Brachiopod morphology,life mode,and crushing predation in the Ordovician Arnheim Formation,Kentucky, USA

Predation on ancient shelled prey is an often-studied topic in paleoecology, but the early Paleozoic and the brachiopods that dominated the seafloor at that time are relatively underrepresented in the predation literature. We assessed predatory repair scar frequencies among the brachiopod genera from the Early Richmondian (Late Ordovician) Oregonia Member (Arnheim Formation) near Flemingsburg, Kentucky. We found higher repair frequencies on the concavo-convex Rafinesquina and Leptaena relative to the bi-convex genera. There were no trends in repair frequency through the stratigraphic section and no relationships between repair frequency and community diversity metrics. It is possible that concavo-convex brachiopods’ flat shape, thin shell profile, and free-lying (no pedicle attachment) lifestyle made them more likely or appealing targets of Ordovician crushing predators. It is also possible that concavo-convex brachiopods were better suited to survive crushing attacks than biconvex taxa. We also found differences in shell ornament that may influence the visibility of repair scars.     

The functional morphology and palaeoecology of the Dinantian brachiopod Levitusia

The functional morphology of Levitusia , based principally upon the type-species L. humerosa (J. Sowerby), is described and compared with living brachiopods. It is concluded that Levitusia species lived in tropical Dinantian seas with agitated waters and at shallow depths, perhaps no more than 50 m. The lithologies in which specimens are found support this view and indicate the need for some revision in our interpretations of some Dinantian sedimentary environments. Further evidence for the synonymy of L. christiani (de Koninck) in L. humerosa is presented.     

The late Sandbian – earliest Katian (Ordovician) brachiopod immigration and its influence on the brachiopod fauna in the Oslo Region, Norway

Bulk samples of brachiopods from the middle and upper parts of the Arnestad Formation and the entire Frognerkilen Formation in the Oslo Region demonstrate major vertical changes in the Sandbian to Katian amphicratonic fauna of this part of the Baltic Province. The main influx of new taxa occurred in the upper part of the Arnestad Formation (uppermost Sandbian), and in the succeeding Frognerkilen Formation (basal Katian). Faunal change is initially reflected in an increase in diversity and not as a replacement of the previously incumbent genera, which remain throughout the studied sequence. The majority of the new genera migrated from the East Baltic Region or through this region from Avalonia, while the marginal Laurentian taxa first occupied the Scandinavian part of Baltica. Six taxa migrated from Laurentia, three from Avalonia, two from Gondwana and one apparently originated on the South China palaeoplate. The faunal shifts and immigration of brachiopods appear to have been in response to an initial transgression together with the movement of Baltica into more warm temperate latitudes, but the diverse faunas developed both against a background of subsequent regression and in response to an offshore shift of biofacies. The new amphicratonic fauna was thus generated by faunal shifts from elsewhere in the Baltic Province to the marginal environments of the Oslo Region together with more exotic elements from neighbouring continents and microcontinents.     

Microwear and morphology: functional relationships between human dental microwear and the mandible

Microscopic pits and scratches form on teeth during chewing, but the extent to which their formation is influenced by mandibular morphology is unknown. Digitized micrographs of the base of facet nine of the first, second, and third mandibular molar were used to record microwear features from an archaeological sample of modern humans recovered from Semna South in northern Sudan (n=38; 100 BC to AD 350). Microwear patterns of the molar row are correlated with mandibular corpus width and depth, and with mandibular length. Variations in shear and compression at the base of facet nine during chewing were inferred. It may be that some correlations between microwear and mandibular morphology are predictable, reflecting similar aspects of masticatory loading, though the full extent of the relationship remains to be resolved.     

Functional adaptation of spiriferide brachiopod morphology

It has been suggested that spiriferide morphologies have evolved to adapt to a variety of environmental conditions. Through a computational fluid dynamics approach, we examined how the spiriferide original form was optimized for a lotic condition, specifically addressing the functionalization of the Devonian spiriferide brachiopod Paraspirifer bownockeri to generate passive feeding flows. The results using four models, each of which differed in the development of the spiriferide shell depression, i.e. sulcus, showed that a deeper sulcus functions to create strong spiral flows so as to align on the feeding organ inside the shell. Among the sulcus‐developed models, only the mimic of the natural form could generate comparative slow flows with a stable inflow area. The fossil record of spiriferides shows a morphological trade‐off between the development of the sulcus and wing form. We concluded that spiriferide shells with such a morphological combination evolved to produce various feeding strategies, resulting in diversification.     

Morphology,ontogeny and affinities of the Hirnantian triplisiid brachiopod Streptis undifera from Baltoscandia

New silicified topotypic material of the Upper Ordovician Streptis undifera (Schmidt, 1858) from the stratotype of the Porkuni Regional Stage in Estonia provides important data on triplesiid morphology and ontogeny, which has substantial implications for our understanding of the affinity of this group of brachiopods. In particular, the new material shows that the early ontogeny of Streptis includes evidence for a cicatrix attachment and colleplax‐like structure in the ventral valve. It is likely also that a lecithotrophic feeding habit evolved in triplesiidines sometime in the Ordovician, which sets them apart from all other known strophomenates. A neotype is selected among the specimens of Streptis undifera from the type locality in Porkuni. In Baltoscandia, Streptis undifera appears first in Norway in late Katian below the strata with the pentamerid brachiopod Holorhynchus. In Estonia, Streptis undifera is an index species of the early Hirnantian and occurs in association with stromatoporoid‐coral reefs and related inter‐reef deposits, which overly the strata with Holorhynchus. In Baltoscandia, the distribution areas of Streptis and Holorhynchus are more or less identical in spite of some differences in age.     

The fish tail as a derivation from axial musculoskeletal anatomy: an integrative analysis of functional morphology

The adult morphology of the tail varies greatly among extant fishes despite sharing both ontogenetic similarities and the functional need to propel the body through a fluid medium. Both sharks (Chondrichthyes) and ray-finned fishes (Actinopterygii) control caudal fin musculature independently of axial body myomere activity to modify the stiffness and shape of their tails. For example, sharks and bony fishes possess different structural elements and muscles and move their tails in different ways, resulting in different locomotory hydrodynamic effects and a range of performance variables including speed and maneuverability. The stiffness of the heterocercal, lobate tail of the shark can be modulated during the tail beat resulting in nearly continuous thrust production. In contrast, the highly flexible tail of ray-finned fishes can be manipulated into many different shape conformations enabling increased maneuverability for these fishes. Consequently, the developmental, morphological, and functional derivation of the tail from the axial trunk has resulted in a diversity of form, the attributes of which may be of ecological and evolutionary significance.     

Fine structure and functional morphology of the mouthparts of Bittacus planus and Terrobittacus implicatus (Insecta: Mecoptera: Bittacidae)

The Bittacidae are unique in Mecoptera for their adults being predaceous. However, their mouthparts have not been well documented for functional morphology to date. Here, we investigated the mouthpart morphology of the hangingflies Bittacus planus Cheng and Terrobittacus implicatus (Huang & Hua) using scanning electron microscopy. The mouthparts are of the mandibulate type and situated at the tip of an elongated rostrum. The labrum is greatly elongated, roughly twice as long as the subquadrate clypeus. The epipharynx is furnished with a row of basiconic sensilla arranged evenly as a median band extending from the apex to the base. The mandibles are slender and elongated, bearing a sharp lateral and a small mesal tooth. The maxillae are well developed, each consisting of a partially sclerotized cardo and a stipes, a hirsute galea and a lacinia, and a five-segmented maxillary palp. The sensillar pattern on the distal segment of the maxillary palp differs slightly between the two bittacid species. The labium is composed of a postmentum, a prementum, and a pair of two-segmented labial palps. The feeding mechanism of bittacids is briefly discussed in combination with the mouthpart morphology and their feeding habits.     

Theoretical approach to the functional optimisation of spiriferide brachiopod shell: Optimum morphology of sulcus

Evidence suggests that biological forms that provide physiological and autecological functions have evolved to adapt to environmental conditions and to optimise requisite morpho-functions. We examined whether shell morphology is functionally optimised to generate passive feeding flow in the Devonian spiriferide brachiopod Paraspirifer bownockeri. This study was based on quantitative results from a computational fluid dynamics simulation and the Lagrangian multiplier method. We estimated the optimum development of the ventral median shell depression, which is called the sulcus, by minimising the pressure difference along the gape. This estimation was made under the constraint that the number of spiral flow rotations must be greater than one, which is effective for spiriferide feeding because of its alignment with the spiral lophophore. During mathematical optimisation, the equation resulted in a suitable flow velocity of approximately 0.1 m/s. At this velocity, the pressure difference was minimised, regardless of sulcus development. The constraint equation showed that the number of spiral flow rotations increased with sulcus development. The optimal solution was similar to the original sulcus form of Paraspirifer under an ambient flow of approximately 0.1 m/s. This result suggests that the variation of shell outline in spiriferids could provide a variety of preferential conditions for ambient flow and that the flow intensity could be adjusted by sulcus development to generate a robust passive feeding flow along the spiral feeding organs.     

Survival and repair of surgical and natural shell damage in the articulate brachiopod Terebratulina retusa (Linnaeus)

Living specimens of Terebratulina retusa from the Firth of Lorn, Scotland, were surgically damaged by drilling 2 mm diameter holes or narrow slits one cm long in the anterior portion of one valve, by bevelling the anterior margin of both valves, or by amputation of the anterior third of one valve. These injuries to the shell and mantle simulated the type of repaired shell damage seen in Paleozoic species, i.e., scalloped, divoted, cleft, and embayed valves. Less than ten percent of the 200 damaged specimens survived until the 25th week after surgery. Specimens of T. retusa showed the ability to repair drill holes, slits, and bevelled anterior shell regions, but not the most severe damage, i.e., amputations of the anterior third of one valve. Shell‐repair was initiated in the fourth week after surgery by the development of a membrane across the wound. The development of caeca in the new shell layer secreted to plug the drill holes became apparent by the eighth week. The punctate pattern was complete in the new, translucent shell material of bevelled and drilled specimens by the 25th week following surgery. Failure of any specimens to survive amputation of the anterior portion of a valve for more than seven weeks after surgery, and the absence of initiation of the repair process, suggests that terebratulids do not have the tolerance for, nor the ability to repair, the severe injuries (embayed valves) which were sustained and mended by extinct strophomenids.     

A reconstruction of Telephina bicuspis, a pelagic trilobite from the Middle Ordovician of the Oslo Region, Norway

Well preserved parts of Telephina bicuspis described from dark limestone concretions of Llanvirn age, in the Oslo Region, Norway, exhibit many features associated with a pelagic life style for this trilobite. The free cheek has a huge holochroal eye with perfectly square lenses and a long genal spine which is unique in being directed vertically upwards rather than ventrally. Additional spines include an occipital spine, the macro spine on the rachis of the 6th thoracic segment and paired spines from each of the two rachial rings of the pygidium. These spines may have had a hydrodynamic and stabilizing function. The genal spine could have had a sensory function or been a dimorphic feature. The hypostoma is figured for the first time and a reconstruction is given for the complete exoskeleton.     

Goings‐on inside a worm: functional hypotheses derived from sexual conflict thinking

Different interests between mating partners regarding the fate of their gametes can lead to sexual conflicts in many species. Although these conflicts can sometimes be dealt with pre‐copulatorily (e.g. by choosing with which partners to mate), they often extend beyond copulation. Post‐copulatory sexual conflicts are expected to be particularly strong in simultaneous hermaphrodites because an individual may have to accept sperm in order to obtain an opportunity to donate sperm, reducing the effectiveness of pre‐copulatory conflict resolution. The present study investigates the post‐copulatory interactions between male and female sexual traits of a highly promiscuous simultaneous hermaphrodite, the free‐living flatworm Macrostomum lignano. Using light and electron microscopy, we show the different levels of complexity of the sperm and the genitalia, and derive hypotheses about how the different traits may represent evolutionary responses to such sexual conflicts. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 370–383.     

Paradigm lost: reconsidering functional form and group hypotheses in marine ecology

Although functional form and functional group models for marine algae have been used extensively, there is little general literature support for these models, and many studies have shown that associated hypotheses are often incorrect. In functional form/group models, a wide range of ecological and physiological functions are assumed to be correlated with general algal form or morphology. In contrast, functional group approaches have been used most successfully in terrestrial and aquatic systems when groupings are based on a particular function rather than overall plant morphology, and when addressing ecosystem-level questions. In this type of functional group approach, a given set of species would likely be grouped differently depending on the function under consideration. Functional groupings are appropriate for many situations and questions, but not all. Certainly, grouping taxa by a particular function can be very useful and often necessary for many ecosystem-level questions and modeling, especially where qualitative results are more important than quantitative predictions, and when there are too many species in a system to consider them all individually. However, when one considers species–species interactions or questions about population biology, the specific responses of individual species must be considered. To make functional group models more useful, we recommend that groupings be based on specific functions (e.g. nutrient uptake rates, photosynthesis rates, herbivore resistance, disturbance resistance, etc.) rather than gross morphology. Explicit testing of performance of a particular function should be made before generalizations can be assumed, and groupings should be used for questions/approaches where they are most appropriate. If models fail when tested, they should be modified using the additional information to generate new hypotheses and models, and then retested.     

The brachiopod family Linoproductidae stehli from the lower Moscovian of the Middle Carboniferous of the Moscow Region

The phylogeny and taxonomy of the brachiopod family Linoproductidae are revised. The change in the distributional pattern of spines and distinctive structure of the median lobe of the cardinal process are shown to be the main apomorphies in the family evolution. A new subfamily, Linispininae, and the included new genera, Linispinella and Linispinus, are described. In the nominotypical subfamily Linoproductinae, a new genus Linoproductoides and two new species included in this genus are described from the Vereiskian deposits of the Moscow Region. The new species Linispinus riparius (Trautschold), L. longus sp. nov., and L. staricensis (Ivanov) are described from the Kashirskian deposits. Another new species is tentatively described as “Linoproductus” kabanovi.     

Functional morphology and evolution of the cystoid Echinosphaerites

Echinosphaerites , known from the Lower and Middle Ordovician, has branched biserial brachioles. Such features are so far unknown in the Rhombifera. Echinosphaerites had a skeletal meshwork like that of other blastozoan echinoderms, with a fine outer mesh layer and an inner coarse mesh layer. During evolution the number and location of brachioles, including the pattern of brachiole branching, changed by increase in the number of brachioles and increased complexity of the branching pattern. The exothecal pore structures increased in the complexity of patterns of tangential canals. The pattern of skeletal growth in Echinosphaerites is discussed. The Echinosphaeritidae and Caryocystitidae families are closely related and show parallel development.     

Discussion of the Early Devonian rhynchonellid brachiopod genus Borealirhynchia Su, 1976

The Early Devonian brachiopod genus Borealirhynchia was established by Su, 1976. Phylum Brachiopoda (Cambrian to Devonian). In: Paleontological Atlas of North China, Inner Mongolia Volume 1. Geological Publishing House Beijing, pp. 155-227 (in Chinese). It has not been previously described in detail, nor have transverse serial sections of the internal features of Borealirhynchia delerensis, the type species, been published. In this paper Borealirhynchia, along with a few species previously assigned to it, is analysed and discussed. Detailed transverse serial sections of the internal features of B.? lata Su, 1976, are provided, based on well-preserved specimens collected from the Lower Devonian strata of Dong Ujimqin Qi, northeastern Inner Mongolia. Borealirhynchia? gigantea Su, 1976 and Latonotoechia multicosta Su, 1976 are considered to be synonyms of B.? lata Su with the same external and internal characters present in all three species. Some Lower Devonian strata, in which Borealirhynchia was found and reported, from Dong Ujimqin Qi, northeastern Inner Mongolia, are fully described.     

Bouchardia rosea,a vanishing brachiopod species of the Brazilian platform: taphonomy,historical ecology and conservation paleobiology

Dead-live faunal comparisons can offer powerful data to detect natural or human-induced population changes in the late Holocene. Here, we document dead–live comparisons for death assemblages of the brachiopod Bouchardia rosea in nearshore (0–45 m) environments along the northern coast of São Paulo State, Brazil. The sampling programme included 30 stations (14 at Ubatuba, 16 at Picinguaba bay). The bottom was sampled via Van Veen grab sampler, and also dredged. Out of 30 stations, 22 yielded brachiopods. The fidelity estimates were obtained by direct comparisons of live biota with dead shells. A total of 6627 brachiopods were recovered, 5339 (80.6%) from Ubatuba and 1288 (19.4%) from Picinguaba. Out of these, 6621 (99.9%) were empty, dead shells, while only six individuals (0.1%) were found alive, all in the Picinguaba Bay. These results suggest extremely poor dead–live compositional fidelity for B. rosea assemblages. The spatial data suggest that the distribution of B. rosea accumulations has been highly patchy in the region, whereas the great scarcity of live brachiopods may point to a recent decline in local populations. Several lines of evidences indicate that changes in water temperature, nutrient availability, population history and even pollution, may have all affected spatio-temporal dynamics of B. rosea populations.     

Filling the gap in functional trait databases: use of ecological hypotheses to replace missing data

Functional trait databases are powerful tools in ecology, though most of them contain large amounts of missing values. The goal of this study was to test the effect of imputation methods on the evaluation of trait values at species level and on the subsequent calculation of functional diversity indices at community level using functional trait databases. Two simple imputation methods (average and median), two methods based on ecological hypotheses, and one multiple imputation method were tested using a large plant trait database, together with the influence of the percentage of missing data and differences between functional traits. At community level, the complete‐case approach and three functional diversity indices calculated from grassland plant communities were included. At the species level, one of the methods based on ecological hypothesis was for all traits more accurate than imputation with average or median values, but the multiple imputation method was superior for most of the traits. The method based on functional proximity between species was the best method for traits with an unbalanced distribution, while the method based on the existence of relationships between traits was the best for traits with a balanced distribution. The ranking of the grassland communities for their functional diversity indices was not robust with the complete‐case approach, even for low percentages of missing data. With the imputation methods based on ecological hypotheses, functional diversity indices could be computed with a maximum of 30% of missing data, without affecting the ranking between grassland communities. The multiple imputation method performed well, but not better than single imputation based on ecological hypothesis and adapted to the distribution of the trait values for the functional identity and range of the communities. Ecological studies using functional trait databases have to deal with missing data using imputation methods corresponding to their specific needs and making the most out of the information available in the databases. Within this framework, this study indicates the possibilities and limits of single imputation methods based on ecological hypothesis and concludes that they could be useful when studying the ranking of communities for their functional diversity indices.     

The latest Ordovician Hirnantia brachiopod fauna of Myanmar: Significance of new data from the Mandalay Region

A new, latest Ordovician brachiopod fauna is systematically described from the Hwe Mawng Purple Shale Member (Hirnantian) of the Naungkangyi Group of the Pa-thin area, Mandalay Region, Myanmar, revealing one of the most diverse representatives of the typical Hirnantia Fauna. Rhynchonelliformean, craniiformean and linguliformean brachiopods studied belong to 23 genera and a few indeterminate taxa, among which the most abundant genus Kinnella, along with the four common genera (Paromalomena, Pseudopholidops, Fardenia, and Dalmanella), and other genera (such as Cliftonia, Draborthis, Hindella, Hirnantia, Leptaena, Mirorthis, Plectothyrella, Skenidioides, and Xenocrania) are recorded for the first time. This high diversity Hirnantia Fauna represents an ecological differentiation within the benthos of the Sibumasu Terrane during the end Ordovician global crisis. The palaeoeco-unit named herein as the Kinnella-Paromalomena Association is assigned to lower BA3. This paper includes a taxonomic revision with the following conclusions: 1) Sinomena Zeng et al. and Yichangomena Zeng et al. are treated as junior synonyms of Eostropheodonta Bancroft; 2) Hubeinomena Zeng et al. is regarded as a juvenile form of Coolinia Bancroft; 3) Paramirorthis Zeng et al. is considered an immature growth stage of Mirorthis Zeng; 4) Shanomena Cocks and Fortey is treated as a junior synonym of Paromalomena Rong; 5) Scenidium? medlicotti Reed is redesignated as the type species of Kinnella Bergström to replace Hirnantia? kielanae Temple. The distribution of the Hirnantia Fauna in Myanmar, Thailand and western Yunnan of the Sibumasu Terrane highlights the distinctive nature of the Mandalay fauna, and the analyses of the Hirnantia Fauna and others of Sibumasu shows that Sibumasu was not located far from the South China and Lhasa palaeoplates during late Middle and Late Ordovician.