Fossils, sediments, tectonics

Tectonic displacement and small-scale tsunamis apparently affected deposition of the Kolymbia limestone, Cape Vagia, Rhodes, Eastern Mediterranean. Coarse beds interrupt the sequential build-up of this Pliocene–Pleistocene bryomol limestone. Celleporid bryozoans, bivalves, and brachiopods dominate these beds. The palaeoecology of the thicket-forming Celleporaria palmata is re-evaluated and subsequently revised. The limestone comprises two parasequences in a transgressive systems tract, and deposition occurred at palaeodepths between 30 and 120 m. At intervals, tectonic movements lowered relative sea level and sent slumps of shallow-water fauna downslope. The depositional history was validated using independent sets of data: sediment structure and grain size, palaeobathymetry using bryozoan growth forms and occurrences of modern representatives of bryozoans and other taxa, basin configuration, and regional tectonics. Concordance of these lines of evidence provides a means of evaluating confidence in palaeoenvironmental inferences.     

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.     

Agelacrinitid Edrioasteroidea (Echinodermata) from the Middle Devonian of the Eifel (Rhenish Massif, Germany)

The few isolated reports of agelacrinitid edrioasteroids (“Timeischytes”) from the Middle Devonian of the Eifel (Rhenish Massif, Germany) are revised. In this context, a specimen-rich but low-diversity edrioasteroid association is described from the threshold of the Eisen and Latistriatus Members of the Junkerberg Formation (Eifelian, Middle Devonian) of the Blankenheim Syncline. Predominantly, the autochthonously preserved edrioasteroids settled fixosessile-epibenthically on the frequent-occurring orthid brachiopod Schizophoria schnuri blankenheimensis and represent a wide spectrum of ontogenetic growth stages. They are associated with epibenthic brachiopods, microconchids, bryozoans, and auloporids—exemplifying synecological interactions. Roundish, shallow concavities with the shape and size of preserved adjacent edrioasteroids within encrusting colonies of the bryozoan Eostenopora clivosa are of special interest. It is suggested that the depressions represent former positions of the adjacent edrioasteroids. The new Bellastrella eifeliana n. gen. n. sp. is described, and genus Krama Bell, 1976 is reported from the Middle Devonian of Germany for the first time.     

Structure of bryozoan communities in an Antarctic glacial fjord (Admiralty Bay,South Shetlands)

Bryozoans are among the most important groups of the Southern Ocean benthic macrofauna, both in terms of species richness and abundance. However, there is a considerable lack of ecological research focused on their distribution patterns and species richness on smaller scale, especially in the soft bottom habitats of Antarctic glacial fjords. The aim of this study was to describe those patterns in the Admiralty Bay. Forty-nine Van Veen grab samples were collected at the depth range from 15 to 265 m, in the summer season of 1979/1980, at three sites distributed along the main axis of the fjord. Among 53 identified species of bryozoans, 32 were recorded in the Admiralty Bay for the first time. The most common and abundant species were Himantozoum antarcticum, Inversiula nutrix and Nematoflustra flagellata. Genera such as Arachnopusia, Cellarinella and Osthimosia were the most speciose taxa. It was demonstrated that depth was important for the distribution of the bryozoans. More than half of the recorded species were found only below 70 m. An influence of glacial disturbance was reflected in the dominance structure of colony growth-forms. The inner region of the fjord was dominated almost entirely by encrusting species, while the diversity of bryozoan growth-forms in less disturbed areas was much higher. In those sites the highest percentage of branched, tuft like species represented by buguliform and flustriform zoaria was observed.     

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.     

Bryozoans, corals, and microfacies of Lower Eifelian (Middle Devonian) limestones at Kierspe, Germany

The Lower Eifelian Meinerzhagener Korallenkalk (= upper Cultrijugatus Beds) at Kierspe, Sauerland, contains a rich reefal fauna. Eight bryozoan species are described, two of them are new: the cystoporate Fistuliporella kierspensis n. sp. and the trepostome Leptotrypella sophiae n. sp. The bryozoans from the Meinerzhagener Korallenkalk shows distinct similarities to the Lower–Middle Devonian of Spain (Santa Lucía Formation, Emsian–Eifelian), and to the Middle Devonian (Eifelian) of Transcaucasia. The coral fauna comprises five tabulate corals and one rugose coral that document a paleobiogeographic relationship between Central and Eastern Europe and Central Asia. The associated fauna is represented by brachiopods, ostracods, and echinoderms. The studied limestones also commonly contain calcimicrobes represented by three species. The faunal and microfacial characteristics indicate a shallow marine depositional environment just above the storm wave base, with a supposed depth of 20 m, within the photic zone. The nutrient regime was at least a mesotrophic. The upper boundary of the Cultrijugatus Beds coincides with the Chote?-Event that strongly affected brachiopods, whereas corals and bryozoans were insensitive to this event.     

Bryozoaires et Brachiopodes des “Calcaires bajociens a Bryozoaires” des Causses (France) et leur paléoécologie

Two Lower Bajocian new localities, with many silicified bryozoans and brachiopods have been discovered in Causses (France): Saint-Rome-de-Cernon (Aveyron) and Mende (Lozère). The study of the faunas allows to increase the anterior knowledges supplied by the study of localities in the South of Causses (neighbourhood of Alzon and Sumène = North of the Seuil sud-caussenard).The interest is, at first, the distance between the localities: Mende in the North, Saint-Rome-de-Cernon in the West. Moreover, in the second, well-preserved faunas were fossilized without transport. Bryozoans and brachiopods are very numerous. Large and complete bryozoan colonies have been get out by action of HCl. Fine internal structures like rhynchonellid Nannirhynchia or like thecidean transversarium and hemispondylium have been studied. At last, observations and paleoecological hypothesis are drawn, likewise comparison between three localities or locality-groups known in the Causses basin.The fauna from Saint-Rome-de-Cernon is biocoenosis, the substratum was soft, the hydrodynamism low and the depth, according to lamellibranchs, about 200 meters. The beds with bryozoans and small brachiopods contain a fauna with two predominant bryozoan species: Ceriocava straminea and Mesenteripora wrighti. These two species could built large colonies, in spite of fine granulometry of the sedimentary bottom. These colonies provided supports to less adapted bryozoans, to thecidids and to lamellibranch Lopha. The beds with terebratulids and bryozoans contain lesser bryozoans and thecidids. Terebratulids are the main group and are fixed with their peduncle to organic support above the irregular deposits. Beside these benthic animals, the rhynchonellid Nannirhynchia campestriensis is epibiontic on gorgonids or epiphytic on floating sea-weeds.On the contrary, all fossils from Mende have been more or less carried, excepted large sponges perhaps. The most numerous bryozoans (Ceriocava straminea, Mesenteripora wrighti) and numerous Lenticulines (Foraminifera) lived on hard bottom with moving water in community with crinoids. It is possible that a smaller part of bryozoans, requiring lower hydrodynamism, lived, either in sheltered parts of the crinoid community or closer to the sedimentation zone. Thecidids were fixed either on bryozoans in crinoid communities or on shelly remains and on bryozoans nearer to the sedimentation zone. Like in Saint-Rome-de-Cernon, Nannirhynchia campestriensis was epibiontic, or epiphytic, or both.In comparison with the whole localities from the Seuil sud-caussenard, we notice the simplicity of Saint-Rome-de-Cernon and Mende. The localities from the Seuil sud-caussenard were assemblages proceeding from 2 or 3 biotops. On the contrary, at Saint-Rome-de-Cernon we have a biocoenosis and at Mende a thanatocoenosis supplied by one biotop almost.     

Species- and site-specific bacterial communities associated with four encrusting bryozoans from the North Sea, Germany

Epi- and endozoic bacterial communities associated with four bryozoan species from the Jade (North Sea, Germany) were investigated by the combined application of molecular tools and electron microscopic visualisation of zooids. Denaturing gradient gel electrophoresis of PCR-amplified 16S rRNA gene fragments of associated bacteria displayed specific bacterial community profiles in the examined species Aspidelectra melolontha, Conopeum reticulum, Electra monostachys and Electra pilosa. Actual bacterial epibiosis was only observed on C. reticulum whilst the other bryozoans under investigation were largely free of microbial epibionts. These observations indicated that bryozoan-associated bacteria identified by molecular methods originated from internal cavities of bryozoan zooids. Cluster analysis of DGGE band patterns revealed species-specific bacterial communities in A. melolontha, E. monostachys and E. pilosa. Bacteria associated with C. reticulum were seemingly influenced by site-specific parameters. A comparison of bacterial community profiles between reference and invertebrate surfaces allowed for an interpretation of conspicuous group-specific differences. Operational taxonomic units (OTUs) obtained from a single set of bryozoan replicates that were absent on the inorganic reference samples (mussel shells) were hypothesized to be favourable endobionts. Contrary, OTUs present in the references but absent in bryozoan samples could be assumed to stem from bryozoan-specific defenses against ubiquitous bacterial colonizers. Although there was no experimental evidence for a mutual relationship between prokaryotes and their eukaryotic bryozoan hosts, this study demonstrated that in three out of four bryozoans under investigation associated bacterial communities were characteristically shaped by host attributes.     

Age and distribution of the Late Devonian brachiopod genus Dzieduszyckia Siemiradzki, 1909 in southern China

Conodont biostratigraphical work was done at four sections recently found with occurrence of the rhynchonellide brachiopod genus, Dzieduszyckia Siemiradzki, in southern Guangxi and in the border area between Dushan County of Guizhou and Nandan County of Guangxi, South China. These sections represent two different types of facies, i.e., carbonate platform and intraplatform basin. The conodont analysis reveals that this genus occurs in the Upper triangularis Zone and the Middle crepida Zone at the Dazhai Section, through the Lower to Upper crepida zones at Dalong, and is restricted within the Upper rhomboidea Zone at the two intraplatform basin sections (Changtang and Duli). This result demonstrates that the occurrence of these peculiar rhynchonellide brachiopods in South China, regardless of the depositional environments, is within the Lower Famennian instead of the previously suggested Upper Famennian. Furthermore, this brachiopod genus in South China began to inhabit on the carbonate platform almost since the beginning of the Famennian and did not extend to the intraplatform basin facies until the late Early Famennian.Available biostratigraphic data indicate that during the Early and Middle Famennian, Dzieduszyckia is widely distributed not only in South China, but also throughout the world, such as Morocco and southern Ural.Observation on the new collections from the four studied sections reveals that the peculiar rhynchonellide brachiopods have a great morphological variation within each section. Significant differences existed among the collections from different sedimentary settings and localities, probably reflecting the environmental and geographic constraint on the morphology of Dzieduszyckia. Samples from different layers in the same section have nearly identical morphological variation, suggesting the temporal inheritance in morphology of the rhynchonellide brachiopod.     

Marine faunal assemblages in the Silurian-Devonian Keyser Limestone of the central Appalachians

Sedimentological analysis of the Keyser Limestone (Upper Silurian - Lower Devonian of the central Appalachians) indicates that its sediments were deposited in a range of marginal and shallow marine environments. Major depositional environments include: tidal flat, lagoon, barrier bar and island, and open marine shelf. Each major environment is represented by a lithofacies which is lithologically and faunally distinct. Tidal flat lithofacies are characterized by eurytopic organisms, including ostracodes, gastropods, stromatoporoids and blue-green algae. Lagoon lithofacies are dominated by bryozoans, brachiopods, ostracodes and stromatoporoids. Barrier lithofacies are characterized by rooted crinoids, encrusting bryozoans and robust brachiopods. Open shelf lithofacies contain a diverse fauna of cystoids, crinoids, bryozoans and brachiopods.
The distributions of faunal assemblages in the Keyser show no simple relationship to either water depth or distance from shore. They are, in general, related to the distributions of depositional environments.
Recurring associations of brachiopod genera were not found in the Keyser. With few exceptions, any genus may be found in any subtidal environment. Abundance of brachiopods is related to the abundance of local hard substrates (usually bryozoans).     

Organogenesis during budding and lophophoral morphology of Hislopia malayensis Annandale, 1916 (Bryozoa, Ctenostomata)

Background  

Bryozoans represent a large lophotrochozoan phylum with controversially discussed phylogenetic position and in group relationships. Developmental processes during the budding of bryozoans are in need for revision. Just recently a study on a phylactolaemate bryozoan gave a comprehensive basis for further comparisons among bryozoans. The aim of this study is to gain more insight into developmental patterns during polypide formation in the budding process of bryozoans. Particular focus is laid upon the lophophore, also its condition in adults. For this purpose we studied organogenesis during budding and lophophoral morphology of the ctenostome bryozoan Hislopia malayensis.     

Modern Data on the Innervation of the Lophophore in Lingula anatina (Brachiopoda) Support the Monophyly of the Lophophorates

Evolutionary relationships among members of the Lophophorata remain unclear. Traditionally, the Lophophorata included three phyla: Brachiopoda, Bryozoa or Ectoprocta, and Phoronida. All species in these phyla have a lophophore, which is regarded as a homologous structure of the lophophorates. Because the organization of the nervous system has been traditionally used to establish relationships among groups of animals, information on the organization of the nervous system in the lophophore of phoronids, brachiopods, and bryozoans may help clarify relationships among the lophophorates. In the current study, the innervation of the lophophore of the inarticulate brachiopod Lingula anatina is investigated by modern methods. The lophophore of L. anatina contains three brachial nerves: the main, accessory, and lower brachial nerves. The main brachial nerve is located at the base of the dorsal side of the brachial fold and gives rise to the cross neurite bundles, which pass through the connective tissue and connect the main and accessory brachial nerves. Nerves emanating from the accessory brachial nerve account for most of the tentacle innervation and comprise the frontal, latero-frontal, and latero-abfrontal neurite bundles. The lower brachial nerve gives rise to the abfrontal neurite bundles of the outer tentacles. Comparative analysis revealed the presence of many similar features in the organization of the lophophore nervous system in phoronids, brachiopods, and bryozoans. The main brachial nerve of L. anatina is similar to the dorsal ganglion of phoronids and the cerebral ganglion of bryozoans. The accessory brachial nerve of L. anatina is similar to the minor nerve ring of phoronids and the circumoral nerve ring of bryozoans. All lophophorates have intertentacular neurite bundles, which innervate adjacent tentacles. The presence of similar nerve elements in the lophophore of phoronids, brachiopods, and bryozoans supports the homology of the lophophore and the monophyly of the lophophorates.     

新疆西准噶尔晚泥盆世法门期的苔藓虫动物群及其形态特异性和古环境意义*

本文重新研究了产自新疆西准噶尔晚泥盆世法门期洪古勒楞组和铁列克提组的苔藓虫动物群, 重点关注其形态学特征和古生态学意义。该苔藓虫动物群由隐口目(Cryptostomata)、窗格苔藓虫目(Fenestrata), 变口目 (Trepostomata)和泡孔目(Cystoporata)组成。优势类群隐口目苔藓虫是具有中轴或中板的细枝状和双叶状群体。 它们的虫室具有加厚的外区体壁, 弯曲强烈的自虫室, 数量众多的大虫室(metazooecia)或隐蔽虫室(tectizooecia), 以及径向刺突和/或棘状刺突。多样化的异虫室、刺突、自虫室等结构, 以及不同生长环境中的不同生态组合是新疆西准噶尔法门期苔藓虫成功演化的基础。洪古勒楞组下段和铁列克提组下部的苔藓虫动物群可以划分为三类生态组合(即: No. 1, No. 2和No. 3), 代表前滨—近岸—远岸的沉积环境。其中, 近岸环境的枝状—双叶状—网状苔藓虫组合的属种丰度、分异度最高。     

新疆西准噶尔晚泥盆世法门期的苔藓虫动物群及其形态特异性和古环境意义*

本文重新研究了产自新疆西准噶尔晚泥盆世法门期洪古勒楞组和铁列克提组的苔藓虫动物群, 重点关注其形态学特征和古生态学意义。该苔藓虫动物群由隐口目(Cryptostomata)、窗格苔藓虫目(Fenestrata), 变口目 (Trepostomata)和泡孔目(Cystoporata)组成。优势类群隐口目苔藓虫是具有中轴或中板的细枝状和双叶状群体。 它们的虫室具有加厚的外区体壁, 弯曲强烈的自虫室, 数量众多的大虫室(metazooecia)或隐蔽虫室(tectizooecia), 以及径向刺突和/或棘状刺突。多样化的异虫室、刺突、自虫室等结构, 以及不同生长环境中的不同生态组合是新疆西准噶尔法门期苔藓虫成功演化的基础。洪古勒楞组下段和铁列克提组下部的苔藓虫动物群可以划分为三类生态组合(即: No. 1, No. 2和No. 3), 代表前滨—近岸—远岸的沉积环境。其中, 近岸环境的枝状—双叶状—网状苔藓虫组合的属种丰度、分异度最高。     

Chemo–ecological interactions in Antarctic bryozoans

The role of bioactive metabolites in ecological interactions involving Antarctic bryozoans has been scarcely studied. Bryozoans are one of the most abundant and diverse members of the Antarctic benthos and are preyed upon by diverse kinds of predators. They seem to be casual food items of the common Antarctic sea urchin Sterechinus neumayeri and the ubiquitous omnivorous amphipod Cheirimedon femoratus. In this study, the cytotoxic activity against embryos and sperm of the sea urchin S. neumayeri and the substrate preferences of the amphipod C. femoratus were assessed using organic extracts from Antarctic bryozoans, in order to determine the presence of chemical defenses. New adapted protocols were designed using a solidifying gel for simulating bryozoans’ surface. We analyzed 32 organic extracts from 16 samples that belonged to 13 different bryozoan species. No cytotoxicity was detected against embryos of S. neumayeri, while 12 of the 13 bryozoan species were cytotoxic to sperm at natural concentrations. In the substrate preference assays, the amphipod C. femoratus was repelled by ten species. The variable bioactivities found in both types of organic partitions of extracts indicated the presence of both lipophilic and hydrophilic defenses. Inter- and intraspecific variability of chemical defenses was detected also, suggesting environmental-induced responses, symbiotic production, and/or genetic variability. Possible alternative defensive mechanisms are also discussed for species with low or no chemical defense. Our results clearly support the fact that chemically mediated bioactivity in Antarctic bryozoans is common, and there is a likely ecological role of cytotoxic and repellent compounds for their protection.     

New insights into the larval development of Macandrevia cranium (Müller, 1776) (Brachiopoda: Rhynchonelliformea)

The terebratulid Macandrevia cranium (Müller, 1776) is a representative of articulate brachiopods. However, little is known about its embryology and larval development. In order to obtain reproducible results we used a strict protocol of artificial fertilization under controlled temperature conditions as a basis for our morphological study. Sampling of embryos or developing larvae at frequent intervals led to the most comprehensive collection of preserved developmental stages, ranging from early zygotes to late three-lobed stage larvae. SEM studies of all these stages showed that the development of M. cranium is similar to that of other terebratulid brachiopods. This includes the presence of four bundles of larval setae in three-lobed stage larvae. Our results contradict earlier observations on the development of M. cranium and show that this species exhibits more typical features of articulate brachiopod development than previously thought.     

The life and death of torquirhynchia inconstans (brachiopoda,upper jurassic) in England

Torquirhynchia inconstans, a rhynchonellid brachiopod, shows a curious asymmetric commissure. This is interpreted as an adaptation to life in tidal environments. Type of preservation, growth line, and size frequency analysis indicate that the population analyzed consist of dominantly mature individuals, which by analogy with recent brachiopod populations is a primary feature of the original population, and not due to selective destruction, selective transport, or selective predation of smaller individuals. The possible functions of the asymmetry of Torquirhynchia inconstans are considered, and it is concluded that the brachiopod was adapted to life in tidal environments, a conclusion supported by sedimentological evidence. Asymmetric brachiopods are considered to have developed from colonies of partly asymmetric, variable brachiopods by selection of extreme variants.     

New bryozoan species from the stratotype section of the Upper Kazanian Substage

Representatives of nine bryozoan genera of four orders are identified for the first time from the carbonate deposits of the stratotype section of the Upper Kazanian Substage of the Middle Permian. Among them four new species are established and described: Permopora volgensis sp. nov., Stellahexaformis burovi sp. nov., Alternifenestella esaulovae sp. nov., and Wjatkella definita sp. nov. A brief lithological characterization is provided of the carbonate deposits that contain the remains of these bryozoans and were formed under the conditions of transgression-regression cycles of the Late Kazanian marine basin. Some questions concerning the taphonomy, ecology, and morphology of the bryozoans studied are considered.     

PHYLOGENETIC RELATIONSHIPS AMONG EXTANT BRACHIOPODS

Abstract— The monophyletic status of the Brachiopoda and phylogenetic relationships within the phylum have long been contentious issues for brachiopod systematists. The relationship of brachiopods to other lophophore-bearing taxa is also uncertain; results from recent morphological and molecular studies are in conflict. To test current hypotheses of relationship, a phylogenetic analysis was completed (using PAUP 3.1.1) with 112 morphological and embryological characters that vary among extant representatives of seven brachiopod superfamilies, using bryozoans, phoronids, pterobranchs and sipunculids as outgroups. In the range of analyses performed, brachiopod monophyly is well supported, particularly by characters of soft anatomy. Arguments concerning single or multiple origins of a bivalved shell are not relevant to recognizing brachiopods as a clade. Articulate monophyly is very strongly supported, but inarticulate monophyly receives relatively weak support. Unlike previous studies, the nature of uncertainties about the clade status of Inarticulata are detailed explicitly here, making them easier to test in the future. Calcareous inarticulates appear to share derived characters with the other inarticulates, while sharing many primitive characters with other calcareous brachiopods (the articulates). Experimental manipulation of the data matrix reveals potential sources of bias in previous hypotheses of brachiopod phylogeny. Although not tested explicitly, lophophorate monophyly is very tentatively supported. Molecular systematic studies of a diverse group of brachiopods and other lophophorates will be particularly welcome in providing a test of the conclusions presented here.