Biostratigraphical value of some Strunian (Devonian, uppermost Famennian) Productidina, Rhynchonellida, Spiriferida, Spiriferinida brachiopods

The space of Devonian time considered in this paper corresponds to the uppermost part of the Famennian or the latest Famennian as a fourfold subdivision of the stage, usually called by many authors Strunian in neritic facies and Wocklumeria Stufe or Wocklumian in pelagic facies. Here, we examine the biostratigraphical value of certain brachiopod genera and species as bio-markers of the uppermost Famennian throughout the world (Europe, the former USSR, Middle East, Asia, North America, North Africa, and Australia). We have focused our study for species which stratigraphical range has been firmly established in correlation with the conodont biozones (Upper expansa, Early, Middle and Late praesulcata), the ammonoid biozones (do VI = “Wocklumeria Stufe”), or the foraminifera biozones (kobeitusana Zone). Other data for which the stratigraphical range is not based on standard biozonation are provided in the annex. The brachiopod bio-markers discussed in this paper belong to 14 Productidina genera (Acanthatia, Araksalosia, Ericiata, Hamlingella, Mesoplica, Nigerinoplica, Orbinaria, Ovatia, Rugauris, Semiproductus, Sentosia, Spinocarinifera, Steinhagella, Whidbornella), 7 Rhynchonellida genera (Araratella, Centrorhynchus, Hadyrhyncha, Megalopterorhynchus, Novaplatirostrum, Rozmanaria, Tchanakhtchirostrum), 13 Spiriferida genera (Brachythyris, Cyrtospirifer, Dichospirifer, Eochoristites, Imbrexia, Parallelora, Prospira, Rigauxia, Sphenospira, Tenisia, Toryniferella, Tylothyris, Voiseyella) and one Spiriferinida genus (Syringothyris). Other orders have not been studied in this paper. The main features of the uppermost Famennian brachiopod taxa represented in this paper are commented. All these taxa are listed as completely as possible throughout the world (with complements in the annex). A quick summary of the geographic distribution of the more represented taxa is given in conclusion.     

SULCIPENTAMERUS (PENTAMERIDA, BRACHIOPODA) FROM THE LOWER SILURIAN WASHINGTON LAND GROUP, NORTH GREENLAND

Abstract:  The brachiopod Sulcipentamerus , previously considered a common endemic pentameride genus in the Lower Silurian (Aeronian) of South China, is reported from largely coeval strata in the Hauge Bjerge, Ymers Gletcher and Odins Fjord formations (Washington Land Group) of North Greenland. Two species are present. In eastern North Greenland, S . cf. dorsoplanus occurs in carbonate strata of Aeronian – early Telychian age, whereas in the western region, a new species, S . lunatus sp. nov., is described. The identification of both forms as Sulcipentamerus is indicated by their relatively long ventral median septum and dorsal inner hinge plates. The new species, Sulcipentamerus lunatus sp. nov., is characterized by a nonlobate shell, a narrow but sharply defined interarea bordering the delthyrium and a deeply concave pseudodeltidium. Despite the two known occurrences of the genus in South China and Greenland, the Sulcipentamerus -bearing Harpidium Fauna of North Greenland (northeastern part of Laurentia) was quite distinct from both the Aeronian–Telychian brachiopod faunas of South China and the southern half of Laurentia. In general, the Aeronian pentameride brachiopod faunas of these three regions are strongly endemic, in contrast to their more tightly clustered counterparts from Avalonia, Baltica, Siberia, Kazakhstan, Tadzhikistan, Kolyma and Novaya Zemlya.     

Early Cambrian organophosphatic brachiopods from the Xinji Formation,at Shuiyu section,Shanxi Province,North China

Abundant and diverse small shelly fossils have been reported from Cambrian Series 2 in North China, but the co-occurring brachiopods are still poorly known. Herein, we describe seven genera, five species and two undetermined species of organophosphatic brachiopods including one new genus and new species from the lower Cambrian Xinji Formation at Shuiyu section, located on the southern margin of North China Platform. The brachiopod assemblage comprises one mickwitziid (stem group brachiopoda), Paramickwitzia boreussinaensis n. gen. n. sp., a paterinide, Askepasma toddense Laurie, 1986, an acrotretoid, Eohadrotreta cf. zhenbaensis Li and Holmer, 2004, a botsfordiid, Schizopholis yorkensis (Holmer and Ushatinskaya in Gravestock et al., 2001) and three linguloids, Spinobolus sp., Eodicellomus cf. elkaniiformis Holmer and Ushatinskaya in Gravestock et al., 2001 and Eoobolus sp. This brachiopod assemblage suggests a late Age 3 to early Age 4 for the Xinji Formation and reveals a remarkably strong connection with coeval faunas from East Gondwana, particularly the Hawker Group in South Australia. The high degree of similarity (even at species level) further supports a close palaeogeographic position between the North China Platform and Australian East Gondwana during the early Cambrian as indicated by small shelly fossil data.     

STROPHOMENIDE BRACHIOPODS FROM THE CHANGWU FORMATION (LATE KATIAN, LATE ORDOVICIAN) OF CHUN'AN, WESTERN ZHEJIANG, SOUTH-EAST CHINA

Abstract:  Late Ordovician strophomenide brachiopods (superfamilies Strophomenoidea and Plectambonitoidea) from the upper Changwu Formation (mid Ashgill, late Katian) of Jianglütang, Chun'an County, western Zhejiang Province, consist of ten genera and 12 species. Five new species of three new genera are recognized: Chunanomena triporcata , Chunanomena sembellina , Cheramomena subsolana , Lateriseptomena modesta , and Lateriseptomena rugosa . The strophomenide brachiopods from the upper Katian strata described in this study and those from the border region of Zhejiang and Jiangxi provinces reported in previous work contain 16 strophomenoid and 12 plectambonitoid genera, and most of the strophomenoids are endemic to South China. Numerical analysis of well-documented late Katian strophomenide brachiopod faunas indicates a strong provincialism, characterized by the highly distinct North American province (Laurentia), the South China-Kazakhstan province, and the Avalonia-Baltica province (Wales, Belgium and Sweden). Surprisingly, the Girvan district of Scotland, which was a peri-Laurentian terrane during the Ordovician, contains a late Katian brachiopod fauna that is more closely related to the contemporaneous brachiopods of Avalonia-Baltica than to those of North America.     

Recent brachiopods collected during the “SEAMOUNT 1” CRUISE off Portugal and the Ibero-Moroccan Gulf (Northeastern Atlantic) in 1987

The Seamount 1 expedition, on board R.V. Noroit in 1987, explored the Lusitanian seamounts off Portugal and the Ibero-Moroccan Gulf of the Northeast Atlantic. A rich brachiopod fauna of about 15 species has been identified in 52 of the 117 stations sampled. The species living around these seamounts are examined, observed using scanning electron microscopy (SEM) to complete their micro-morphological characteristics, and classified in accordance with the Treatise on Invertebrate Paleontology (Brachiopoda Revised, Treatise on Invertebrate Paleontology, Geological Society of America Inc. and University of Kansas, Boulder, Colorado and Lawrence vol. 1 (1997), vols. 2, 3 (2000) and forthcoming volumes). The location of these seamounts, between the Mediterranean and the Caribbean Seas, allows some faunal comparisons to be made. Comparisons are also made with the brachiopod fauna from Seamount 2, which allows taxonomic assessment of the status of one species, Stenosarina crosnieri (COOPER), found in both cruises. Apart from the species characteristics, and the occurrence of early diagenetic transformations of the shells, this paper describes the depth-ranges, the substrate relationships, and highlights the wide colonisation of these seamounts due to the free larval stage of these benthic species.     

滇西保山地区丁家寨组生物群的时代

首次系统描述保山地区丁家寨组的类化石,详细讨论丁家寨组所产类和腕足类动物群及孢粉植物群的时代和性质,提出丁家寨组的时代应与华南早二叠世的紫松阶相当,即相当于国际上的Ａｓｓｅｌｉａｎ阶至Ｓａｋｍａｒｉａｎ阶。确认丁家寨组不存在不同时代化石的再沉积混杂或大量化石时代倒置的异常现象。丁家寨组出现特提斯型Ｅｏｐａｒａｆｕｓｕｌｉｎａ动物类;腕足类组合属介于西澳区和泰马区之间的过渡动物群,西澳区的色彩较浓;     

EARLY CAMBRIAN BRACHIOPODS FROM NORTH-EAST GREENLAND

Abstract:  A diverse assemblage of late Early Cambrian brachiopods is described from the Bastion and Ella Island formations of North-East Greenland. The fauna includes nine species, representing all three extant brachiopod subphyla in addition to the stem group brachiopod Mickwitzia cf. occidens . Four linguliforms: Eoobolus priscus , Botsfordia caelata , Micromitra bella , Vandalotreta sp., three rynchonelliforms: Obolella crassa , Kutorgina reticulata , and an unidentified chileid plus a possible craniiform species occur. The fauna shows similarities to late Early Cambrian (Dyeran Stage) brachiopod faunas of eastern Canada and the United States, but also to faunas from the late Early Cambrian (Botomian–Toyonian equivalent) of Australia, Antarctica and Siberia.     

The Early Silurian brachiopod Eocoelia from the Hudson Bay Basin, Canada

Eocoelia akimiskii sp. nov. from the Lower Silurian (upper Telychian) Attawapiskat Formation of Akimiski Island, James Bay, Nunavut, is the first and oldest Eocoelia to be described from the Hudson Bay Basin, one of three largest inland basins of North America. The new species lacks dental plates, dental cavities, and marginal deflection or lip, which indicates a post-Aeronian form of the well-known Eocoelia lineage. In rib numbers, Eocoelia akimiskii falls between E. curtisi and E. sulcata , being closer to E. sulcata . In rib strength, however, the new species is more closely allied to E. curtisi . Although Eocoelia is currently assigned to the Rhynchonellida because of its lack of spiralia, several features (particularly the lack of a septalium, the presence of a unique notothyrial platform and cardinal process, and dense, free-hanging fibrous growth frills) of the genus and other leptocoeliids are distinctly atypical of the rhynchonellides. The new species occurs in an inter-reef, shelly packstone facies within the Attawapiskat Formation, which is characterized by coral-stromatoporoid reefs with abundant, diverse, reef-dwelling brachiopods and other shelly organisms. The close association of Eocoelia akimiskii with the Attawapiskat reefs supports a shallow subtidal (BA2) setting generally assigned to the Eocoelia Community. The reefs themselves, however, host an extremely abundant brachiopod fauna dominated by Pentameroides , Trimerella , Septatrypa , and Gypidula . Four species of Clorinda are also common elements of the reef-dwelling brachiopods. This demonstrates that the concept of the classic Early Silurian level-bottom brachiopod communities cannot be directly applied to reef-dwelling brachiopod communities.     

滇东曲靖上志留统关底组若干腕足动物居群的生态特征

依据居群生态学的理论，采用大小－频率直方图和生存曲线，对采自滇东曲靖上志留统关底组的４种１４个腕足动物居群进行分析对比，尝试性地阐述了居群结构与沉积环境、居群形态变异与沉积环境之间的关系。     

Latest Guadalupian brachiopods from the Guadalupian/Lopingian boundary GSSP section at Penglaitan in Laibin, Guangxi, South China and implications for the timing of the pre-Lopingian crisis

A brachiopod fauna comprising nine species in eight genera from three closely spaced stratigraphic horizons of the same stratigraphic section is described for the first time from the Laibin Limestone in the uppermost part of the Maokou Formation in the Guadalupian/Lopingian (G/L) GSSP section at Penglaitan, Guangxi Autonomous Region, South China. The brachiopod assemblages are bracketed between two conodont zones: Jinogondolella xuanhanensis Zone below and Jinogondolella granti Zone above and, therefore, they can be safely assigned to the latest Capitanian in age. However, all but one of the nine brachiopod species from the Laibin Limestone carry strong early Lopingian (Wuchiapingian) aspect. Thus, the discovery of this brachiopod fauna not only suggests that some Lopingian brachiopod species had already appeared in the late Guadalupian (Capitanian); more importantly, it has also highlighted the fact that both the previously noted pre-Lopingian life crisis (or end-Guadalupian or Middle Permian mass extinction) and Lopingian recovery/radiation actually occurred in late Capitanian times, sometime before the G/L chronostratigraphic boundary. So far, the Penglaitan GSSP section provides the highest-resolution disappearance patterns of different fossil groups around the G/L boundary.     

Late Ordovician brachiopods from Peru and their palaeobiogeographical relationships

A Sandbian brachiopod association from the Calapuja Formation, in the Peruvian Altiplano, north‐west of Lake Titicaca, has allowed a re‐examination of the palaeobiogeographical relationships between Gondwana and Avalonia during the Late Ordovician, when the palaeocontinents are considered to be already very distant from one another. The brachiopod fauna includes the new species Onnizetina calapujensis sp. nov., Horderleyella chacaltanai sp. nov., Drabovinella minuscula sp. nov. and Tasmanella curtiseptata sp. nov., as well as Caeroplecia sp., Dinorthis cf. flabellulum and Tunariorthis cardocanalis. In addition, Colaptomena expansa expansa and Heterorthis retrorsistria, known from the British Burrellian Stage of the Caradoc Series (late Sandbian) in Wales and the Welsh Borderlands, have also been identified. The brachiopod collection is the most diverse known from a single locality in the whole Central Andean Basin. Within it, forms with clear Gondwanan links occur, such as the new species of Onnizetina, Drabovinella and Horderleyella, and typical representatives of the Avalonian faunas, such as the Welsh Colaptomena expansa expansa and Heterorthis retrorsistria. The brachiopod species exchange between the Proto‐Andean margin of Gondwana and Avalonia, now believed to be possible during the late Sandbian, allows a reconsideration of the global taxonomic affinities of both regions. With this in mind, detrended correspondence analysis (DCA) and cluster analysis have been applied to an updated rhynchonelliformean brachiopod matrix consisting of presence/absence data. The scatter plot resulting from the DCA allows a vivid visualization of the grouping and geographical trends of the South American localities with respect to Avalonia–Baltica and the Mediterranean margin of Gondwana during the Sandbian. Our results agree with previous palaeogeographical reconstructions, depicting Avalonia very close to Baltica and already distant from Gondwana. As a few brachiopod species, with low dispersal potential, would have been able to migrate between those distant palaeocontinents, the existence of intermediate islands in the Rheic Ocean, permitting the transit by island hopping of eurythermal species, must be considered.     

Revision of some spiriferide and spiriferinide brachiopods from the historical type area of the Tournaisian stage (Carboniferous,southern Belgium)

Spiriferides, and to a lesser extent the spiriferinides, rank among the major elements of the Tournaisian brachiopod fauna from the mixed argillaceous–carbonate succession of the Tournai area (Hainaut sedimentation area, southern Belgium), along with productides (chonetidines and productidines) and athyridides. Based on well-preserved and silicified material from the Tournai Formation (Ivorian), representatives of the families Ambocoeliidae (Crurithyris goldfussiana), Strophopleuridae (Voiseyella roemeriana), Mucrospiriferidae (Tylothyris laminosa), and Cyrtinidae (Cyrtina koninckiana, C. sp. indet.) are described. Cyrtina hibernica is reported for the first time in the Visean of Visé (Visé Formation, Belgium). Biotic interactions between spiriferides and diverse organisms are illustrated, notably the diverse epizoans fixed on brachiopods (e.g., microconchids, cornulitids, crinoid holdfasts), single and multiple drill holes, and regeneration traces after predator attacks.     

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.     

Mitochondrial COI sequences of brachiopods: genetic code shared with protostomes and limits of utility for phylogenetic reconstruction

A 1230-bp region of the cytochrome c oxidase subunit I (COI) gene of mitochondrial DNA of each of 16 brachiopod species, representing all five living orders, was amplified by polymerase chain reaction and sequenced. Pairwise comparisons of sequence differences plotted against divergence times estimated from the brachiopod fossil record revealed that, although there are considerable variations in the expected substitution rate among different lineages, amino acid substitutions of the COI sequences may largely become saturated in 100 Ma, due mostly to multiple substitutions at the same site. Coinciding with this result, phylogenetic analysis indicated low bootstrap values for nodes corresponding to divergence events that occurred before 100 Ma, suggesting that COI sequences are suitable only for inference of phylogenetic events subsequent to the Mesozoic. Examination of brachiopod codons corresponding to invariant amino acids in the COI of various other animals suggest the nonuniversal codon relationships UGA = Trp, AUA = Met, AAA/G = Lys, and AGA/G = Ser. These are identical to those in mollusks, annelids, and arthropods, consistent with the conclusion that brachiopods are protostomes, as indicated by previous molecular analyses.     

Brachiopods, biostratigraphy, and correlation of the Permian marine deposits of Mongolia

Six horizons and 11 brachiopod zones are distinguished in the Permian of the Khangai-Khentei (Boreal) and South Mongolian (Tethys) marine basins of Mongolia on the base of monographic study of brachiopods and literary data on other faunal groups. The distinguished in the Permian basins of Mongolia biostratons are correlated with zonal brachiopod scales of neighboring regions of Northeast Asia. Possible bio-geographical relations of the Mongolian Permian marine basins and channels of brachiopods are analyzed. Forty-five brachiopod species are figured including species, which were first described from Mongolia, zonal index species and typical representatives of zonal assemblage.     

Givetian Stringocephalid brachiopods from eastern Yunnan of Southwest China with notes on global distribution of the family Stringocephalidae

Stringocephalid brachiopods are widely known in the Givetian, but little knowledge is obtained concerning their palaeobiogeographical patterns globally, therefore further studies with new updates and details are required. In this paper, we describe two new stringocephalid brachiopod genera: Yangirostra asiatica n. gen. n. sp. (subfamily Stringocephalinae) and Chinellirostra rara n. gen. n. sp. (subfamily Bornhardtininae), and a new species Stringocephalus sinensis n. sp., together with an indeterminate species Parastringocephalus sp., from the Givetian (late Middle Devonian) of eastern Yunnan, Southwest China. Moreover, with information of the stringocephalides from North Africa, Alaska, North America, Europe and Northeast Eurasia, we compile a dataset of family Stringocephalidae containing 32 genera in 7 subfamilies globally. Based on our data, subfamily Stringocephalinae brachiopods show cosmopolitism and considerably wide distribution from Siberia to the northern Gondwana margins (i.e., North Africa and Australia). Nevertheless, the Boreal Realm and Palaeotethyan Realm are depicted in this paper at the subfamily level, which is much different from the previous palaeobiogeographical schemes in the Givetian. Furthermore, palaeobiogeographical links between Siberia, the Urals and western North America (Alaska, Canada, Nevada and Sonora) are confirmed by diversification of the subfamilies Omoloninae and Rensselandiinae. Whereas in Eurasia (i.e., western and eastern Europe, North and South China), many endemic species of the subfamilies Bornhardtininae and Geranocephalinae are present, as well as the Kaplexinae and Leioseptathyridinae.     

Brachiopod palaeobiogeography in the western Tethys during the Early Jurassic diversity maximum: introduction of a Pontic Province

The biogeography of Pliensbachian (Early Jurassic) brachiopods in western Tethys is investigated using complementary multivariate tools including metric and non-metric ordination, additive cluster and bootstrapped spanning network analyses, as well as one-way analysis of similarity and similarity percentage analysis. All analyses were conducted using the Dice and Simpson similarity indices for presence/absence data on occurrence datasets involving 24 assemblages homogenized at the species level, including (403 taxa) or excluding (210 taxa) species found in only one assemblage. The analyses present a highly consistent biogeographical picture involving three main clusters: the Euro-Boreal, Mediterranean and Pontic biochores. The brachiopod species typical of the newly defined Pontic biochore are illustrated. The three assemblages from the Atlas area are interpreted as a fourth biochore, compositionally intermediate between those of the Euro-Boreal and Mediterranean. The Mediterranean biochore can be further divided into an intra- and a peri-Mediterranean group. These five, palaeogeographically well-constrained biochores show moderate to high degrees of species endemicity, ranging from 20% (Atlas) to 58% (Euro-Boreal). Based on available evidence, and after a reasonable cutback of the customary scale of ranks, the following biogeographical categories and names are suggested for the western Tethyan Pliensbachian brachiopod biochores: Euro-Boreal Province, Mediterranean Province (including an intra-Mediterranean and a peri-Mediterranean Subprovince), Pontic Province and Atlas Subprovince. In addition, a still poorly documented brachiopod biochore occurs on the Gondwana margin as a possible precursor of the extensive Middle to Late Jurassic Ethiopian Province.     

Phylogeny and taxonomy of the damselfly genus Enallagma and related taxa (Odonata: Zygoptera: Coenagrionidae)

Abstract The zygopteran genus Enallagma has been the subject of numerous behavioural and ecological studies, but phylogenetic relationships among species have been examined only within eastern North America, and even the composition and diagnosis of the genus are unclear on a world-wide basis. Most authorities currently recognize about seventy species within Enallagma , comprising two major radiations, in North America and Africa. This study, using morphological data, demonstrates that the North American and a few related Palaearctic species form a monophyletic group that is quite distinct from the African species. The latter are themselves divided into at least three, and probably four, separate clades, one of which may be related to E. parvum of India. Consequently, three of Kennedy's long disused genera, Africallagma , Amphiallagma and Proischnura ( Kennedy, 1920 ) are resurrected and two new genera, Azuragrion gen.n. and Pinheyagrion gen.n. are established for the remaining African taxa. Finally, Enallagma is divided into two subgenera, Enallagma s.s ., the typical 'bluets', including many North American, Holarctic and Palaearctic species, and Chromatallagma subgen.n., comprising a group of species of more variable colour that is confined to North America, the Caribbean and northernmost South America.     

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

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

A relict Ordovician brachiopod fauna from the Parakidograptus acuminatus Biozone (lower Silurian) of the English Lake District

At Yewdale Beck in northern England, a brachiopod fauna dominated by species of Hindella, Kinnella, Mirorthis, Paromalomena and Plectothyrella , occurs interbedded with a graptolite fauna which includes Akidograptus ascensus , Atavograptus ceryx , Persculptograptus parvulus and Normalograptus spp. (including probable examples of N. normalis , N. angustus and N. medius ) within the lower part of the Skelgill Formation. This suggests that taxa of the terminal Ordovician Hirnantia fauna occur within the lower P. acuminatus Biozone, representing the youngest documented occurrence of the Hirnantia brachiopod fauna. Biostratigraphy, brachiopods, graptolites, Ordovician-Silurian boundary.