Silurian (Wenlock–ludlow) Graptolites from Bolivia

Well preserved middle to upper Silurian (Wenlock–Ludlow) graptolites from Bolivia are described for the first time. Generally monospecific graptolite faunas, of species largely endemic to South America, are found in a few levels in the lower part of the Kirusillas, Rio Carrasco and Uncía formations. The oldest identified level yields specimens of Pristiograptus praedeubeli (Jaeger) and is referred to the upper Wenlock. Younger faunas belong to the Ludlow and include Saetograptus , Monograptus and Neodiversograptus specimens. These may be referred to the Gorstian (lower Ludlow). The fauna includes Saetograptus argentinus robustus ssp. nov. and Monograptus bolivianus sp. nov.     

Pattern of some Silurian bioevents

The best‐known Silurian bioevent occurred at the end of the Wenlock: the lundgreni event, together with the nassa‐ludensis crisis, was established among planktic graptolites. The East Baltic data show several peaks of high diversity (especially triangulatus, turriculatus, scanicus zones) and three levels of low diversity of graptolites (antennularius, radians, ludensis zones). These are more or less coincident with sea‐level changes. Energetic innovation of the shallow shelf corals started in the early Silurian. The late Wenlock regression seems not to affect them seriously but the late Silurian decline coincides with the aridization of the climate and a regression of the shelf seas. The Agnatha had strong radiations in the Wenlock (ludensis Zone) and Ludlow (leintwardinensis Zone), the fishes in the Pridoli. Many Silurian vertebrates were long‐ranging and extinction rate was relatively low; only at the late leintwardinensis level and in the latest Ludlow did considerable extinctions occur.     

Decay and composition of the hemichordate Rhabdopleura: implications for the taphonomy of graptolites

Although the graptolites lacked biomineralised tissue, their skeletons are abundantly preserved in deeper-water mudstones. Decay experiments and observations on the closely related living hemichordate Rhabdopleura demonstrate that the periderm and stolon are highly resistant to decay, remaining intact for months, whereas the zooids are unrecognizable within days. The extreme rarity of the preservation of traces of the zooids in graptoloids reflects their planktic lifestyle; the zooids had normally decayed before burial. Curie-point-gas-chromatography (Py-GC) and Curie-point-gas-chromatography-mass spectrometry (Py-GC-MS) of the periderm of Rhabdopleura confirms that proteinaceous organic matter is a major constituent. Ultrastructurally preserved graptolite periderm (Ordovician, Oklahoma; Silurian, Arctic Canada), on the other hand, is a highly altered kerogen-like substance rich in aliphatic biomacromolecules. The composition of the preserved graptolite periderm reflects diagenetic replacement by components probably mainly derived from algal cell walls.     

Foraminiferal turnover across the Eocene–Oligocene transition at Fuente Caldera, southern Spain: No cause–effect relationship between meteorite impacts and extinctions

We studied planktic and small benthic foraminifera from the Fuente Caldera section, southern Spain, across the Eocene–Oligocene transition. Benthic foraminifera indicate lower bathyal depths for the late Eocene and earliest Oligocene. Detailed high-resolution sampling and biostratigraphical data allowed us to date precisely layers with evidence for meteorite impact (Ni-rich spinel), which occur in the lower part of the planktic foraminiferal Globigerapsis index Biozone and in the middle part of the small benthic foraminiferal Cibicidoides truncanus (BB4) Biozone (middle Priabonian, late Eocene). Major turnovers of foraminifera occur at the Eocene/Oligocene boundary, only. The impact did not occur at a time of planktic or benthic foraminiferal extinction events, and the late Eocene meteorite impacts did thus not cause extinction of foraminifera. The most plausible cause of the Eocene/Oligocene boundary extinctions is the significant cooling, which generated glaciation in Antarctica and eliminated most of the warm and surface-dwelling foraminifera.     

Phylogenetic analysis reveals that Rhabdopleura is an extant graptolite

A phylogenetic analysis of morphological data from modern pterobranch hemichordates (Cephalodiscus, Rhabdopleura) and representatives of each of the major graptolite orders reveals that Rhabdopleura nests among the benthic, encrusting graptolite taxa as it shares all of the synapomorphies that unite the graptolites. Therefore, rhabdopleurids can be regarded as extant members of the Subclass Graptolithina (Class Pterobranchia). Combined with the results of previous molecular phylogenetic studies of extant deuterostomes, these results also suggest that the Graptolithina is a sister taxon to the Subclass Cephalodiscida. The Graptolithina, as an important component of Early–Middle Palaeozoic biotas, provide data critical to our understanding of early deuterostome phylogeny. This result allows one to infer the zooid morphology, mechanics of colony growth and palaeobiology of fossil graptolites in direct relation to the living members of the clade. The Subdivision Graptoloida (nom. transl.), which are all planktic graptolites, is well supported in this analysis. In addition, we recognize the clade Eugraptolithina (nov.). This clade comprises the Graptoloida and all of the other common and well‐known graptolites of the distinctive Palaeozoic fauna. Most of the graptolites traditionally regarded as tuboids and dendroids appear to be paraphyletic groups within the Eugraptolithina; however, Epigraptus is probably not a member of this clade. The Eugraptolithina appear to be derived from an encrusting, Rhabdopleura‐like species, but the available information is insufficient to resolve the phylogeny of basal graptolites. The phylogenetic position of Mastigograptus and the status of the Dithecoidea and Mastigograptida also remain unresolved. □ Biodiversity, Cambrian, Hemichordata, Deuterostomia, Ordovician.     

Volkhov graptolites from the lower-middle Ordovician boundary beds of the St. Petersburg region,Russia

The lower and lowermost Middle Ordovician strata of the Volkhov Stage exposed in the vicinity of St. Petersburg, Russia, yield a graptolite fauna at several stratigraphic levels, being more often confined to the mud-mound clays than to the normally stratified, Condensed carbonates. Overall taxonomic diversity of the Volkhov graptolites is relatively low and associations include no more than seven species in ail localities. TheTetragraptus, Didymograptus andXiphograptus species found are pandemic and have quite extensive stratigraphie ranges within the mid-Arenig Series. Based on the graptolite zonation, the Volkhov Stage can be correlated to thePseudophyllograptus angustifolius elongatus andDidymograptus (Expansograptus) hirundo graptolite biozones of Baltoscandia and to the upperDidymograptus simulans toDidymograptus (Expansograptus) hirundo biozones of the British graptolite succession. TheIsograptus victoriae lunatus lower zonal boundary probably falls within the graptolite-bearing interval of the lower Volkhov Stage. As the result of indirect biostratigraphic and interpretative lithological correlations, the first appearance ofUndulograptus austrodentatus at the base of the Darriwilian Stage is deemed to be close to the boundary of theMegistaspis simon andM. limbata trilobite biozones and the lowerBaltionodus norrlandicus conodont Biozone of the upper Volkhov Stage. Seven dichograptid species belonging to four genera are described, one of them,Azygograptus volkhovensis n. sp., new.      

扬子北缘钟祥地区奥陶系弗洛阶笔石动物群研究

湖北钟祥下奥陶统弗洛阶岩性以灰黑色页岩为主,夹粉砂岩及灰岩透镜体, 化石丰度和分异度高。通过系统的化石采集和鉴定, 于钟祥温峡口剖面识别出笔石7属31种, 大体上为扬子地台区的常见属种。自下而上分为Didymograptellus bifidus带、Corymbograptus deflexus带和Azygograptus suecicus带。该序列与宜昌?南漳地区一致, 但在笔石动物群组成面貌上差异较大,可能与钟祥地区在早奥陶世位于扬子台地边缘, 富集的营养物质和低氧的环境更利于笔石动物群的繁盛和笔石化石的保存有关。     

Discovery of Anticostia uniformis from the Xiazhen Formation at Zhuzhai,South China and its stratigraphic implication

The Xiazhen Formation is an Upper Ordovician lithostratigraphic unit in the Jiangshan-Changshan-Yushan (JCY) area, which contains series of Late Ordovician reef successions. The reef successions of the Xiazhen Formation at Zhuzhai are critical for evaluation of the Late Ordovician marine diversity and palaeoecology. However, their age has long been uncertain and generally is regarded as of upper Katian, based on the occurrences of shelly fossils and correlation with the stratigraphic equivalent Changwu Formation. The newly discovered graptolite species Anticostia uniformis, in the Xiazhen Formation, together with the combined evidence of brachiopods and sedimentology, indicates an age range for the graptolite locality from the Dicellograptus complanatus Biozone to the Diceratograptus mirus Subzone of late Katian, but the graptolites do not rule out the possibility that it is Hirnantian.     

Dissepiments in the graptolite Monograptus turriculatus

An example of the graptolite Monograptus turriculatus (Barrande, 1850) from the crispus Biozone (Silurian, Llandovery) of the Girvan district, Scotland, shows the first recorded instance of dissepiment formation in graptoloid graptolites. The dissepiments are formed by the fusion of thecal spines with the dorsal wall of the adjacent part of the succeeding whorl of this spiral graptolite. The formation of these structures appears easiest to envisage using models of the graptolite colony that do not involve extrathecal tissue. □ Graptolithina , Monograptus, morphology, functional morphology, Silurian, Llandovery.     

Early Tremadocian (Ordovician) graptolite biostratigraphy and correlation

A revision of the early Tremadocian graptolites of the genus Rhabdinopora shows that Rhabdinopora proparabola from the Xiaoyangqiao section, Dayangcha, North China is the oldest species of the genus. Early Tremadocian species of Rhabdinopora can be differentiated easily by the construction of their nemata. The earliest species, Rhabdinopora proparabola, lacks a nema, while a multibranched nema is present in Rhabdinopora campanulatum (= Rhabdinopora parabola). The Rhabdinopora proparabola, Rhabdinopora campanulatum, Rhabdinopora flabelliformis (Anisograptus matanensis Biozone) and Rhabdinopora anglica biozones can be correlated worldwide. A short interval at the base of the Ordovician System does not contain any planktic graptolites.     

Ecostratigraphy, zonation and global correlation of earliest Ordovician planktic graptolites

The succession of earliest Ordovician planktic graptolites (graptoloids) in 11 key sequences around the world is reviewed and used to assess their distribution in depth facies, in paleolatitude and in time. An ecostratigraphic model shows the pattern of distribution of species across the shore-to-ocean profile in time. Fluctuations in this pattern are related to eustatic change; the Acerocare, Black Mountain, Ceratopyge and possibly, the Peltocare Regressive events are recognized. Surface-water temperature is found to have had little significant effect on distribution of Early Tremadoc species, and once the effects of facies is allowed for, their biostratigraphic utility can be assessed. A global suite of four chronozones is proposed for the Early Tremadoc and a further five zones comprise a provisional suite for the Late Tremadoc. The nine zones enable precise correlation of Tremadoc graptolitic sequences. The zones replace the previously used graptolite assemblages, are correlated with key conodont horizons and are, in upward sequence, the Zones of Rhabdinopora praeparabola, R. flabelliformis parabola, Anisograptus matanensis, R. f. anglica, Adelograptus, Paradelograptus antiquus, Kiaerograptus, Araneograptus murrayi-Araneograptus pulchellus , and Hunnegraptus copiosus.     

Restudy of some Ordovician–Silurian boundary graptolites from Anticosti Island, Canada, and their biostratigraphic significance

The Ellis Bay Formation on Anticosti Island has long been recognized for the biostratigraphic importance of its latest Ordovician conodont, palynomorph, and shelly fossil assemblages. However, a sparse record of graptolites has made it difficult to correlate these assemblages with the graptolite biozonation. Restudy of some previously described and examination of newly collected normalograptid specimens from the Ellis Bay and lower Becscie formations, however, shows that biostratigraphically important taxa are present and these species provide important constraints on the age of the strata. Normalograptus parvulus and N. minor have been identified in the upper half of the Ellis Bay Formation, suggesting that these strata are late Hirnantian in age ( N. persculptus Biozone). Normalograptus imperfectus and Normalograptus sp. aff. N. acceptus occur in the basal beds of the Becscie Formation, indicating that these strata are earliest Silurian. These graptolite data support the hypothesis that the positive carbon isotope excursion seen in the uppermost strata of the Ellis Bay Formation is isochronous with that seen in the Hirnantian strata at Dob's Linn, Scotland, and does not span much of the lower to mid-Hirnantian, as is the case in Arctic Canada and Nevada, USA.     Anticosti Island , Hirnantian, graptolites, Ordovician, Silurian.     

Proposal for adoption of the base of the Undulograptus austrodentatus Biozone as a global Ordovician stage and series boundary level

The base of the Undulograptus austrodentatus Biozone appears to be a synchronous event that is widely recognizable within graptolitic facies around the world. It occurs within an interval in which graptolite species ranges are now well known and in which there is a rapid turnover in the composition of graptolite faunas. This turnover reflects the rapid evolutionary radiation of the Diplograptacea simultaneously with the appearance of several distinctive pseudisograptid and glossograptid species. These events provide the basis for the recognition of two thin but widely applicable subzones; a lower Arienigraptus zhejiangensis Subzone and an upper U. sinicus Subzone. The occurrence of the lower boundary of the U. austrodentatus Biozone within a succession of first appearances also permits accurate and reliable identification of the boundary as well as assessment of stratigraphic completeness across the boundary interval in correlated sections. Diverse graptolite faunas of late Yapeenian and early Darriwilian age occur in association with the Histiodella altifrons Biozone of the North American midcontinent conodont zonation and the Paroistodus originalis and Microzarkodina parva biozones of the North Atlantic conodont zonation. They also occur in association with the shelly-fossil zonations developed for several different continents. These features of the base of the U. austrodentatus Biozone make it a suitable level for use as the boundary level for a global stage. Its stratigraphic position within the Ordovician System relative to other likely global stages as well as its coincidence with one of the major events in graptolite evolutionary history suggest that this level also may be a suitable level for the base of a global Middle Ordovician Series.Ordovician System, Ordovician stages, graptolite zonation, chronostratigraphy, international correlation. Charles E. Mitchell and Jörg Maletz, Department of Geology, State University of New York at Buffalo, Buffalo, New York 14260-1550, USA; 13th July, 1994; revised 22nd May, 1995.     

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.     

Preservation of soft tissues in Silurian graptolites from Latvia

The contractile stalks of graptoloid zooids are preserved as organic carbon residues in thecae of the middle Llandovery graptoloid graptolites Rastrites geinitzii and Neolagarograptus? sp. from the Aizpute‐41 core, Latvia. The contractile stalks are surrounded by equant pyrite crystals, resulting in three‐dimensional preservation of the graptolite rhabdosomes, and are associated with sediment of similar composition to, and derived from, the adjacent matrix. Matrix entered the thecae after pyrite crystal growth and filled some of the space left by collapse of the contractile stalks and some intercrystalline cavities; other space is partially infilled by diagenetic minerals. The contractile stalks are parallel‐sided and occupy up to one‐half the metathecal width, which is not inconsistent, assuming post‐mortem shrinkage, with the suggestion that graptoloid zooids filled their thecal tubes in life. The location of the preserved soft tissues, towards the distal ends of the metathecae, is very different from that predicted by decay experiments on the extant pterobranch hemichordate Rhabdopleura; the latter's soft tissues may thus not be a reliable analogue for those of these Silurian graptoloids.     

A refined graptolite biostratigraphy for the late Ordovician–early Silurian of central Wales

Morphometric analysis of graptolites from the persculptus and acuminatus biozones of central Wales identifies four successive morphospecies of normalograptids. These graptolites can be used for biostratigraphical subdivision of these strata as follows: (i) an early persculptus Biozone interval containing broad forms with geniculate thecae that have the morphology of Persculptograptus persculptus with an early insertion point for the full median septum (theca 1¹); (ii) a supra‐adjacent level of early persculptus Biozone age, with narrower, parallel‐sided forms, that have been referred to as Normalograptus? aff. parvulus and have a slightly later insertion point for the full median septum (theca 1²); (iii) a third interval, encompassing the later part of the persculptus Biozone to the early acuminatus Biozone, with Normalograptus? cf. parvulus, which has the full median septum delayed to the level of theca 3–7; and (iv) a younger interval, in the mid‐acuminatus Biozone with Persculptograptus cf. persculptus specimens that do not display the median septum on its reverse side. These taxa can be used for refined biostratigraphy and correlation in the late Ordovician and early Silurian of central Wales. The progressive delay in the insertion of the median septum in these taxa may have wider application for the correlation of the interval immediately after the Hirnantian glacial maximum.     

Population structure of graptolite assemblages

Graptolite rhabdosomes display a diverse suite of morphologies. The range of morphotypes present within most moderate- to high-diversity assemblages from the Ordovician and Silurian is similar, despite the different taxonomic composition of the faunas at different times. Survivorship analyses of graptolite faunas from the Ordovician and Silurian demonstrate strong similarities in the mortality rates of unrelated graptolites of similar functional morphology. They also show a strong correlation between decreasing mortality rates among more mature colonies with increasing rhabdosome complexity. This similarity in both functional morphology and life history of graptolites suggests that they lived within a very stable planktic community structure.     

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.     

Dictyonema Hall and its importance for the evolutionary history of the Graptoloidea

Dictyonema retiforme has been regarded as the benthic ancestor to the planktic Graptoloidea, represented by the earliest planktic Rhabdinopora flabelliformis and its descendants. The revision of the type material of Dictyonema retiforme, the type species of the genus Dictyonema, from the Silurian of New York State shows compound stipes formed by the complexly growing and overlapping tubular thecae of acanthograptid type. The connections between adjacent stipes are formed by thecal tubes or thecal bridges and not by dissepiments. Thus, the species has to be transferred to the Acanthograptidae and cannot be regarded as being related to the early planktic Graptoloidea. The tubarium meshwork of Dictyonema is phylogenetically unrelated to the meshwork of the planktic Rhabdinopora, and represents a case of convergent evolution. The origins of the planktic graptoloids lie among members of middle to upper Cambrian Dendrograpidae, as can be seen from the thecal style and the triad budding patterns with regularly developed bithecae in this group.     

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.