiopods, from the Middle Devonian Hamilton Group of New York State, revealed distinct patterns of epibiont encrustation, that provide insight into taphonomy and paleoautecology of the host brachiopod shells and depositional environments. The concavo‐convex orthid, Tropidoleptus carinatus (Conrad), as well as strophomenid, and smooth athyrid brachiopods are among the most heavily encrusted. However, terebratulids of nearly identical size and shape are relatively clean of epibionts. This selective distribution strongly suggests that epibionts were discouraged from settling on punctate brachiopods. Brachiopods with small spines and frills were also nearly clean of epibionts, possibly because of entrapment of a mud layer, which made the outer layer of the host inhospitable for larval settling. Concavo‐convex taxa reveal high percent coverage and diversity of epibionts on the convex valve, which probably rested on the substrate during the life of brachiopod. This pattern is observed even on brachiopods that were buried with the convex valve downward. This implies complex post‐mortem histories involving multiple episodes of reorientation and colonization.     

Overgrowth on ammonite conchs: environmental implications for the Lower Toarcian Posidonia Shale

Ammonites of the south–west German Posidonia Shale (Early Toarcian) are occasionally overgrown by bivalves, brachiopods or serpulids. Attachment to both sides of the conchs in some specimens suggests attachment during the ammonite's lifetime, with a pseudoplanktic mode of life for the epizoans. However, dense overgrowth restricted to the upper surface of some ammonite shells indicates post–mortem colonization. Our study shows that apart from oxygen supply as a main factor controlling benthic colonization, substrate consistency also played an important part. Unfavourable living conditions prevailed during deposition of the organic–rich sediments, excluding macrofauna in the benthic environment. However, less than 0.1 per cent of the ammonite conchs found during the excavation show overgrowth, indicating that pseudoplanktonism was an infrequently adopted strategy for inhabiting surface waters.     

Variable pedicle morphology in a population of the Recent brachiopod Terebratulina septentrionalis

Extremely variable pedicle morphology is described in a large sample of the Recent articulate brachiopod Terebratulina septentrionalis (Couthouy) from off the coast of Nova Scotia. The majority of these specimens were attached to the shells of living or dead scaphopods by a dense network of pedicle rootlets. Other brachiopods had been lying on the sea-floor anchored solely by the weight of sediment enmeshed within the bush-like network of pedicle rootlets. Some brachiopod larvae had settled on the exposed pedicle rootlets of adults, presumably because of the scarcity of other suitable substrate. Such a mode of life is thought 'to indicate that these brachiopods could survive a further reduction in the available substrate and it is suggested that, should such conditions persist, changes in hard-part morphology would enable descendants of the species to adopt a predominantly free-lying mode of life. Likely morphological adaptions are suggested, based on examples from the fossil record. This evolutionary trend, from attached to free-lying and perhaps vice versa, has occurred many times in the history of the Phylum Brachiopoda, and it is suggested that the mode of life of this T. septentrionalis population provides an important insight into at least one of the possible evolutionary pathways which bring about such transformations.     

Distribution of pedicle boring traces and the life habit of Late Paleozoic leiorhynchid brachiopods fiom dysoxic habitats

Alexander, R.R. 1994 10 15: Distribution of pedicle boring traces and the life habit of Late Paleozoic leiorhynchid brachiopods from dysoxic habitats.
Pedicle boring traces, Podichnus isp., are concentrated on the anterolateral commissure of both valves of Leiorhynchoidea carboniferum and L. weeksi from the black ferruginous micrites of the Chainman Formation (Upper Carboniferous; west-central Utah, USA) and black phosphatic ('False Cap') limestone of the Phosphoria Formation (southeastern Idaho, USA), respectively. The absence of Podichnus isp. from the posterior of shells of both species indicates that the beak of the shell was buried in the mud, inaccessible to colonization by conspecific larvae. The concentration of pedicle boring traces near the anterolateral incurrent regions of leiorhynchid shells with a well-developed central fold further suggests that settling conspecific larvae behaved rheotaxically. Larvae were induced to metamorphose near the commissure of the host shell, where suspended food was drawn to the incurrents of the host. The piggybacked mode of life on the anterior of conspecific hosts provided a refuge for juveniles above the dysoxic black sediments. In contrast, modem brachiopods that live as epibionts on skeletal substrates display either a random or posteriorly concentrated distribution of pedicle boring traces. Permian, Carboniferous, brachiopods, leiorhynchids, Podichnus, dysoxic, rheotaxis .     

Reasons of micromorphism in modern or fossil brachiopods

Mass samples of the micromorphic recent (Hemithyris psittacea, Eucalathis murrayi, Macandrevia cranium, Megathyris detruncate, Argyrotheca cuneata, Argyrotheca cordata) and fossil Silurian (Microsphaeridiorhynchus sp.) brachiopods were studied. New location of Eucalathis murrayi on the Kitovyi Ridge in the Atlantic Ocean is described. Rates of growth of shells were revealed by the study of the sizes and age of brachiopods. Reasons of dwarfism in brachiopods are discussed. Three strategies of growth of the micromorphic brachiopods are distinguished. Appearance of micromorphic brachiopods is connected with the changes in the rates of shell growth and life expectancy.     

The monophyletic origin of the Brachiopoda

Although it is commonly acepted that the brachiopods descended from phoronid-like ancestors there is dispute over their origin. Traditionally they have been regarded as a monophyletic group, a clade. More recently it has been claimed that brachiopods are polyphyletic and that several of the orders arose independently from separate phoronid-like stocks. The latter point of view implies that brachiopods are not a taxon but merely a grade of organization. Traditional stratophenetic approaches do little to resolve the problem, which may be outside their domain. It is possible, even probable, that the initial radiation involved organisms that lacked mineralized shells. Cladistic analysis of both living forms and Lower Paleozoic taxa strongly supports the contention that brachiopods are monophytetic and closely related to the phoronids. It suggests, however, that the 'inarticulate' Paterinida and Kutorginida are genealogically more closely related to the Articulata than they are to the remaining Inarticulata. □ Brachiopoda, Lophophorata, cladistics, Cambrian.     

Morphogenetic regulation in the shells of bivalves and brachiopods: evidence from the geometry of the spiral

Ackerly. S. C. 1992 07 15: Morphogenetic regulation in the shells of bivalves and brachiopods: evidence from the geometry of the spiral.
Analyses of the spiral geometry in shells of the mollusc Pecten maximus and the brachiopod Terebratulina retusa indicate a relative reduction in morphological variability within the population during growth. The spiral, as measured by a model of exponential radial expansion, tends to converge on a particular adult form, irrespective of irregularities during early growth phases. The recurrence of this pattern of variability in populations from two separate phyla (molluscs and brachiopods) suggests a common mechanism controlling shell form. Brachiopoda. Mollusca, coiling, spiral. morphogenetic regulation, growth .     

Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

A three‐dimensional geometric morphometric study of the development of sulcus versus shell outline in Permian neospiriferine brachiopods

The morphological variation of the sulcal development and shell outline in large Permian neospiriferine brachiopods including Fasciculatia Waterhouse, 2004 is investigated using geometric morphometrics. The sulcal tongues of spiriferide brachiopods can be, in a qualitative sense, categorized into three types according to the degree of their development: short sulcal tongue, long sulcal tongue and geniculated sulcal tongue. All three types have been noted within Fasciculatia striatoparadoxa, regardless of the nature of the substrate which they originally inhabited. To quantify its morphological variation both in sulcal development and shell outline, 51 brachiopod shells were scanned with a three‐dimensional (3‐D) surface imaging device, and their 3‐D models were reconstructed. Using two landmarks and 58 semilandmarks designated on the surface of the reconstructed 3‐D models, a landmark‐based morphometric analysis was performed. Our result demonstrates a significant intraspecific variation of sulcal development in F. striatoparadoxa and its relatives. Local environmental factors, especially the intensity of ambient water flow, are invoked as the most likely cause for this intraspecific variation. Additionally, this study also shows that there are considerable interspecific distinctions in shell outline among Fasciculatia species, independent of the high variation in the sulcal development. The strong stability of overall shell outline at species level implies a decoupled morphological development between sulcal tongue and whole shell outline. The 3‐D morphometric approach applied here demonstrates its great utility as a tool for quantifying and analysing the morphological variation of highly convex brachiopod shells.     

Determination of paleoseasonality of fossil brachiopods using shell spiral deviations and chemical proxies

Brachiopods have been extensively used in paleoclimatic and paleoecological reconstructions, but their utility would greatly increase if paleoseasonality information could be obtained from their shells. Determining seasonal seawater temperature variations from fossil brachiopods requires knowledge of specimen ontogenetic ages, which is difficult to determine compared to other organisms secreting a shell by accretion. In this study, the combination of the spiral deviation methodology and chemical proxies is tested for determining specimen ontogenetic ages and paleoseasonality using two species of fossil brachiopods, Laqueus rubellus and Terebratula terebratula, of Pleistocene and Late Miocene age, respectively. Spiral deviations were obtained for Laqueus and Terebratula using an R program developed for modern taxa, and well-preserved shells were analyzed using oxygen isotopes and Mg/Ca ratios as chemical proxies for past seawater temperature. Results reveal that locations of spiral deviations on shells of L. rubellus displayed a strong direct relationship with Mg concentrations, and resulting Mg/Ca-derived paleotemperatures were seasonal. Conversely, specimens of T. terebratula did not show a consistently strong relationship between Mg concentrations and spiral deviations, although resulting paleotemperatures agreed with those from previous studies. Overall, the results from this study indicate that the spiral deviation methodology combined with chemical proxies presents great potential for utility in past seasonal seawater temperature reconstructions in pristinely preserved, biconvex fossil 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.     

Epizoans on late Ordovician brachiopods from Southeastern Indiana

The frequency of epizoans (cornulitids, inarticulate brachiopods, bryozoans, solitary and colonial rugosan corals) on over 8000 specimens of articulate brachiopods (four strophomenids, five orthids, one rhynchonellid) was calculated for four stratigraphic horizons in the Dillsboro Formation of southeastern Indiana. Frequency of shells encrusted correlates significantly with the surface area of the valves. Punctae in brachiopod shells (Onniella meeki) may have deterred larval settlement of epizoans. Coarse ribbing on articulates deterred encrustation by the inarticulate brachiopod. The horn coral shows a preference for attachment to the anterior of Hiscobeccus capax. Bryozoans show a preference for the incurrent lateral margins of inferred living hosts, suggesting rheotropic behavior by settling larvae. Inarticulate brachiopods are concentrated around the sloping commissure of the brachial valve of strophomenids, suggesting geotropic behavior and/or selective survival of settling larvae. Inarticulates deterred overgrowth by bryozoans. High frequencies of encrustations of the medial region of pedicle valves of orthids and strophomenids probably reflect post‐mortem encrustations. Alternating intervals of slow sediment accumulation punctuated by tropical storms and rapid shell burial may account for the high frequency of shells with either their entire surface veneered or only a very small area encrusted by bryozoans.     

Growth rate and substrate-related mortality of a benthic brachiopod population

Collins, M. 1991 01 15: Growth rate and substrate-related mortality of a benthic brachiopod population. Lethaia , Vol. 24, pp. 1–11. Oslo. ISSN 0024–1164.
Vital staining and careful examination of potential substrates enabled an accurate census of Terebratulina retusa to be made and prompted a study of their population dynamics. Seasonal samples of T. retusa from a deep water Modiolus -brachiopod assemblage were always dominated by small (>1 mm) individuals as growth rate of the post-larvae, estimated from changes in mean cohort length. was unexpectedly slow. Six months after settlement the animals had barely doubled in length, a rate of increase consistent with laboratory studies, but an order of magnitude less than conventional estimates. As the attainment of a size refuge is the only documented strategy by which articulate brachiopods counter overgrowth or disturbance this observation has profound implications for survival. Mortality of T. retusa in the Firth of Lorn, from different substrates, followed an unexpected pattern. Virtually the only substrate on which adult T. rehusa were recorded was the surface of M. modiolus shells, although juvenile T. retusa attached to this substrate suffered enhanced levels of mortality. Grazing pressures and spatial competition, believed to be reduced on complex surfaces, may account for the elevated mortality levels of M. modiolus-attached post-larvae prior to the apparent size refuge at a length of 2 mm. * Brachiopoda, Terebratulina, growth rate, disturbance, ecology, population structure .     

Internal structure and paleoecology of the lower Permian Uzunbulak reef complex of the Tarim Basin,Northwest China

Summary The internal construction and biotic communities of the Uzunbulak reef of the northwestern Tarim Basin are studied for the first time. The reef was built during the Sakmarian, while the reef substrate and capping beds are of latest Asselian and earliest Artinskian ages, respectively. The reef substrate beds are composed of skeletal and oncoid grainstone. Those fusulinid-dominated skeletal shoals and oncoid banks indicate a high-energy environment and produced local topographic highs on which the reef grew. Reef framework consists mainly of calcisponge bafflestone, calcisponge-Thartharella framestone, and Tubiphytes, Archaeolithoporella and Girvanella boundstones. Calcisponges were the primary frameconstructors that baffled high-energy currents. Archaeolithoporella, Tubiphytes, Girvanella and possibly microbes acted as the primary binders for the boundstone framework. Fusulinids and brachiopods were common reef dwellers. The interreef facies sediments are composed of skeletal-crinoid wackestone-packstone. Most of bioclasts have thick, micritized envelopes. The back-reef facies deposits consist of alternating skeletal packstone to wackestone and black shale. Sea-level fluctuations were probably accountable for the reef growth and demise. Of the reefal dwellers, brachiopods are extraordinarily abundant in Uzunbulak. They are assignable to five distinctive associations, one each from the reef substrate, framework and inter-reef facies, respectively, and two from the reef capping facies. The brachiopods in the substrate beds were mostly attached to hard substrates by a pedicle, while a few species rested on soft substrates by support of halteroid spines. Cementation of the ventral valve on hard substrates characterizes attachment of the reef framework brachiopods. All inter-reef species were anchored into the substratum comprising hard material by a strong pedicle. Back-reef brachiopods dominantly rested on the soft substrates by support of halteroid spines. the framework brachiopods had the strongest wave-resistant capability;those from both substrate and inter-reef facies were moderately capable of withstanding agitation; and the backreef species preferred to live in calmwater, organic-rich muddy environments.     

Study of fossil and recent brachiopods,using a skyscan 1172 X-ray microtomograph

Fossil and recent brachiopods were studied with the aid of a Skyscan 1172 microtomograph. The capabilities of this method at different stages of studying, X-ray scanning and producing slices and 3D models are described. The method enables the study of punctuation, microornamentation, and inner structures of the brachiopod shells and soft tissues. The contrast of shell structures of fossil brachiopods is discussed; it depends on differences in the mineral composition of the shell and surrounding matter. This method allows studying the inner structure of the holotypes of brachiopod species without damaging their shells. The data on the efficiency of the method are provided.     

In search of the microbe/mineral interface: quantitative analysis of bacteria on metal surfaces using vertical scanning interferometry

To understand the development of biofilms on metal surfaces, analysis of initial bacterial attachment to surfaces is crucial. Here we present the results of a study, using Shewanella oneidensis MR-1 as a model organism, in which vertical scanning interferometry (VSI) was used to investigate the initial stages of cell attachment to glass, steel and aluminium surfaces. It was found that while VSI gave unambiguous results with opaque surfaces, when reflective surfaces were used, an artifact sometimes appeared, with the bacteria appearing as rod-shaped pits rather than as cells on the surface. When the bacteria were altered to increase opacity, this artifact disappeared, and upon further investigation, it was found that the observational artifact was the result of a conflict between light reflected from the bacteria and the light reflected from the bacteria–metal interface. These results suggest that not only can bacteria be measured on surfaces using VSI, but with some modifications to the analytical software, there may be a unique window for studying the bacterial/substrate interface that can be used for quantitative observations. Imaging and characterization of the bacteria–substrate interface in vivo (previously invisible) will provide new insights into the interactions that occur at this important juncture.     

Marine benthic communities in the Upper Devonian of New York

Four distinctive benthic communities occur in the mid-shelf deltaic deposits of the Upper Devonian Sonyea Group: the Bellerophon Community, dominated by herbivorous gastropods and deposit-feeding bivalves; the Prodrtctella Community, dominated by brachiopods with adaptations for life on muddy substrata; the Cypricardella Community, dominated by bivalves and brachiopods with a variety of adaptations; and the Rhipidomella Community, dominated by brachiopods adapted for life on sandy substrata. The geographic and stratigraphic occurrence of the communities suggest that their distribution was controlled by two principal sets of environmental variables: (1) onshore to offshore gradients in salinity, dissolved oxygen concentrations, and sedimentary substrata, and (2) rate of delta progradation. A change in the rate of delta build-out during mid-Sonyea time led to a dramatic change in onshore to offshore community patterns. The change suggests that great care must be used in constructing generalized community models for longer intervals of Paleozoic time.     

THE COMPOSITION AND STRUCTURE OF LOWER SILURIAN MARINE COMMUNITIES

Five benthic communities occupied the shelf regions of the British Isles, Norway, and North America in Upper Llandovery times. The communities are listed below in order of increasing distance from shore.

• 1 The Lingula Community is the least diverse; it has both infaunal elements, including a protobranch, and two lingulids, and epifaunal elements, including a rhynchonellid, a pterioid, and a cornulitid. A restricted and protected near-shore environment, such as a bay or estuary, is postulated.
• 2 The Eocoelia Community shares elements in common with the former community, but is more diverse and is dominated by epifaunal forms; the many small pedunculate brachiopods probably lived attached to the large leptostrophiid brachiopod.
• 3 The Pentamerus Community is dominated by this genus which lived free and upright on the bottom; smaller pedunculate brachiopods probably attached to this large neighbor.
• 4 The Costistricklandia Community was similar in structure to the former community with the many small pedunculate brachiopods being attached to the large Costistricklandia.
• 5 The Clorinda Community is the most diverse, with a great variety of small brachiopods which were probably able to attach to small objects in this quiet off-shore environment, or to some moderately sized brachiopods, such as Clorinda and Cyrtia, which apparently lived free on the bottom.

The brachiopod dominated communities of the Silurian clearly inhabited the ‘level bottom’, an area now occupied mainly by infaunal forms. The main attachment surfaces for the epifaunal elements of the Silurian communities were disarticulated, convex-upward shells.     

Functional morphology of chonetidine (Brachiopoda) spines: biomechanical tests of a potential key innovation

Chonetidine brachiopods were a significant component of Paleozoic marine life; chonetidines experienced a major adaptive radiation during the Late Silurian and Early Devonian. The Chonetidina clade is united by the presence of spines on the hingeline but the function of these spines has not been clearly demonstrated. The present study performs a biomechanical experiment, using specimens of Neochonetes granulifera with and without artificial spines in a recirculating flume, to test if the spines may have inhibited entrainment in higher energy settings. Specimens with spines were less likely to be overturned or transported than those without spines, and were overturned or transported at significantly higher velocities than specimens without spines. Performance improved with the addition of another pair of spines. In addition, spinose specimens reoriented such that their commissures faced upstream. The results suggest that chonetidines could have survived in higher-energy environments, even if the brachiopods were not physically attached to the substrate. As such, it is functionally plausible that the evolution of hinge-spines may have possibly facilitated the adaptive radiation of the clade by enabling chonetidines to inhabit previously unoccupied habitats.