Phylogenetic relationships of leptellinid brachiopods

The relationships among the diverse genera comprising the family Leptellinidae (Brachiopoda) are reviewed in the light of the revised edition of the Treatise on Invertebrate Palaeontology. Taxonomic work reassessed all the genera identified as Leptellinidae in the most current classification. Four genera were discarded, namely Bekkerella, Benignites, Leptastichidia and Nikitinamena. Cladistic analysis reveals the paraphyly of these genera; their abandonment leading to more morphologically coherent subdivisions of the family. Two subfamilies, Leptellininae and Palaeostrophomeninae, are emended and taxonomically restructured. The palaeogeographical history of the Leptellinidae is complicated. The Leptellinidae are first recorded in Baltica in the late Floian (Early Ordovician) and rapidly dispersed to circum-Iapetus palaeocontinents by the Dapingian, thence to most terranes composing Gondwana by the Darriwilian (Middle Ordovician).     

Trilobites from the Middle Ordovician Stairway Sandstone,Amadeus Basin,central Australia

Abstract

During the Middle Ordovician (Darriwilian) sandstones and siltstones were deposited in the epicontinental Larapintine Sea, which covered large parts of central Australia. The Darriwilian Stairway Sandstone has, for the first time, been sampled stratigraphically for macrofossils to track marine benthic biodiversity in this clastic-dominated shallow-water palaeoenvironment situated along the margin of northeastern Gondwana. The faunas from the Stairway Sandstone are generally of low diversity and dominated by bivalves but include several animal groups, with trilobites representing 25% of the entire shelly fauna. Thirteen trilobite taxa are described from the Stairway Sandstone; the fauna displays a high degree of endemism. One new species, Basilicus (Parabasilicus) brumbyensis sp. nov. is described.     

Revision of two phragmodontid species (Conodonta) from the Darriwilian (Ordovician) of the Canning Basin in Western Australia and phylogeny of the Cyrtoniodontidae

Zhen, Y.Y. 9 July 2019. Revision of two phragmodontid species (Conodonta) from the Darriwilian (Ordovician) of the Canning Basin in Western Australia and phylogeny of the Cyrtoniodontidae. Alcheringa XX, xxx–xxx. ISSN 0311-5518

Based on re-examination of the material used in the original study from the subsurface Goldwyer and Nita Formations (middle Darriwilian, Middle Ordovician) of the Canning Basin, Western Australia, two phragmodontid species (Phragmodus polystrophos Watson and Ph. spicatus Watson) are revised as having a septimembrate apparatus including geniculate (Ph. polystrophos) or nongeniculate (Ph. spicatus) M, triform alate Sa, modified tertiopedate (Ph. polystrophos) or tripennate (Ph. spicatus) Sb, modified bipennate Sc, modified quadriramate Sd, carminate Pa and pastinate Pb elements. Characterized by a carminate Pa element in their respective species apparatuses, these two species demonstrate a close phylogenetic relationship with Phragmodus cognitus Stauffer from the Late Ordovician (Sandbian) of North America. These distinctive shared characters have allowed their accommodation within a new genus, Protophragmodus gen. nov., which represents an evolutionary lineage separated from species of Phragmodus Branson & Mehl (sensu stricto). In addition, it is postulated that the Cyrtoniodontidae might have originated in the early–middle Darriwilian from ‘Plectodina’ in shallow-water settings, with Phragmodus (sensu stricto), the most derived part of the family, perhaps directly evolving from Protophragmodus gen. nov. in the late Darriwilian and then becoming cosmopolitan, deeper-water dwellers in the Late Ordovician.

Y.Y. Zhen [yong-yi.zhen@planning.nsw.gov.au], Geological Survey of New South Wales, W.B. Clarke Geoscience Centre, 947–953 Londonderry Road, Londonderry, NSW 2753, Australia.     

Middle Ordovician (Darriwilian) conodonts from the Goldwyer Formation of the Canning Basin,Western Australia

Zhen, Y.Y., Normore, L.S., Dent, L.M. & Percival, I.G., 11 July 2019. Middle Ordovician (Darriwilian) conodonts from the Goldwyer Formation of the Canning Basin, Western Australia. Alcheringa 44, 25–55. ISSN 0311-5518

Middle Ordovician conodonts attributed to 46 species were recovered from a stratigraphic interval spanning the Willara, Goldwyer and Nita formations in core sections from the Sally May-2 and Theia-1 petroleum exploration wells in the Canning Basin, Western Australia. The Histiodella serrata, Histiodella holodentata and Eoplacognathus pseudoplanus biozones are recognized in the lower and middle part of the Goldwyer Formation, indicative of an early–middle Darriwilian age. This revised conodont biostratigraphy enables more precise correlation with North America and North and South China. Several biogeographically distinctive conodont species, most likely of North Chinese origin, are recorded from the Goldwyer Formation. Their presence signals a strong palaeobiogeographic connection between the Sino-Korean Craton and the Canning Basin on the western margin of eastern Gondwana during the late Middle Ordovician.

Y.Y. Zhen* [yong-yi.zhen@planning.nsw.gov.au], W.B. Clarke Geoscience Centre, Geological Survey of New South Wales, 947–953 Londonderry Road, Londonderry NSW 2753, Australia; L.S. Normore [leon.normore@dmirs.wa.gov.au]; L.M. Dent [louisa.dent@dmirs.wa.gov.au], Department of Mines, Industry Regulation and Safety, Mineral House, Geological Survey of Western Australia, 100 Plain Street, East Perth, WA 6004, Australia; I.G. Percival [ian.percival@planning.nsw.gov.au], W.B. Clarke Geoscience Centre, Geological Survey of New South Wales, 947–953 Londonderry Road, Londonderry NSW 2753, Australia;     

Late Ordovician conodont biozonation of Australia—current status and regional biostratigraphic correlations

Zhen, Y.Y. & Percival, I.G. March 2017. Late Ordovician conodont biozonation of Australia—current status and regional biostratigraphic correlations. Alcheringa 41, xxx–xxx. ISSN 0311-5518.

Seven conodont biozones are recognized in the Upper Ordovician of Australia. The Pygodus anserinus, Belodina compressa and Phragmodus undatus–Tasmanognathus careyi biozones are successively represented in the Sandbian. Although the Erismodus quadridactylus Biozone of the late Sandbian North America Midcontinent succession was previously recognized in the Stokes Siltstone of the Amadeus Basin and the Mithaka Formation of the Georgina Basin in central-north Australia, we argue for a middle–late Darriwilian age for these two units. Four conodont biozones, from oldest to youngest the Taoqupognathus philipi, T. blandus, T. tumidus–Protopanderodus insculptus and Aphelognathus grandis biozones, are established in the Katian of eastern Australia. Taoqupognathus species are particularly useful in correlation of the lower–middle Katian successions of eastern Australia with contemporary rocks in other parts of eastern Gondwana and peri-Gondwana, such as with the three major terranes of North and South China and Tarim. These regions, together with Sibumasu and eastern Australia, were part of the Australasian Superprovince during the Late Ordovician, with a strong palaeobiogeographic identity signalled by domination of Taoqupognathus, Tasmanognathus and Yaoxianognathus. Longstanding difficulties for precise correlation with the well-established North American Midcontinent or Baltoscandian successions in the Late Ordovician, owing mainly to strong endemism of the Australian faunas particularly from shallow-water settings, have been resolved by integration of regional conodont biostratigraphic schemes. The conodont biozonation of the Australian Upper Ordovician reviewed herein also provides a crucial chronological reference for better constraining the temporal and spatial range of Late Ordovician tectonostratigraphic events across the intracratonic basins of northern and western Australia and orogenic belts of eastern Australia.

Yong Yi Zhen* [yong-yi.zhen@industry.nsw.gov.au] and Ian G. Percival [ian.percival@industry.nsw.gov.au], Geological Survey of New South Wales, W.B. Clarke Geoscience Centre, 947–953 Londonderry Road, Londonderry NSW 2753, Australia.     

Ichnofacies of the Stairway Sandstone fish-fossil beds (Middle Ordovician,Northern Territory,Australia)

The Stairway Sandstone is a 30–560 m thick succession of Middle Ordovician siliciclastic sedimentary rocks within the Amadeus Basin of central Australia, deposited in the epeiric Larapintine Sea of northern peri-Gondwana. The Stairway Sandstone is significant as one of only two known Gondwanan successions to yield articulated arandaspid (pteraspidomorph agnathan) fish. Herein we use the ichnology of the Stairway Sandstone to reveal insights into the shallow marine habitat of these early vertebrates, and compare it with that of other known pteraspidomorph-bearing localities from across Gondwana. The Stairway Sandstone contains a diverse Ordovician ichnofauna including 22 ichnotaxa of Arenicolites, Arthrophycus, Asterosoma, Cruziana, Didymaulichnus, Diplichnites, Diplocraterion, ?Gordia, Lockeia, Monocraterion, Monomorphichnus, Phycodes, Planolites, Rusophycus, Skolithos and Uchirites. These ichnofauna provide a well-preserved example of a typical Ordovician epeiric sea assemblage, recording the diverse ethologies of tracemakers in a very shallow marine environment of flashy sediment accumulation and regularly shifting sandy substrates. New conodont data refine the age of the Stairway Sandstone to the early Darriwilian, with ichnostratigraphic implications for the Cruziana rugosa group and Arthrophycus alleghaniensis.     

New,early ostracods from the Ordovician (Tremadocian) of Iran: systematic,biogeographical and palaeoecological significance

New Tremadocian ostracod material from the Alborz Mountains of Iran confirms the early and widespread occurrence of the Ordovician genus Nanopsis, and the apparently simultaneous first appearance of ostracods in the fossil record at the level of the P. deltifer conodont biozone (485.5 Ma) from China to Argentina. Nanopsis pairidaeza sp. nov. adds to the pool of species diversity for the Early Ordovician, though documented Tremadocian ostracod generic diversity remains low, with only four genera. The presence of Early Ordovician ostracods in Alborz, their occurrence elsewhere in palaeocontinental Gondwana, Baltica and China coupled to their marked absence from the Tremadocian of Laurentia and Siberia, supports the notion of the earliest occurrence of ostracods centred on Gondwana/Baltica.     

Conodonts and tabulate corals from the Upper Ordovician Angullong Formation of central New South Wales,Australia

Zhen, Y.Y., Wang, G.X. &; Percival, I.G., August 2016. Conodonts and tabulate corals from the Upper Ordovician Angullong Formation of central New South Wales, Australia. Alcheringa 41, xxx–xxx. ISSN 0311-5518.

The Angullong Formation is the youngest Ordovician unit exposed in the Cliefden Caves area of central New South Wales. Its maximum age is constrained by a Styracograptus uncinatus graptolite Biozone fauna at the very top of the underlying Malongulli Formation, but the few fossils previously reported from higher in the Angullong Formation are either long-ranging or poorly known. From allochthonous limestone clasts in the middle part of the formation, we document a conodont fauna comprising Aphelognathus grandis, A. solidum, Aphelognathus sp., Aphelognathus? sp., Belodina confluens, Drepanoistodus suberectus, Panderodus gracilis, Panderodus sp., Phragmodus undatus, Pseudobelodina inclinata and Pseudobelodina? sp. aff. P. obtusa, which supports correlation with the Aphelognathus grandis Biozone (late Katian) of the North American Midcontinent succession. The species concepts of Aphelognathus and Pseudobelodina are reviewed in detail. Associated corals are exclusively tabulates, dominated by agetolitids, including Agetolites angullongensis sp. nov., Heliolites orientalis, Hemiagetolites breviseptatus, Hemiagetolites sp. cf. H. spinimarginatus, Navoites sp. cf. N. circumflexa, Plasmoporella bacilliforma, P. marginata, Quepora sp. cf. Q. calamus and Sarcinula sp. Affinities of the coral fauna from the Angullong Formation are closer to faunas from northern NSW and northern Queensland than to the locally recognized Fauna III of late Eastonian age in central NSW. We propose a subdivision of Fauna III to account for this difference, with the late Katian Fauna IIIB characterized by the incoming of agetolitid corals. The currently known distribution of representatives of this group with adequate age constraints suggests that agetolitids possibly originated in North China, subsequently migrating to Tarim, South China and adjacent peri-Gondwanan terranes while also spreading eastward to northern Gondwana, where they progressively moved through eastern Australia to reach the central NSW region by the early Bolindian.

Yong Yi Zhen* (yong-yi.zhen@industry.nsw.gov.au) and Ian G. Percival (ian.percival@industry.nsw.gov.au), Geological Survey of New South Wales, W.B. Clarke Geoscience Centre, 947–953 Londonderry Road, Londonderry, NSW 2753, Australia; Guangxu Wang (gxwang@nigpas.ac.cn), State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, 39 East Beijing Road Nanjing 210008 PR China.     

The coral Cothonion from the lower Cambrian of North Greenland

Originally described from the Coonigan Formation of New South Wales, Australia, a second occurrence of the operculate coral Cothonion sympomatum is here described from the Paralledal Formation of North Greenland. Both finds are of late early Cambrian age, Series 2, Stage 4 in the emerging fourfold classification of the Cambrian. The new find supports widespread distribution patterns seen in early Cambrian Small Shelly Fossils, although associated trilobites belong to traditional redlichiid (Australia) and olenellid (Greenland) realms, respectively.     

Occurrence of the ammonite Goodhallites goodhalli (J. Sowerby) in the Eromanga Basin,Queensland: an index species for the late Albian (Cretaceous)

Mortoniceratid ammonites of the Eromanga Basin of Queensland, although uncommon, are reviewed and reassessed utilising all known collections. Representatives of this group are restricted to the Allaru Formation and almost all specimens are Goodhallites goodhalli, a well-known species from the English Gault. The Allaru Formation overlies the Toolebuc Formation, widely considered to be an essentially isochronous unit because of its unusual sedimentary and geochemical character. Using G. goodhalli, the middle and upper Allaru Formation can be directly correlated with the early late Albian orbignyi and auritus Subzones of the Mortoniceras inflatum Zone as recognized in the reference ammonite zonation embedded in the standard Cretaceous time scale. Overlapping ranges of G. goodhalli and Labeceras and Myloceras allow these common Austral heteromorph genera to be also confidently assigned a late Albian age in Australia, matching their biostratigraphic occurrence in South Africa.     

Conodont fauna and biostratigraphy of the Honghuayuan Formation (Early Ordovician) of Guizhou,South China

Serratognathus diversus An, Cornuodus longibasis (Lindström), Drepanodus arcuatus Pander, and eleven other less common conodonts, including Cornuodus? sp., Oistodus lanceolatus, Protopanderodus gradatus, Protoprioniodus simplicissimus, Juanognathus variabilis, Nasusgnathus dolonus, Paltodus? sp., Scolopodus houlianzhaiensis, Semiacontiodus apterus, Semiacontiodus sp. cf. S. cornuformis and Serratognathoides? sp., are described and illustrated from the Honghuayuan Formation in Guizhou, South China, concluding revision of the conodont fauna from this unit, which comprises 24 species in total. The most distinctive species in the fauna, S. diversus, consists of a trimembrate apparatus, including symmetrical Sa, asymmetrical Sb and strongly asymmetrical Sc elements. This species concept is supported by the absence of any other element types in a large collection represented by nearly 500 specimens of this species. The fauna indicates a late Tremadocian to mid-Floian age (Early Ordovician) for the Honghuayuan Formation, which was widely distributed on the Yangtze Platform in shallow water environments. Previously published biostratigraphic zonations for the Honghuayuan Formation are reviewed, and revised on the basis of our knowledge of the entire conodont fauna, supporting the establishment of three biozones, Triangulodus bifidus, Serratognathus diversus, and Prioniodus honghuayanensis biozones in ascending order. Species of Serratognathus enable correlation between Ordovician successions of South China, North China (North China Platform and Ordos Basin), Tarim Basin, and further afield into Malaysia and northwestern Australia.     

Ordovician trilobites from Deh-Molla,eastern Alborz,Iran

Ghobadi Pour, M., 21 June 2019. Ordovician trilobites from Deh-Molla, eastern Alborz, Iran. Alcheringa 43, 381–405. ISSN 0311-5518

Seventeen species from 14 genera of Tremadocian and Darriwilian trilobites, plus two taxa recognizable only down to family level, have been documented from the Lower to Middle Ordovician succession of the Deh-Molla area, southeast of Shahrud in northern Iran. Two species, Asaphellus intermedius and Conophrys multituberculatus, are new to science. Unlike previously documented Iranian faunas, the early Tremadocian trilobite assemblage is characterized by proliferation of the olenid Chungkingaspis sinensis, which is also known as the eponymous taxon of the basal Ordovician trilobite biozone in South China. This is the first record of the occurrence of the olenid biofacies in the Ordovician of Iran. Overall, both the Tremadocian and Darriwilian trilobite assemblages show distinct similarity to the contemporaneous faunas of South China down to species level. Trilobite-based correlation with the Ordovician succession of South China confirms the existence of a hiatus at the base of the Ordovician succession in the eastern Alborz and a significant gap, with the upper Tremadocian, Floian and Dapingian parts of the succession completely missing in Deh-Molla.

Mansoureh Ghobadi Pour [mghobadipour@yahoo.co.uk and m.ghobadipour@gu.ac.ir], Department of Geology, Faculty of Sciences, Golestan University, Gorgan 49138-15739, Iran. *Also affiliated with Department of Natural Sciences, Natural Museum of Wales, Cardiff, Cathays Park, Cardiff CF10 3NP, UK.     

New material from the Ordovician of China indicates that Inocaulis is a graptolite

Muir, L.A., Zhang, Y.-D. & Lin, J.-P. 2012. New material from the Ordovician of China indicates that Inocaulis is a graptolite. Alcheringa 37, 558–559. ISSN 0311-5518.

The problematic Early Palaeozoic fossil Inocaulis has been regarded as an alga, a graptolite and a hydroid by different authors. A new specimen from the Ordovician (late Darriwilian) of Guizhou Province (China) shows fusellar banding, confirming that it is a benthic graptolite.

Lucy A Muir [lucy@asoldasthehills.org], Yuan-dong Zhang [ydzhang@nigpas.ac.cn], Jih-Pai Lin [jplin@nigpas.ac.cn] (corresponding author), State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, No. 39 East Beijing Road, Nanjing 210008, PR China. Received 13.12.2012; revised 10.5.2013; accepted 13.5.2013.     

The early bivalve radiation in the Ordovician Gondwanan basins of Argentina

The Ordovician diversification is marked by an increase in both marine diversity and ecospace occupation. Bivalves, like other groups, underwent a remarkable diversification in the Early Ordovician. The early phases of such a bivalve diversification took place in the Gondwanan basins of western Argentina. In the Northwestern Argentina (NWA) Basin, three clades originated during late Tremadocian–Floian times. In the Floian successions of the Famatina Basin, a probable basal arcoid is recorded. Genera from these two basins belong to 13 families. Phylogenetic analysis of the NWA heteroconchian bivalves indicates that redoniids and coxiconchinids may have originated during this radiation event. This taxonomic radiation also implies an ecological diversification. Ten guilds are recognized on the basis of bauplan, mode of life, and feeding types. Lifestyles included free endofaunal, free semi-endofaunal, semi-endobyssate, and epibyssate; feeding types included suspensivorous and detritivorous habits. Physiological changes imposed by colonization of low-salinity environments also account for guild definitions. Recent discoveries of Tremadoc to early Darriwilian bivalves from the NWA and Famatina basins indicate that the dominance of higher groups (e.g. Heteroconchia, Pteriomorphia) deviates from the patterns evident in other Gondwanan basins. This agrees with previous ideas supporting the importance of local radiations during the Ordovician diversification. Two new taxa are described, Eoredonia orientalis gen. et sp. nov. and Babinka notica sp. nov., and Coxiconchia sellaensis Sánchez & Babin is first reported from the NWA Basin.     

Late Silurian and Early Devonian biostratigraphy in the Hill End Trough and the Limekilns area,New South Wales

Precordilleran Late Palaeozoic brachiopod genera considered for this palaeobiogeographical analysis belong to four faunal associations: the low diversity Early Carboniferous (Tournaisian) Protocanites scalabrinii-Azurduya chavelensis Zone; the Late Carboniferous (Bashkirian-Moscovian) Levipustula fauna, which appears in sequences associated with glacial intervals; the latest Carboniferous-earliest Permian (Gzhelian-Asselian) Rhipidomella-Micraphelia and Tuberculatella-Aseptella associations, and the Early Permian (mid to late Asselian) Tivertonia jachalensis-Streptorhynchus inaequiornatus Biozone. The brachiopod affinities of the Precordillera are compared with the biogeographical Late Palaeozoic regions previously suggested by other authors using cluster analysis. During the Bashkirian-Moscovian, the Precordilleran faunas show a high affinity with the Austral Realm because of the presence of the Levipustula faunal elements in Eastern Australia. At the same time the affinity with the central and North American Regions is very low. The Early Permian Precordilleran faunas demonstrate highest affinity with the Gondwanan Realm with several genera also linked to the Tethyan and Boreal Realms. After the Late Carboniferous glacial episodes that affected the southwestern Gondwanan margin, the Precordilleran region underwent climatic amelioration. Subsequently, the Early Permian Precordilleran brachiopod assemblages are characterized by typical cool to cold water genera widely developed in the core Gondwanan Realm, with fewer genera (such as Kochiproductus, Rhynchopora and Neochonetes) suggesting a warmer water influence. The Precordilleran faunas demonstrate an increase in brachiopod diversity from the Carboniferous to the Permian, related to water temperature changes and to the palaeogeographical evolution of the southwestern Gondwanan margin and the movement of Gondwana across the South Pole.     

Possible fossil impression in sandstone from the late Palaeoproterozoic-early Mesoproterozoic Semri Group (lower Vindhyan Supergroup), central India

The Lower–Middle Ordovician Zitai Formation of the South China palaeoplate consists of a succession of purple red, nodular argillaceous limestones. Palaeogeographically, it is distributed along the southeastern margin of the Yangtze Platform, and is of late Floian to Dapingian age, correlative with the Dawan Formation of the Middle and Lower Yangtze Platform. In Shitai County, Anhui Province, East China, the Zitai Formation is rich in conodonts, enabling the recognition of four biozones based on first appearance data. Detailed palaeontological and biostratigraphical study of these conodonts reveals that the Ordovician conodont radiation in the Lower Yangtze Platform attained its first diversity peak low in the Oepikodus evae Biozone. This diversification is generally consistent with macroevolutionary trends of brachiopods of South China and graptolites of the Upper Yangtze Platform, but was earlier than that of trilobites and acritarchs of the same palaeoplate. Correlation with the sea-level curve for South China suggests that conodont diversity change during the Ordovician radiation was mainly controlled by regional sea-level fluctuations.     

A new temnospondyl amphibian from the Lower Triassic of South America

A new temnospondyl from the Early Triassic Buena Vista Formation (Paraná Basin) in Uruguay is described. The specimen is represented by a skull table fragment which includes the left orbit, and, although fairly incomplete, shows close affinities with the basal temnospondyl clade (Dvinosaurus + Tupilakosauridae). Previously, members of this clade were only known from Upper Permian and Lower Triassic deposits from the Northern Hemisphere. The new specimen described herein together with a recent record from the Lower Triassic of South Africa (Karoo Basin), support the hypothesis of an invasion of members of the basal (Dvinosaurus + Tupilakosauridae) clade into Gondwana as early as the Late Permian.     

Archaeocyaths from Yorke Peninsula,South Australia and archaeocyathan Early Cambrian zonation

Zhuravlev, A. Yu., & Gravestock, D. I., 1994:03:28. Archaeayaths from Yorke Peninsula, South Australia and archaeocyathan Early Cambrian zonation. Alcheringa 18, 1–54. ISSN 0311-5518.

Two assemblages of archaeocyaths are documented from Lower Cambrian outcrops and drillholes on Yorke Peninsula. South Australia. The older assemblage (11 species) occurs in the uppermost Kulpara Formation and conformably overlying basal Parara Limestone, and is equivalent to Lower Faunal Assemblage II in the Flinders Ranges. The younger assemblage (28 species plus Acanthhcyathus and Rodiocyathus) occurs in the Koolywurtie Member near the top of the Parara Limestone. Equivalent taxa are widespread in the Flinders Ranges, western New South Wales and Antarctica. Archaeocyathan distribution is now sufficiently well known to propose three assemblage zones and two informal assemblages for regional correlation. No new taxa have been added, but Erugatocyathus scutatus (Hill) and Pycnoidocyathus latiloculatus (Hill), hitherto known only from Antarctica, are found in the upper assemblage on Yorke Peninsula. Irregular archaeocyathan systematics are discussed, the ontogeny of Archaeopharetra irregularis (Tylor) clarified, and Kruseicnema Debrenne. Gravestock & Zhuravlev, represented by K. gracilis (Gordon), is fully described.     

A new trilobite species from the upper Rhuddanian (lower Silurian) of South China,and its biogeographical implications

Anacaenaspis yanpingensis sp. nov., from the lower Niuchang Formation (upper Rhuddanian, lower Llandovery, lower Silurian), is the first record of this genus from South China. The biogeographical distribution of Anacaenaspis and some other trilobite genera from South China (e.g., Gaotania, Hyrokybe, Aulacopleura and Raphiophorus) in the Llandovery evidences faunal exchanges between Avalonia–Baltica, Laurentia, Australia and South China. We attribute these dispersals to prevailing ocean currents, and especially equatorial countercurrents, which would have propagated dispersals during the Rhuddanian, shortly after the end-Ordovician mass extinction.     

A new assemblage of graptolites,rhabdopleuran hemichordates and chitinous hydroids from the late Arenig (Ordovician) of the Banestan area,east-central Iran

A rich benthic and planktonic graptolite fauna is associated with encrusting rhabdopleuran hemichordates and chitinous hydrozoans in the late Arenig (Ordovician) part of the Katkoyeh Formation at Banestan, east-central Iran.

New taxa described here are the dendroid graptolites Callograptus huckriedei sp. nov. and Dictyonema bitubulata sp. nov. Other dendroid graptolites Dendrograptus sp. cf. D. flexuosus J. Hall, 1865; Acanthograptus divergens Skevington, 1963; and Thallograptus ?succulentus Ruedemann, 1904 are described. The tuboid graptolite Galeograptus sp., the rhabdopleuran hemichordates Rhabdopleura sp. aff. R. primaevus (Kozlowski, 1967) and Kystodendron sp., and the chitinous hydroid Palaeotuba sp. are also described. Graptoloid graptolites described here are Aulograptus? sp., Didymograptus incertus Perner, 1895, Undulograptus formosus (Mu & Lee, 1958), and Yutagraptus sp. cf. Y. mantuanus Riva, 1994. All but two species-level taxa are described from Iran for the first time.

The fauna is confirmed as being late Arenig or, less likely, early Llanvirn and thus probably correlates with the early part of the Darriwilian stage. It resembles the Atlantic cold water faunas in lacking isograptids and sinograptids.