Extinction and opportunistic evolution among late Wenlock graptolites

The late Homerian was a time of profound importance to graptolites. Following the complete extinction of Cyrtograptus and the near extinction of Monograptus (s.s.)at the end of the lundgrenitestis Zone(lower Homerian, upper Wenlock), the morphologically simple and long-ranging Pristiograptus dubius lineage underwent rapid (opportunistic) cladogenesis during the succeeding upper Homerian. Two clades, each with novel and sometimes complex sicular and thecal structures, appeared. One clade, the 'Pristiograptus' praedeubeli group, gave rise to similar but more complex species, such as ' P.' ludensis and 'P.' deubeli , which in turn gave rise to Ludlow taxa, including Saetograptus (s.l.) and Pseudomonclimacis. The other clade, the Lobograptus group, gave rise to Ludlow taxa Lobograptus, Neolobograptus, Bohemograptus , and Neocucullograptus some with very complex thecal structures. Ludlow and younger monograptid faunas therefore comprise at least three separate lineages: Monograptus (s.s.), the Lobograptus group (clade), and the 'Pristiograptus' praedeubeli group (clade). Cladogenesis, evolution, extinction, Homerian, PRISTIOGRAPTUS, LOBOGRAPTUS     

A conodont, thelodont and acanthodian fauna from the Lower Přídolí (Silurian) of the Much Wenlock area, Shropshire

Conodonts, thelodonts and acanthodians have previously been collected from the P?ídolí of Shropshire (Welsh Borderland) from bone beds such as the Ludlow Bone Bed. In general, specimens collected from these types of deposit are very abraded and make taxonomic studies difficult. This contribution describes well preserved thelodont microelements from two samples from the Downton Castle Sandstone Formation (P?ídolí, Silurian) of the Much Wenlock area of Shropshire. Head scales, a wide variety of transitional scales, and pore scales of the lateral line sensory canals of the thelodont Paralogania ludlowiensis (Gross) are described for the first time. A new acanthodian Nostolepis linleyensis sp. nov. and elements from the apparatus of the conodont Ozarkodina?hemensis sp. nov. are documented from the same samples. The thelodonts in both samples are typical of upper Ludlow–P?ídolí faunas from across the Welsh Borderland, and are also present in smaller numbers in the Ludlow Series of Gotland, Sweden, Saaremaa Island, Estonia, and in Latvia and Lithuania. The conodont O? hemensis sp. nov. is restricted to the Much Wenlock area of Shropshire. The co-occurrence of well preserved elements of conodonts and thelodonts suggests that they were deposited fairly rapidly with little or no re-working and that a restricted marine environment prevailed in the early P?ídolí in the Much Wenlock area of Shropshire.     

Early Ludlow monograptids with Devonian morphological affinities

Nilssoni–scanicus Zone deposits (Silurian, early Ludlow) from the Leintwardine and Long Mountain districts of Britain, and the Clrasi Borehole, Moesian Platform of Romania, have yielded a species of monograptid which, whilst resembling rare Ludlow Monoclimacis species, has strong morphological affinities with Devonian monograptids such as Monograptus ramstalensis Jaeger. Because of the occurrence of such structures at a surprisingly low horizon the writers urge caution in identifying age from monotypic assemblages. The difficulties surrounding the affinities of the low Ludlow species necessitates a provisional open nomenclature as Monograptus sp. The possible evolutionary significance is discussed.     

The earliest vascular land plants: continuing the search for proof

New collections from Bringewoodian strata (Silurian, Ludlow Series, Saetograptus leintwardinensis incipiens Zone) in Wales have yielded the oldest known specimens of the plant Cooksonia . From evidence at younger (Eř;éolí/Downton) horizons this genus is considered to be a vascular plant; although no tracheids have been isolated from the Bringewoodian specimens, on the basis of comparative morphology they are concluded to be the oldest recorded tracheophytes. The types of evidence relevant to the identification of vascular plants are briefly reviewed, and the relationships of the Welsh Cooksonia flora to other late Silurian floras are discussed. Confident stratigraphical dating is a prerequisite for any discussion of early plant evolution, emphasising the desirability of an integrated approach incorporating both geologial and palaeobotanical investigations. The dating of two published records of diverse Silurian floras, including lycopods, is questioned; recent dating of the Australian Baragwanathia flora as early Ludlow on the basis of graptolites is not supported by the shelly faunal evidence, which indicates a correlation within the Lower Devonian.     

Graptolites from glacial erratics of the Laerheide area,northern Germany

Ordovician and Silurian glacial erratics of the Laerheide area (Lower Saxony, north-western Germany) bear well-preserved graptolites. The faunas provide important information on the origin and transport direction of the sediments preserved in a kame, representing the Drenthe stadial of the Saalian glaciation. The faunas even include species not commonly encountered in the successions of mainland Sweden, from where the erratics presumably originated. The most common graptolites are from Upper Ordovician (Sandbian to Katian) limestones and from Katian black shales. More common, however, are greenish limestones, sand- and siltstones, often combined in the term ‘Grünlich-Graues Graptolithengestein’, in which upper Wenlock to Ludlow (upper Silurian) graptolites are common.     

内蒙古达茂旗巴特敖包地区包尔汉图剖面牙形刺生物地层

内蒙古达茂旗(达尔罕茂明安联合旗)巴特敖包地区志留纪、泥盆纪碳酸岩相地层发育,本区珊瑚、腕足类、层孔虫等底栖大化石的研究工作已有一定基础。但志留纪、泥盆纪地层在时代确定和对比上还存在很多问题,必须用主导化石门类牙形刺加以解决。研究表明：包尔汉图剖面的顶部属泥盆系无疑,应归入阿鲁共组,而不是西别河组;巴特敖包地区的海侵,始于罗德洛统卢德福德阶(Ludfordian)早期。本剖面没有发现文洛克世和罗德洛世早期的海相沉积。本文描写了一个志留纪牙形刺新种：Ozarkodina uncrispa sp．nov．。     

Late Silurian trilobites from the Nuratau and Turkestan ranges, Uzbekistan and Tajikistan

Silurian trilobites of Ludlow and P?ídolí age are described from sections in southern Uzbekistan and from adjacent parts of Tajikistan. They belong to 22 species (10 named and 12 under open nomenclature) distributed among 13 genera belonging to the families Proetidae, Tropidocoryphidae, Aulacopleuridae, Scharyiidae, Cheiruridae, Encrinuridae and Odontopleuridae. The following new species are described: Interproetus pentaxus, Paleodechenella turkestanica, P. zaaminicus, Cromus tamchii and Leonaspis nuratensis. The generic associations from the mid Ludlow bear a striking resemblance to coeval ones from Bohemia, and include genera otherwise known only from that region or from adjacent parts of central Europe. The trilobite faunas from the late Ludlow and P?ídolí series are much less diverse and are dominated by genera of the Warburgellinae (Tropidocoryphidae), which include one species common to Vaigatch Island in the Russian Arctic, and Podolia, Ukraine. The closest links of the diverse central Asian mid Ludlow faunas are therefore to central Europe, whilst the sparser younger Ludlow and P?ídolí faunas, such as they are, suggest closer links to Baltica.     

The rhabdosome structure of aSaetograptus species (Graptoloidea, Monograptacea) from a North German glacial boulder

Saetograptus sp. cf.Saetograptus leintwardinensis from the late Ludlow is described from material isolated from a glacial boulder. The species shows several unexpected rhabdosome structures, especially a considerably widening trumpetshaped sicular aperture with a forked dorsal tongue. The high infraspecific variation of the species is documented.     

Devonian features in Ludlovian graptolites — no guide to ancestry

A diverse group of Ludlovian Monograptus species is characterized by the retarded growth of thecal hoods, either proximally [ M. colonus (Barrande), M. chimaera (Barrande) and M. haupti Kühne] or throughout the rhabdosome [M. micropoma (Jaekel)]. In all these species the hoods are secondary structures, i.e. later growth additions to the thecae. By contrast, in the Devonian Monograptus species the thecal hoods are primary structures, i.e. the outward extension of the dorsal thecal wall. and are formed by undelayed growth that typically is completed before growth of the succeeding theca begins. It is among Ludlovian ( M. uncinatus Tullberg) and Pridolian ( M. simih Přibyl) forms with homologous hoods that the roots of the majority of the Devonian Monograptus species might be found. The Ludlovian M. micropoma and M. haupti should not be affiliated to the group of Monograptus vomerinus (Nicholson), which is highly characteristic of, and restricted to, the late Llandovery and Wenlock graptolite faunas.     

Bats (Chiroptera, Mammalia) from Middle Miocene karstic fissure fillings of Petersbuch near Eichstätt, Southern Franconian Alb (Bavaria)

The bats from the karstic fissure fill sites Petersbuch 6, 10, 18, 31, 35 and 48 include 14 species, four of which are new. All samples are numerically dominated by rhinolophids, especially by Rhinolophus delphinensis. The rich samples from Petersbuch 6-18 are striking in their high species diversity (seven species each). Myotis bavaricus nov. sp. is characterised by a series of plesiomorphous characters, while Myotis reductus nov. sp. is more derived. A special trait of Submyotodon petersbuchensis nov. gen. nov. sp. is the coexistence of nyctalodont and (sub-) myotodont molars in one dentary. Miniopterus rummeli nov. sp. is characterised by its large size, a high coronoid process and its widely spaced lower premolars. Composition and origin of the chiropteran faunas are discussed. They are interpreted as an accumulation of bats that perished in the cave or rock crevice and to a negligible extent as remains of owl pellets. The stratigraphic correlation of the faunas, based on rodents, is late Middle Miocene (MN 7/8). This study is based on several thousand specimens.     

内蒙古上新世高特格地点的仓鼠化石(英文)

对历年来采自高特格地点3个不同层位的共227件仓鼠类标本进行了分类描述和研究,这些标本可以归入2属3种,分别为进步中华仓鼠Sinocricetus progressus,大中华仓鼠(新种)S.major sp.nov.和蒙古微仓鼠Nannocricetus mongolicus。作为内蒙古中部地区新近纪动物群序列中的最晚代表,上新世高特格动物群存在一些特有的进步的小哺乳动物种类,S.major新种就是仓鼠类中的一例。大中华仓鼠新种特征为:个体显著大,齿冠高,下臼齿发育有较发达的(假)下中脊,其中m1的下中脊指向前内方向,m2的假下中脊(或称下原尖后臂)前内向与下后尖后壁连接。古地磁的测定表明,Sinocricetus progressus和Nannocricetus mongolicus在高特格剖面上的年限大致为4.2～3.9Ma,S.major只出现于剖面下部,其年限约为4.2Ma.Sinocricetus和Nannocricetus两属目前主要发现于中国北方晚中新世早期至最晚上新世的地层中,它们可能都是中国特有的新近纪仓鼠。对内蒙古中部地区最晚中新世二登图和哈尔鄂博、早上新世比例克和高特格地点的材料对比表明,Sinocricetus牙齿特征可能存在如下演化趋势:M1-2原脊I发育频率逐渐增高,M1-2的中脊与后尖前壁连接的程度逐渐增高,m1下前脊从双支分别与唇舌两侧下前边小尖连接向单支与唇侧下前边小尖连接转变,以及m2的假下中脊与下后尖后壁发生连接的频率逐渐增高。Nannocricetus在其演化中牙齿形态变化可能存在的趋势是:M1-2的牙根数由3增加到4,M1-2的后小脊II逐渐退化消失,m1的下前边尖逐渐收缩变窄和分裂,M2-3和m2-3的舌侧(下)前边脊逐渐退化消失,及m3下中脊出现频率逐渐增高。     

秦岭东段李官桥盆地始新世哺乳动物化石新材料

李官桥盆地是始新世地层发育较全、哺乳动物化石丰富的盆地。本文报道了该盆地两个始新世化石新地点,哺乳动物共计4目7属8种。其中有5个新种,分别为Kuanchuanius? danjiangensis sp. nov.、Asiocoryphodon progressivus sp. nov.、Gobiatherium minutunz sp. nov.、Lophialetes? primus sp. nov.、Eomoropus? zhanggouensis sp. nov.。动物群的时代可能为中始新世早期。     

THE ZONES OF MONOGRAPTUS HERCYNICUS AND MONOGRAPTUS FALCARIUS IN PAI-KHOI

In the continuous, tectonically undisturbed section of the Upper Silurian-Lower Devonian carbonatic-terrigenous deposits on the Kara Sea shore, north-western Pai-Khoi, there occurs a succession of graptolite zones, i.e. those of Pristiograptus transgrediens, Monograptus uniformis s.l., and Monograptus hercynicus , with Monograptus praehercynicus at the base. The upper boundary of the Monograptus hercynicus zone is established by the appearance of Monograptus falcarius Koren', M. aequabilis notoaequabilis Jaeger (= M. aequabilis amdermaensis Koren'), and M. aff. thomasi Jaeger in beds just overlying shales with the youngest: records of M. hercynicus. These beds are to be recognized as the new Monograptus falcarius zone, the extent of which corresponds to the vertical range of the zonal species in the section studied. The Monograptus falcarius zone is considered to be older than the Monograptus yukonensis zone and fills the gap in the graptolite sequence in the lower part of the Pragian.     

REMARKS ON THE STRATIGRAPHY AND MORPHOLOGY OF PRAGUIAN AND PROBABLY YOUNGER MONOGRAPTIDS

Praguian and probably younger graptoloids as well as the early Lower Devonian (Lochkovian) graptoloids are known from all continents except Antarctica and South America. At least five to six graptolite zones can now be distinguished in the Lower Devonian. In comparison with the Lochkovian faunas, the medial to late Lower Devonian graptoloid communities are considerably impoverished and composed exclusively of Monograptus species of the M. uncinatus group. Lino-graptus and Abiesgraptus , which are characteristic of much of the Lochkovian, are lacking in Praguian (and younger) strata. Monograptus pacificus from the southeast Alaskan Karheen Formation appears to be the youngest true graptolite known to date, possibly having a stratigraphic position not far away from the Lower/Middle Devonian boundary.     

Tabulate Corals after the Frasnian/Famennian Crisis: A Unique Fauna from the Holy Cross Mountains,Poland

Famennian tabulate corals were very rare worldwide, and their biodiversity was relatively low. Here we report a unique tabulate fauna from the mid- and late Famennian of the western part of the Holy Cross Mountains (Kowala and Ostrówka), Poland. We describe eight species (four of them new, namely ?Michelinia vinni sp. nov., Thamnoptychia mistiaeni sp. nov., Syringopora kowalensis sp. nov. and Syringopora hilarowiczi sp. nov.); the whole fauna consists of ten species (two others described in previous papers). These corals form two assemblages—the lower, mid-Famennian with Thamnoptychia and the upper, late Famennian with representatives of genera ?Michelinia, Favosites, Syringopora and ?Yavorskia. The Famennian tabulates from Kowala represent the richest Famennian assemblage appearing after the F/F crisis (these faunas appear some 10 Ma after the extinction event). Corals described here most probably inhabited deeper water settings, near the limit between euphotic and disphotic zones or slightly above. At generic level, these faunas show similarities to other Devonian and Carboniferous faunas, which might suggest their ancestry to at least several Carboniferous lineages. Tabulate faunas described here represent new recruits (the basin of the Holy Cross mountains was not a refuge during the F/F crisis) and have no direct evolutionary linkage to Frasnian faunas from Kowala. The colonization of the seafloor took place in two separate steps: first was monospecific assemblage of Thamnoptychia, and later came the diversified Favosites-Syringopora-Michelinia fauna.     

Thelodonts and distribution of associated conodonts from the Llandovery–lowermost Lochkovian of the Welsh Borderland

Thelodont scales from the Middle Llandovery–lowermost Lochkovian of southern Britain are described with three new taxa, Loganellia? unispinata sp. nov., Nethertonodus prodigialis gen. et sp. nov. and Paralogania tarranti sp. nov. established, and Paralogania kaarmisensis Märss identified for the first time from Britain. Thelodonts are rare in the Llandovery and Wenlock series where predominantly Loganellia cf. aldridgei Turner, Loganellia scotica (Traquair) and Thelodus sp. are accompanied by coniform conodonts Panderodus Ethington, Decoriconus Cooper and Dapsilodus obliquicostatus (Branson and Mehl). The majority of the material has been recovered from the Ludlow and P?ídolí series. Thelodus laevis (Pander) and Paralogania martinssoni (Gross) occur in the lower Gorstian. Paralogania kaarmisensis and Phlebolepis elegans Pander come from the Upper Gorstian–Lower Ludfordian and are associated with the zonal conodont Polygnathoides siluricus Branson and Mehl in the upper Gorstian. Thelodus parvidens Agassiz and T. trilobatus Hoppe dominate in the Upper Ludfordian in association with the rarer zonal conodonts Ozarkodina snajdri (Walliser), O. crispa (Walliser) and O. remscheidensis eosteinhornensis (Walliser). The basal P?ídolí Series indicates a change to a thelodont fauna dominated by Paralogania ludlowiensis (Gross) and including Nethertonodus prodigialis. Higher in the P?ídolí Series, a succession in faunas includes Katoporodus cf. timanicus (Karataju¯te‐Talimaa), Paralogania tarranti, Loganellia? unispinata and Goniporus alatus (Gross). In the highest P?ídolí beds Paralogania kummerowi (Gross) and Loganellia cuneata (Gross) can be traced before the incoming of the lowermost Devonian taxon Turinia pagei (Powrie) along with Nikolivia gutta Karataju¯te‐Talimaa. Correlations with the Baltic are suggested from the Lower Ludlow, Upper Silurian, up to the Lower Lochkovian, Lower Devonian.     

New Carboniferous brachiopods from the eastern Great Basin, Nevada, USA: implications for loop ontogeny and evolution in Late Palaeozoic terebratuloids

Terebratuloid brachiopods from two localities in the eastern Great Basin, Nevada, USA, include the new taxa Cryptacanthia savagei sp. nov., Fletcherithyris infrequens sp. nov., Cryptonella simplex sp. nov., and Albelenina alvarezi gen. et sp. nov. The faunas are considered to be mid Desmoinesian (late Moscovian) in age. The brachiopods were found associated in clusters, and many of the specimens are well preserved. Records of Pennsylvanian terebratuloid brachiopods are uncommon, and these faunas have enabled an understanding of internal features previously unknown or poorly understood. Systematic analyses of the faunas have generated some new concepts on the ontogeny and evolution of the loop in Late Palaeozoic terebratuloids. From ontogenetic analyses, it is inferred that the stage of loop development may be the most important aspect in taxonomic classification. Analyses of loop evolution suggest the possibility of a transition from a teloform stage to long-flanged deltiform and deltiform stages. These transitions may be possible through reduction in length of the descending lamellae of the crura by resorption of calcium carbonate or posteroventral folding of flanges. Subsequently, these changes in loop morphology are retained by paedomorphosis within long-term evolutionary processes.     

Upper Silurian microplankton of the Leintwardine Group, Ludlow Series, in the type Ludlow area and adjacent regions

The acritarchs and prasinophyte algae from the Upper Bringewood, Lower Leintwardine, Upper Leintwardine and Lower Whitcliffe formations of the Ludlow type area and surrounding regions are described. The following new taxa are proposed: Cymatiosphaera pumila sp. nov., Melikeriopalla pustula sp. nov., Cheleutochroa beechenbankensis sp. nov., Cymbosphaeridium molyneuxii sp. nov., Flammulasphaera bella gen. et sp. nov., Percultisphaera incompta sp. nov., Salopidium aldridgei sp. nov. and Umbellasphaeridium? wicanderi sp. nov. A holotype is defined for Lophosphaeridium galeatum Hill and a further 34 new taxa are described under open nomenclature. Two biozones, identified by the first appearance of the zone taxon, are defined in the Sunnyhill section, Ludlow [Global Stratotype Section and Point (GSSP) for the base of the Ludfordian Stage, Ludlow Series]. The base of the Leoniella vilis Biozone is identified at 4.52m below the top of the Upper Bringewood Formation at Beechenbank, Aymestrey. The base of the Triangulina sanpetrensis Biozone is identified at 15.49m above the base of the Lower Leintwardine Formation and Ludfordian Stage at the Sunnyhill section. These biozones allow correlation with sections in north–west Spain, Podolia and Gotland. The possibility of using Visbysphaera whitcliffense and U.? wicanderi as biozonal indicators is suggested.     

WUCHIAPINGIAN (LOPINGIAN, LATE PERMIAN) BRACHIOPODS FROM THE EPISKOPI FORMATION OF HYDRA ISLAND, GREECE

Abstract:  Twenty-three species of silicified brachiopods are described from four samples in the middle and upper parts of the Episkopi Formation from Hydra Island, Greece. These brachiopods are newly recorded from the region and together with previously described brachiopods from the same localities constitute the most diverse Lopingian (Late Permian) brachiopod fauna reported in southern Europe. The brachiopod fauna is Wuchiapingian as indicated by the associated conodonts. The fauna from Hydra exhibits strong palaeobiogeographical links with the faunas from South China. In addition, palaeobiogeographical affinities with the faunas of Thailand and the northern peri-Gondwanan region are also present, which implies a peri-Gondwanan origin for Hydra. Palaeoecologically, the brachiopod assemblage from sample EP in the middle part of the Episkopi Formation is dominated by pedically-attached and cementing genera and reflects moderate energy conditions above storm wave base and an abundance of hard substrates provided by sponges in the biohermal habitat. By contrast, the brachiopod assemblage in the other three samples from the upper part of the Episkopi Formation is dominated by spinose genera with a free-resting life habit, suggesting soft substrates in a quiet water environment below storm wave base on the outer part of the shelf. New taxa are Petinospiriferina gen. nov., Hustedia episkopiensis sp. nov., Waterhouseiella hydraensis sp. nov. and Xenosaria tenuis sp. nov.