Linking micromorphism,brooding, and hermaphroditism in brachiopods: Insights from Caribbean Argyrotheca (Brachiopoda)

In extant brachiopods, parental brooding of the larvae occurs exclusively within Rhynchonelliformea. Methods of larval protection range from simple retention of the larvae within the mantle cavity, to sophisticated brood care within highly specialized brood pouches found in Argyrotheca and Joania (Terebratulida, Megathyridoidea), Gwynia (Terebratulida, Gwynioidea), and all Thecideoidea (Thecideida). Previous studies on the reproductive biology of Argyrotheca yielded contrasting results on the epithelial origin of the brood pouches in this genus. Here, representatives of different species of Argyrotheca from the Belize Barrier Reef were examined using histological section series. Brood pouches of four species, A. cf. schrammi and Argyrotheca sp. 1–3, are of the same basic structure, formed by invaginations of the anterior body wall and connected to the visceral cavity via the metanephridia. The same four species are simultaneously hermaphroditic, suggesting that fertilization is achieved, at least partly, through selfing. One species, Argyrotheca rubrocostata, differs significantly from all others as it has no brood pouch and gonochoric gonads. Thus, the presence of brood pouches and simultaneous hermaphroditism are concluded to be correlated within Megathyridoidea and proposed to be homologous traits of Joania and several but not all species of Argyrotheca, questioning the monophyletic status of both genera. In contrast to the brood pouches of Thecideoidea, lophophoral epithelium is not involved in the formation of the pouches of Argyrotheca and Joania. Therefore, megathyridoid and thecideoid brood pouches are not homologous but evolved independently within rhynchonelliform brachiopods. All brachiopods with brood pouches share a micromorphic form and a short life span, limiting the space and time available for gamete and larval development. We suggest that the brood pouches and the hermaphroditic gonads of Argyrotheca spp. and Joania compensate these limitations by minimizing the loss of gametes and larvae, and by maximizing the chances of successful fertilization. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

BRACHIOPOD PALAEOECOLOGY IN MIDDLE DANIAN LIMESTONES AT FAKSE, DENMARK

The Danian limestone in the Fakse quarry displays banks of coral limestone interfingering with banks of bryozoan limestone. (1) Previous and present studies on the brachiopods show that ‘Rhynchonella’ flustracea Buch, ‘R.’ faxensis Posselt, ‘Terebratula’ mobergi Lundgren, and Argyrotheca pindborgi (Nielsen) are restricted to the coral limestone of Fakse. (2) Crania (Danocrania) transversa (Lundgren), C. (D.) tuberculata (Nilsson), Isocrania aff. egnabergensis (Retzius), ‘Terebratula’ fallax Lundgren, and Carneithyris incisa (Buch) are known from bryozoan limestone at Fakse and other Middle Danian localities. (3) Common to both kinds of sediments in Fakse quarry and found at other Middle Danian bryozoan limestone localities as well are Terebratulina aff. chrysalis (Schlotheim), Argyrotheca scabricula (Koenen), Megathiris bruennichi (Rosenkrantz), Platidia sp., and Thecidellina? groenwalli (Nielsen). (4) ‘Terebratula cincta’ Nielsen is restricted to Fakse quarry and occurs in both kinds of sediments. (5) Some of the species restricted to the coral limestone appear to be morphologically adapted to fit in between the branches of the dominant coral in Fakse quarry, Dendrophyllia candelabrum Hennig.     

A ‘Problematic fossil’ revealed:Pycnoporidium ? eomesozoicum Flügel, 1972 (Late Triassic,Tethys)—not an enigmatic alga but a strophomenid brachiopod (Gosaukammerella n.g.)

Summary The microproblematicumPycnoporidium ? eomesozoicum Flügel, 1972, from Upper Triassic reefs of the Alpine-Mediterranean region, Turkey Oman and Iran (originally interpreted as possible alga) represents the type species of a new strophomenid brachiopod genus (Gosaukammerella n.g.). The genus is characterized by a very small, millimeter-sized plano-convex shell, whose ventral valve is attached to the substratum (mainly sponges) by symmetrically arranged outgrowths developing from a pseudopunctate, lamellose foliated shell wall and composed of densely spaced subparallel ‘tubes’ comparable with productide spines secreted by papillose extensions of the mantle.Gosaukammerella seems to be the only reliable candidate for the existence of post-Paleozoic strophomenid (productid ?) brachiopods. Gosaukammerella eomesozoica is restricted to possibly cryptic, shaded reef environments inhabited predominantly by sponges serving as substrates for micromorphic brachiopods.     

Recent brachiopods from the southern Brazilian shelf: palaeontological and biogeographical implications

Until recently, the rhynchonelliform (articulated) brachiopod fauna from the Brazilian continental shelf (western South Atlantic) was represented only by the endemic species Bouchardia rosea (Mawe), reported from coastal waters of the states of São Paulo and Rio de Janeiro. The present study, based on samples from coastal (<30 m), shelf, and continental slope waters (99–485 m), documents the South Atlantic brachiopod fauna and shows that this fauna is more widespread, diverse, and cosmopolitan than previously thought. Based on a total of 16,177 specimens, the following brachiopods have been identified: Bouchardia rosea (Family Bouchardiidae), Platidia anomioides (Family Platidiidae), Argyrotheca cf. cuneata (Family Megathyrididae), and Terebratulina sp. (Family Cancellothyrididae). In coastal settings, the fauna is overwhelmingly dominated by Bouchardia rosea . Rare juvenile (<2 mm) specimens of Argyrotheca cf. cuneata were also found at two shallow-water sites. In shelf settings (100–200 m), the fauna is more diverse and includes Bouchardia rosea , Terebratulina sp., Argyrotheca cf. cuneata , and Platidia anomioides . Notably, Bouchardia rosea was found in waters as deep as 485 m, extending the known bathymetric range of this genus. Also, the record of this brachiopod in waters of the state of Paraná is the southernmost known occurrence of this species. The genera Platidia and Terebratulina are documented here for the first time for the western South Atlantic. The Brazilian brachiopod fauna shares similarities with those from the Atlantic and Indian shelves of southern Africa, and from the Antarctic, Caribbean and Mediterranean waters. The present-day brachiopods of the western South Atlantic are much more cosmopolitan than previously thought and their Cenozoic palaeobiogeographic history has to be reconsidered from that perspective.     

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.     

An unusual brachiopod assemblage in a Late Jurassic (Kimmeridgian) stromatactis mud-mound of the Eastern Carpathians (Hăghimaş Mountains), Romania

An unusual fossil assemblage dominated by superabundant rhynchonellid brachiopods in a stromatactis mud-mound is recorded from the Hăghimaş Mountains (Eastern Carpathians), Romania. The mound mainly consists of bioclastic wackestones to packstones with a very rich macrofauna including crinoids, sponges, juvenile ammonites, and echinoids. The brachiopods represent a low-diversity but high-abundance association, dominated by the rhynchonellids Lacunosella and Septaliphoria. The taphonomical features of the fossil assemblage indicate an autochthonous fauna, with successive generations of brachiopods in life position and complete well-preserved individuals in different growth stage alongside an accessory population of crinoids and sponges. Brachiopod-brachiopod endosymbiotic life strategy is documented for the first time from a post-Paleozoic brachiopod assemblage. The mound reveals abundant stromatactis, filled by radiaxial fibrous or drusy calcite cement and internal polymud sediments. This is the first Late Jurassic (Kimmeridgian) stromatactis mud-mound identified in the Eastern Carpathians.     

山东潍坊寒武系苗岭统乌溜阶馒头组上部的腕足动物

寒武系腕足动物属种多样性高、个体数量丰富、形态差异明显、地理分布广泛,具有辅助寒武系三叶虫生物地层划分和对比的潜力.华北板块寒武系苗岭统沉积和化石记录发育良好,是中国苗岭统的经典研究区之一.前人己针对华北寒武系苗岭统乌溜阶腕足动物的系统古生物学开展了一系列基础工作,但这些相关研究主要集中于辽宁地区,目前对华北其他地区苗...     

Brachiopods from cryptic coral reef habitats in the northern Red Sea

In contrast to the Palaeozoic to Jurassic fossil record, modern tropical and subtropical shallow-water brachiopods are typically small-sized and mostly restricted to cryptic habitats in coral reefs, but information on microhabitat-composition is scant. At Dahab, northern Red Sea, living brachiopods of the genus Argyrotheca were only detected on massively encrusted coral colonies attached to encrusting foraminifers and coralline red algae. Three samples from autochthonous sediments underneath coral colonies are comparatively rich in the brachiopod genera Megerlia and Argyrotheca, and additionally show low numbers of Novocrania and Thecidellina. Based on a coarse-grain analysis including more than 16,000 components >1 mm, these brachiopod shells co-occur with skeletal components of 11 higher taxa. Decapods, fixosessile foraminifers, molluscs, scleractinians, and coralline red algae clearly dominate the assemblages. Brachiopods in this study always contribute less than 2% to the sediment composition. This confirms previous results that even in brachiopod habitats the contribution of brachiopod shells to the total sediment composition is almost negligible. Our study indicates that brachiopods co-occur with pteriomorph bivalves and other epifauna in the cryptic habitats with limited space for encrusters or epibionts on the undersides of scleractinians and it tentatively supports the hypothesis of brachiopods preferring habitats with low grazing pressure, because shelly components of grazers (polyplacophorans and regular echinoids) are rare in our samples.     

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.     

Epizoic bryozoans and corals as indicators of life and post-mortem orientations of the Devonian brachiopod Meristella

The in-life and post-mortem orientations of the Lower Devonian brachiopod Meristella atoka from the Haragan Formation (Lower Devonian; south-central Oklahoma) are inferred from the distribution of epizoic bryozoans and the orientations of base plates of epizoic corals. Three bryozoans, Cyphotrypa corrugata, Fistuliporella maynardi and Leioclema pulchellum, and one coral, Favosites conicus, are considered. Most zoaria that contact the commissure terminate at the commissurc, and a few zoaria terminate at growth lines. This suggests that the bryozoans were primarily life associates of Meristella atoka. Collectively, these three bryozoan species most extensively encrusted marginal sectors of the brachial valve and are very rare on the posteromedian sector (the umbo) of the pedicle valve. This distributional pattern indicates that the preferred living orientation of Meristella atoka was umbo-down (posteromedian sector of the pedicle valve resting on or buried in the substratum) with the commissure steeply inclined to the sediment water interface. Most coralla of Favosites conicus that contacted the commissure encrusted over the commissure. This indicates that Favosites conicus either preferentially encrusted Meristella atoka post-mortem or colonized living brachiopods but subsequently caused them to die. Furthermore, Favosites conicus most extensively encrusted anterior sectors of the brachial valve, especially the fold. The lateral and anterior orientations of the commissure with respect to the base plates (holothecae) of Favosites conicus indicates that the brachiopods were oriented approximately horizontal with respect to the base plates. This suggests that the preferred post-mortem orientation of Meristella atoka was resting nearly horizontally on the substratum. These data and interpretations confirm previously inferred in-life orientations of Meristella atoka and are consistent with post-mortem orientations hypothesized for elongate-oval athyrid brachiopods. □Brachiopoda, Athyridacea, Meristella atoka , palaeoecology, autecology, epizoans, bryozoans, corals, sedimentology, Lower Devonian, North America, Oklahoma.     

Traces of predation/parasitism recorded in Eocene brachiopods from the Castle Hayne Limestone,North Carolina,USA

Schimmel, M., Kowalewski, M. & Coffey, BP. 2011: Traces of predation/parasitism recorded in Eocene brachiopods from the Castle Hayne Limestone, North Carolina, USA. Lethaia, Vol. 45, pp. 274–289. The Castle Hayne Limestone (Middle Eocene, North Carolina), noted for its diverse macro‐invertebrate fossils, was sampled to assess if Early Cenozoic brachiopods from eastern North America record any traces of biotic interactions. Systematic surveys of two North Carolina quarries yielded 494 brachiopods dominated by one species: Plicatoria wilmingtonensis (Lyell and Sowerby, 1845). Despite subtle variations in taphonomy, taxonomy and drilling patterns, the two sampled quarries are remarkably similar in terms of quantitative and qualitative palaeoecological and taphonomic patterns. In both quarries, brachiopods contain frequent drillholes (24.5% specimens drilled). The majority of drillholes were singular, perpendicular to shell surface and drilled from the outside. Ventral valves were drilled slightly more frequently than dorsal ones, but site‐selectivity in drilhole location was not evident. Larger brachiopods were drilled significantly more frequently than smaller ones. However, drillhole diameter did not correlate with brachiopod size. The drillholes are interpreted as records of ‘live‐live’ biotic interactions, representing either predatory attacks or parasitic infestations or a combination of those two types of interactions. A notable fraction of specimens bear multiple drillholes, which is consistent with either parasitic nature of interactions or frequent failed predatory events. The high drilling frequency reported here reinforces other reports (from other continents and other epochs of the Cenozoic), which suggest that brachiopods may be an important prey or host of drilling organisms in some settings. The number of case studies reporting high frequencies of drilling in brachiopods is still limited and thus insufficient to draw reliable generalizations regarding the causes and consequences of these occasionally intense ecological interactions. □Brachiopods, drilling parasitism, drilling predation, Eocene, North Carolina, taphonomy.     

Post‐metamorphic allometry in the earliest acrotretoid brachiopods from the lower Cambrian (Series 2) of South China,and its implications

The earliest growth of post‐metamorphic (post‐larval) shells in two species of Eohadrotreta is described from the Cambrian Shuijingtuo Formation of South China. Two different growth patterns can be observed by quantifying developmental variations in size and shape of successive stages of post‐metamorphic shell growth (including the pedicle foramen forming stage, pedicle foramen enclosing stage and intertrough increasing stage) of Eohadrotreta zhenbaensis and Eohadrotreta ? zhujiahensis . The pedicle foramen is never enclosed within the metamorphic shell of E. zhenbaensis , while the enclosed pedicle foramen of E .? zhujiahensis is located directly outside the metamorphic shell after the pedicle foramen enclosing stage. A strongly allometric growth pattern of E. zhenbaensis is demonstrated by the early enclosure of the pedicle foramen; an accelerated lengthening of the ventral intertrough is associated with the development of a more complex dorsal median septum during the intertrough increasing stage. By contrast, E .? zhujiahensis demonstrates possible paedomorphic development by delayed enclosure of pedicle foramen and an associated decreased lengthening of ventral intertrough during the intertrough increasing stage. This ontogenetic developmental sequence represents the marginal accretionary formation and growth of the pedicle foramen, which resembles that of linguloid brachiopods. Furthermore, the developmental process of the pedicle foramen of Eohadrotreta seems to recapitulate the likely evolutionary transition from the Botsfordiidae, with open delthyrium, to the Acrotheloidea, with an enclosed foramen. This study provides a unique opportunity to obtain a complete understanding of the ontogenetic development of the earliest acrotretoids, and casts new light on the phylogeny of lingulate brachiopods.     

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.     

First record of a bivalved larval shell in Early Cambrian tommotiids and its phylogenetic significance

Abstract: Brachiopods are marine Lophotrochozoa whose soft parts are enclosed in a bivalved shell. Although brachiopods are represented by a rich record from the Early Cambrian to the present, the origin of their bivalved body plan remains controversial. The Early Cambrian organophosphatic tommotiids Micrina and Paterimitra from Australia have been proposed as stem brachiopods. Here, we describe their earliest ontogeny, indicating that tommotiids possessed bivalved planktotrophic larvae. The curious combinations of characters in Micrina and Paterimitra indicate that they may belong to the stems of the Linguliformea and Rhynchonelliformea, respectively. The bivalved shell of adult living brachiopods may represent a plesiomorphic character retained from planktic tommotiid larvae; the crown group body plan of the Brachiopoda may have evolved through the paedomorphic retention of a bivalved larval state.     

A new morphometric model in distinguishing two closely related extinct brachiopod species

Pseudoatrypa (Atrypida, Brachiopoda) are extinct Paleozoic brachiopods that are the closest relatives of the modern brachiopods, rhynchonellids. Two Paleozoic brachiopod species were qualitatively distinguished based on close examination of phenotypic characters and traditional morphometrics. These species were recovered for the first time from the Givetian localities of Traverse Group of Michigan. The efficacy of geometric morphometric analyses in discriminating species has been proven for various organisms, but rarely studies have been performed in distinguishing brachiopod species. The advanced morphometric model developed here using the Procrustes-based landmark method is based on the analysis of valve shape in two extinct brachiopod species: Pseudoatrypa lineata and Pseudoatrypa devoniana. Results from quantitative analyses reinforce the qualitative identifications of the two species. The application of this method on four different orientations (dorsal, ventral, posterior and anterior regions) of the brachiopods permits the two species to be clearly separated, and this study proposes that the model can be effectively used for future identification and distinction of fossil species. Additionally, this study will aid biologists, paleontologists and neontologists in making appropriate advancement in systematic studies and further assist with gauging biodiversity congruously.     

In deep water: a crinoid–brachiopod association in the Upper Oligocene of Antigua,West Indies

Extant brachiopods and stalked crinoids are found together in the deeper waters of the Caribbean Sea. Analogous brachiopod/crinoid associations have been reported from diverse palaeoenvironments in the Neogene of the region. Studied examples include the Pleistocene of Jamaica (deeper water fore reef), and the Miocene of Jamaica (island slope chalks), Barbados (accretionary prism) and Carriacou (turbiditic siliciclastic shelf). Comparison with analogous modern environments indicates deposition in 150+m water depth. This association has now been extended back into the Late Oligocene. Crinoids and brachiopods both occur in the Antigua Formation of Antigua; both occur high in the formation, implying deeper water in this retrograde succession. They have received little attention from systematists, although the brachiopods Cistellarcula dubia Cooper and Tichosina foresti Cooper have previously been described from the Antigua Formation; to these, we add Cistellarcula sp., Argyrotheca sp. and Tichosina sp. At Half Moon Bay in southeast Antigua, high in the Antigua Formation, we have found columnals of isocrinid crinoids (cf. Isocrinus sp.) associated with rare brachiopods (Terebratulina sp.) in island slope deposits. These taxa provide independent evidence for the deeper water aspect of this part of the Antigua Formation, in beds that also yield large, thin‐walled fossil sponges.     

Rerooting the rDNA gene tree reveals phoronids to be ‘brachiopods without shells’; dangers of wide taxon samples in metazoan phylogenetics (Phoronida; Brachiopoda)

Molecular phylogenetics has resulted in conflicting accounts of the relationship between phoronids and brachiopods. Taxonomically comprehensive analyses of brachiopod and phoronid ribosomal DNA sequences (rDNAs) rooted with short‐branched mollusc sequences uniformly find that phoronids nest within brachiopods as the sister of the three extant inarticulate lineages. Here, this is called the ‘alternate’ topology because it does not match traditional, morphology‐based ideas. Many other analyses of protein‐coding genes and/or rDNAs place phoronids elsewhere, often as the sister group of all brachiopods, better matching ‘traditional’ ideas. However, these analyses generally are based on data from small selections of brachiopods and phoronids, include data from a wide range of other metazoan taxa, and are rooted with distant outgroups. Here, I show that outgroup rooting of brachiopods and phoronid rDNAs is unreliable, and instead find the root position with procedures that are free from all distortions caused by distantly related taxa, i.e. by Bayesian and maximum likelihood relaxed‐clock analyses of a purely ingroup alignment. All such analyses confirm the ‘alternate’ topology: phoronids belong within the Brachiopoda as the sister group of the inarticulates. In addition, nine factors are identified that (singly or in combination) can cause misreporting of the phylogenetic signal in wide taxon‐range analyses of both rDNA and amino acid sequence data. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012.     

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.     

Phoronid phylogenetics (Brachiopoda; Phoronata): evidence from morphological cladistics,small and large subunit rDNA sequences,and mitochondrial cox1

A matrix of 24 morphodevelopmental characters and an alignment of small subunit (SSU) and large subunit (LSU) rDNA nuclear and cox1 mitochondrial gene sequences (～4500 sites) were compiled from up to 12 phoronids including most named taxa, but probably constituting only a portion of worldwide diversity. Morphological data were analysed by weighted parsimony; sequence data by maximum and Bayesian likelihood, both with Phoronis ovalis as the local outgroup. Morphological and sequence‐based phylogenies were similar, but not fully congruent. Phoronid rDNAs were almost free from mutational saturation, but cox1 showed strong saturation unless distant outgroups and P. ovalis were omitted, suggesting that many phoronid divergences are old (≥100 Myr). rDNA divergence between named phoronid taxa is generally substantial, but Phoronopsis harmeri (from Vladivostock) and Phoronopsis viridis (from California) are genetically close enough to be conspecific. In another alignment, of 24 taxa, phoronid rDNAs were combined with data from brachiopods and distant (molluscan) outgroups. The relative ages of divergence between phoronids and their brachiopod sister‐groups, of the split between the P. ovalis and non‐ovalis lineages, and of other phoronid splits, were estimated from this alignment with a Bayesian lognormal uncorrelated molecular clock model. Although confidence limits (95% highest probability density) are wide, the results are compatible with an Early Cambrian split between phoronids and brachiopods and with the Upper Devonian latest age suggested for the P. ovalis/non‐ovalis split by the putative phoronid ichnofossil, Talpina. Most other ingroup splits appear to be ～50–200 Myr old. Inclusion of phoronids with brachiopods in the crown clade pan‐Brachiopoda suggests that a distinctive metamorphosis and absence of mineralization are ancestral phoronid apomorphies. Worldwide diversity and possible associations between character‐states and life‐history attributes deserve comprehensive further study.     

A new family of Cambrian rhynchonelliformean brachiopods (Order Naukatida) with an aberrant coral‐like morphology

Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian (Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian (Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral‐like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube‐like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian (Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef‐building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well‐developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle.