Middle Telychian (upper Llandovery, Silurian) graptolites from boreholes of northwestern Libya: Their biostratigraphic significance and palaeogeographical implication

A low-diversity but stratigraphically and palaeobiogeographically important graptoloid association comprising Metaclimacograptus flamandi (Legrand), Parapetalolithus meridionalis (Legrand), and Torquigraptus australis Štorch was identified in the core samples of wells A1-66, F1-66 and A1-43 in the Ghadamis Basin of Libya. The former two species have been originally recovered from Algeria. All three taxa are common and widespread in the middle Telychian strata of Spain (Central Iberian Zone and western Iberian Cordillera) and France (Brittany). In European sections, the species are associated with other, moderately diverse graptoloid fauna, which enables a more precise and worldwide biostratigraphic correlation of Saharan boreholes and surface sections. The present faunal association indicates a middle Telychian age (upper crispus, griestoniensis and lowermost crenulata/tullbergi biozones) for the strata and suggests palaeobiogeographical links between North African pericratonic basins and Ibero-Armorican shelves. M. flamandi has not been found outside this NW-Gondwanan realm.     

Late Ordovician-Early Silurian fluctuations in sea level from eastern Anticosti Island, Quebec

Over 1,000 m of Upper Ordovician to Lower Silurian mixed carbonate and clastic strata on Anticosti Island are nearly tectonically undisturbed, despite their proximity to the Northern Appalachians fronting Quebec's Gulf of St. Lawrence. Natural cliffs exposed along the coast and rivers in the eastern part of the island make a relatively conformable sequence belonging to the Ashgill and Llandovery Series. Fossil communities interpreted as depth-associated in life are especially repetitious in the Becscie, Gun River, Jupiter, and Chicotte Formations (Llandovery Series), and to a lesser degree in the Upper Vaureal and Ellis Bay Formations (Ashgill Series). Preliminary study of the pattern of changeovers in Eocoelia, Pentamerus and Stricklandia communities suggests that Anticosti seas deepened and shallowed three and a half times during the Early Silurian. High water peaks were reached during B₁-B₂, C₁-C₂ and C₄-C₅ times, with a final deepening trend beginning in late C₅ time. Age determinations of these events are based on the occurrence of graptolites (with some new records from Anticosti) keyed to the standard graptolite zones, and species of the Eocoelia lineage are also useful for correlation. The profile of the Anticosti sea-level curve compares well with other curves reconstructed from the Lower Silurian of New York, Michigan, and Iowa. Widespread synchronism in sea-level changes on the North American platform is thus corroborated.     

Paleoecologic,temporal, and spatial analysis of early Silurian Reefs of the Chicotte Formation,Anticosti Island,Quebec, Canada

Summary Reefs of the Lower Silurian Chicotte Formation are the largest and most faunally diverse known on Anticosti Island, Quebec. They reach up to 25 m in thickness and 250 m in diameter and are present predominantly at two intervals, forming a lower and upper reef cluster. Remnants of bioherms are represented on the present-day wave-cut terrace as 60 to 100 m diameter, subcircular erosional depressions known as Philip structures or as outcrop. The bioherms were relatively low structures, with approximately 3 to 5 m maximum synoptic relief, some of which developed on hardgrounds and possible paleokarst surfaces of crinoidal wackestone and packstone. Dominant skeletal framework builders and sediment producers within all of the reefs are laminar to low domical stromatoporoids, colonial cerioid and fasciculate rugose corals, colonial tabulate corals, and cryptostome bryozoans. Vertical zonation of reef biota is evident within well-exposed reefs of the lower reef cluster. Three to four stages are recognizable:1) a low-diversity tabulate coral-dominatedpioneering community including large tabulate coral colonies (halysitids and favositids), and few stromatoporoids (clathrodictyids, ecclimadictyids), fasciculate rugosans, large generally monotypic stalked crinoids, and shelly benthos (brachiopods, few ostracodes and trilobites);2) an intermediate- to high-diversity, mixed tabulate coral-stromatoporoid-dominatedreef-core community;3) a slightly lower diversity stromatoporoid-tabulate coral-dominatedclimax community with laminar coenitids and alveolitids; and,4) in a few localities, a capping, low-diversity tabulatecoral-dominated (alveolitid and coenitid), and stromatoporoid-bearing community comprising laminar forms. Amelioration of Early Silurian climates, following Late Ordovician glaciation, allowed gradual reestablishment of extensive shallow-water reef growth, by mainly new and increasingly diverse genera and species of metazoans. Reef development within the Chicotte Formation coincided with global, widespread development of latest Llandovery and earliest Wenlock reefs in subtropical to tropical areas. Chicotte reefs have broad characteristics, in terms of overall biotic composition, vertical successions recognized, and paleogeographic setting, similar to those of equivalent and slightly younger age from intracratonic settings in Baltica (Gotland, Sweden and Estonia) and central and northern Laurentia (Midcontinent, U.S.A.; Hudson Bay, Canada; and North Greenland, Denmark).     

The trace fossil Sinusichnus from the Upper Ordovician of Anticosti Island,eastern Canada

Branched, sinusoidal burrows assigned to Sinusichnus cf. seilacheri Knaust et al. 2016; occur in Upper Ordovician (Hirnantian) carbonates on Anticosti Island. These are abundant and monospecific at the base of the Oncolite Bed of the uppermost Ellis Bay Formation, marking a regional discontinuity surface during the multiphase Ordovician mass-extinction event. In contrast to Mesozoic and Cenozoic records of Sinusichnus, commonly attributed to the activity of decapod and isopod crustaceans, the burrow systems from Anticosti Island were probably produced by other arthropods of unknown affinity. A combined dwelling, locomotion and feeding behaviour of their trace maker is assumed. This is the oldest record of the ichnogenus Sinusichnus, previously only known since the Triassic, and its first evidence from North America.     

A New Endosymbiont in Late Ordovician Tabulate Corals from Anticosti Island,Eastern Canada

Cylindrical tubes of the trace fossil Chaetosalpinx occur within the skeletal walls of Late Ordovician sarcinulid tabulate corals preserved on Anticosti Island. A large host-specific embedment structure, Chaetosalpinx rex isp. nov., is described from the reefal sarcinulid genus Columnopora that occurs in (Rawtheyan, Ashgill) coral-stromatoporoid patch reefs of the upper Vauréal Formation (Mill Bay Member). While locally abundant in Columnopora, this newly described embedment structure is absent in other frame-builders in the reefs.     

Peridermal and interthecal tissue in Silurian retiolitid graptolites: with examples from Sweden and Arctic Canada

Peridermal tissue, and thecal 'floors' are shown to be present in some specimens of Pseudoretiolites, Retiolites and Stomatograptus. The development of a dense reticulum and thecal hoods is also recognized as being a progressive developmental feature in Gothograptus. It is suggested that all these structures, whether internal or external, are the product of mature or old-age development. It appears questionable whether the Rhabdopleura model of rhabdosomal development adequately explains late-stage membranes and other adventitious tissues such as discussed herein.     

Sheinwoodian (Silurian) conodonts and graptolites from NE Anti‐Atlas,Morocco

Männik, P., Loydell, D.K. & Lubeseder, S. 2010: Sheinwoodian (Silurian) conodonts and graptolites from NE Anti‐Atlas, Morocco. Lethaia, Vol. 44, pp. 410–416. Conodonts and graptolites from a limestone at the base of the Tamaghrout Formation indicate that this is of early Wenlock age. This is the first unequivocal dating of a Sheinwoodian limestone from Morocco. All of the conodonts are known also from several other regions and suggest that there was no major difference in conodont faunas in northern Gondwana, Baltica and Laurentia at this time. Biostratigraphy, conodonts, graptolites, Morocco, Silurian, Sheinwoodian.     

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.     

Mustelids from Sackdilling Cave (Bavaria,Germany) and their biostratigraphic significance

Sackdilling Cave is a karst fissure filled with a fossiliferous breccia including abundant fossil remains, mostly snails and small mammals. The taxonomic status of mustelid remains from this cave was previously regarded as controversial and, consequently, different authors listed different species from the locality. A detailed revision of the material shows the presence of five mustelid species: Meles sp., Martes vetus, Mustela strandi, Mustela palerminea, and Mustela praenivalis. Among them, especially noteworthy is the presence of the type specimen of Martes vetus. The Sackdilling holotype resembles Martes foina and differs from that of Martes martes in a short and broad viscerocranium; wide and less extended forward incisor row; compressed, short and wide temporal region; large, strongly inflated and convex tympanic bullae; and narrow P3 with weak lingual bulge. Some dental characters are specific for Martes vetus and show some intermediate values between Martes foina than Martes martes. Among them are the P4 protocone length and the M1 trigon length. The revised material from Sackdilling Cave of Mustela palerminea and Mustela praenivalis was compared with that from other Early and early Middle Pleistocene sites of Europe and showed the presence of intermediate characters. Subsequently, based on the entire mustelid assemblage and other faunal elements (mainly rodents), the age of this fauna was estimated as ca. 0.9–0.7 Ma.     

Llandoverian and Wenlockian Cyrtograptus,and some other Wenlockian graptolites from Northern and Arctic Canada

Llandoverian and Wenlockian graptolite zonesin northern and Arctic Canada comprise the Sakmaricus-Laqueus Zone of latest Llandoverian age, Centrifugus Zone of earliest Wenlockian age, Rigidus Zone of about mid-Wenlockian age, possibly the Perneri Zone of late Wenlockian age, and the Testis-Lundgreni Zone of very late Wenlockian age.Species of Cyrtograptus described and illustratedin this study are C. centrifugus, C. cf. insectus, C. aff. lapworthi, C. laqueus, C. lundgreni, C. sp. (aff. C. lundgreni), C. mancki, C. radians, C. rigidus, C. sakmaricus, C. solaris n. subsp.? and C. sp. (similar to C. murchisoni bohemicus). Other taxa comprise Monograptus cf. capillaceus, M.? aff. deubeli, Gothograptus nassa, Plectograptus praemacilentus, Spinograptus cf. spinosus and Retiolites nevadensis.     

Aeronian (Llandovery, Lower Silurian) palynomorphs and graptolites from the Lipeón Formation, Eastern Cordillera, north-west Argentina

The lower levels of the Lipeón Formation, in the Eastern Cordillera, north-west Argentina, yield a marine-dominated palynomorph assemblage, together with graptolites of mid to late mid Llandovery age (Demirastrites convolutus and probably Stimulograptus sedgwickii zones). The palynomorph assemblage is dominated by acritarchs, but also contains algae and terrestrial cryptospores. Crassiangulina variacornuta, considered a potentially good global biostratigraphical marker for the Upper Llandovery is recovered for the first time from the Silurian of Argentina. The occurrence of this species in strata not younger than late Aeronian, and independently dated by graptolites, indicates an early first appearance for Crassiangulina variacornuta, in the Lipeón Formation, below the Aeronian/Telychian boundary. The lower part of the unit corresponds to a quiet marine environment; thus supporting that Crassiangulina variacornuta is a facies-sensitive acritarch.     

Devonian and Carboniferous dendroid graptolites from Belgium and their significance for the taxonomy of the Dendroidea

Devonian and Carboniferous dendroid graptolites from Belgium are evaluated and partly revised. New finds in two different stratigraphic intervals of the ‘Carrière de Lompret’, an active quarry exploiting Frasnian limestones and shales east of Frasnes-lez-Couvin, allow the identification of Callograptus sp. and Dictyonema fraiponti, both belonging to the dendroid family Acanthograptidae. The relatively high diversity of the dendroid graptolite fauna from the Viséan Marbre noir de Denée, one of the few Carboniferous graptolite faunas in the world, can be shown to be based on astogenetic and preservational aspects. Nearly all known specimens can be included in the highly variable Dictyonema fraiponti, a fan-shaped large dendroid species with complex stipes formed from tubular thecae, possessing simple to complex bridges connecting adjacent stipes. Some of the graptolite material is well preserved and provides important information on the tubarium construction of Devonian to Carboniferous dendroid graptolites and, thus, is highly significant for a taxonomic and phylogenetic understanding of the youngest dendroid graptolite faunas worldwide. The genera Callograptus and Ptiograptus are revised based on their type species (Callograptus elegans from Quebec, Canada; Ptiograptus percorrugatus from the Silurian or Devonian of Kentucky, USA) and referred to the Acanthograptidae.     

Plant assemblages from the Silurian of southern Bolivia and their palaeogeographic significance

Abundant but fragmentary plant fossils are described from two locations in shallow water marine facies of the Lipeón (previously Kirusilla) Formation of southern Bolivia. Field relationships and limited palaeontological data suggest that the rocks are of Ludlow to possibly early P'ídolí age (i.e. late Silurian). The majority of the fossils are sterile coalified compressions or impressions of parallel-sided axes, some with branching typical of Hostinella . No tracheids have been found and such remains are best described as rhyniophytoid. Fragments with irregular branching and variable axial diameters probably belong to algae with some similarities to Hungerfordia and Buthotrephis . Rarely axes terminate in clearly delimited globular or elliptical swellings that are interpreted as sporangia, although no spores have been recorded. The most completely preserved specimens have dichotomous branching ending in predominantly elliptical sporangia with distal borders and closely resemble Cooksonia ca-ledonica . Solitary isolated sporangia are vertically elliptical (cf. Tarrantia ), globose (cf. C. cambrensis, C. hemi-sphaerica ) or laterally extended (cf. C. pertoni ). Those with cup- or funnel-shaped morphologies superficially resemble the rhyniophytoid Steganotheca or dyad-containing Culullitheca . Thus while it is impossible to compare with confidence the taxonomic composition of Bolivian assemblages with coeval ones, their overall morphological grade is closer to material collected from circum-northern Atlantic localities than from assemblages in Australia and Kazakhsta/nChina. Palaeogeographically this translates into floristic similarities between Gondwanan high latitudes and equatorial Laurussia rather than with low latitude, north-eastern Gondwana or with a low latitude Kazakhstan/ Xinjiang micro-palaeocontinent     

New occurrences of Ichniotherium and Striatichnium from the Lower Permian Kildare Capes Formation,Prince Edward Island,Canada: palaeoenvironmental and biostratigraphic implications

Abstract: Tracks and trackways of the vertebrate ichnotaxon Ichniotherium sphaerodactylum and a trace of the invertebrate ichnotaxon Striatichnium bromackerense are described for the first time in association outside of Europe. The tracks are identified as I. sphaerodactylum based on their characteristic rounded digit ends, the ovoid sole‐pad of the pedal imprint and the increase in digit lengths from digits I–IV, and the invertebrate trace is identifiable as S. bromackerense based on the band‐like systems of distally bifurcated striae. The tracks of I. sphaerodactylum are the largest known to date and represent rare evidence of large‐bodied terrestrial vertebrates in the Kildare Capes Formation of Prince Edward Island, Canada. Morphometric differences between ichnospecies of Ichniotherium and other ichnospecies previously collected from Prince Edward Island are examined in a multivariate analysis, and results suggest that the distance between the manus and pes of the same imprint pair, the width of pace and the length of certain digits is useful for species identification. The association between I. sphaerodactylum and S. bromackerense was previously known only from the Bromacker quarry, Tambach Formation, Germany, which is interpreted as a seasonally dry, semi‐arid upland environment. The co‐occurrence of these traces suggests that the Eldon locality of Prince Edward Island is similar in depositional environment to the Bromacker quarry in Germany, and allows for comparison of these two localities for the first time. As diadectids are thought to be the trackmakers of Ichniotherium, the Eldon locality of the Kildare Capes Formation may, with further work, be considered as another example of the rare, herbivore‐dominated palaeoenvironment, which is generally uncharacteristic of the Early Permian of North America.     

Paleogene ostracodes from the Dawenkou Basin,East China and their biostratigraphic significance for the age of mineral resources

The Dawenkou Basin is a Cenozoic terrestrial fault basin in Shandong Province, East China, containing abundant mineral resources, especially gypsum. Recently a borehole was drilled in this basin unearthing the upper and middle members of the Dawenkou Formation. From the upper member, well-preserved ostracodes were discovered, which are entirely non-marine taxa including species of Eucypris, Caspiolla, Candona and Candoniella. Based on the new material, a new species Candona dawenkouensis n. sp. was erected. The ostracode assemblage biostratigraphically indicates an age of middle Eocene to Oligocene of the upper member of the Dawenkou Formation and the strata bearing mineral resources (mainly gypsum) of the middle member of the Dawenkou Formation is likely early Eocene.     

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.      

Early Eocene orthophragminids from the Lower Indus Basin,Pakistan, and their biostratigraphic implication

Orthophragminids are a key biostratigraphic proxy for Paleocene–Eocene sequences in western Tethys, but poorly known in eastern Tethys. The orthophragminids were previously misinterpreted as Paleocene's orbitoidiform foraminifera in the Indus Basin, Pakistan. This study focuses on detailed taxonomy and biostratigraphy of orthophragminids. Nine species/subspecies of orthophragminids are identified, including Discocyclina ranikotensis, D. archiaci bakhchisaraiensis, D. a. ex. interc. staroseliensis bakhchisaraiensis, D. dispansa hungarica, D. d. taurica, D. d. broennimanni and Orbitoclypeus schopeni ramaraoi, and two new species Discocyclina pseudoranikotensis n. sp. and Discocyclina debalensis n. sp. are described. The identified orthophragminid taxa represent the orthophragminid zones OZ2-3, corresponding to the shallow benthic zones SBZ5-7 of early Ypresian (early Eocene), which are correlated with those in western Tethys. However, we found that the first appearances of D. d. hungarica and D. d. taurica occur in the OZ3 or SBZ7 in this study instead of the OZ5 or SBZ10 as in western Tethys. The possible paleoecological setting for the occurrences of recognized species is interpreted as inner to middle shelf.     

Stratigraphical setting of the Silurian thelodonts from Prince of Wales Island, Northwest Territories

Additional and corrected information is given regarding the stratigraphy and palaeoecology of the thelodont-bearing sequence on Prince of Wales Island. The thelodont fauna is not older than latest Wenlockian.     

Geographic biotype and host-associated local adaptation in a polyphagous species, Lambdina fiscellaria (Lepidoptera: Geometridae) feeding on balsam fir on Anticosti Island, Canada

The debate about mechanisms underlying the evolution of host specialization by herbivorous insects remains open. Natural selection may act locally and lead to different patterns of geographic variation in life history traits of polyphagous herbivores. The hypothesis of genetically-based trade-offs in offspring performance on different hosts has been proposed but this has rarely been demonstrated. Under laboratory conditions, the biological performance of two populations of the hemlock looper Lambdina fiscellaria (Guenée), a highly polyphagous lepidopteran, was compared when reared on three different tree host species: balsam fir, eastern hemlock and sugar maple. One population originated from Anticosti Island, Québec, Canada, where the insect has evolved without having access to two of the three tree species tested, the other being from the mainland where all tree species are present. When reared on balsam fir foliage, which was naturally available to each population, larvae from Anticosti Island underwent four instars compared with five for the mainland population, indicating the existence of geographic biotypes in L. fiscellaria. When reared on the foliage of non-naturally available host trees, larvae from Anticosti Island had a higher incidence of supernumerary instars. This is a unique example where local adaptation to environmental conditions of an insect herbivore is expressed through a differential number of larval instars. Moreover, the Anticosti Island population showed a higher growth related index on the host available to both populations indicating that a fitness trade-off was the evolutionary process underlying the local adaptation of this population on balsam fir.     

Latest Permian palynomorphs from Jharia Coalfield,Damodar Basin,India and their potential for biostratigraphic correlation

The objective of the present work is to palynologically date the samples collected from Jharia Coalfield. The samples have been collected from an outcrop section exposed across the Damodar river near NH 32 enroute from Bokaro to Dhanbad (23° 43′06.93″N and 86° 12′06.90″E) near Jharia Coalfield of Damodar Basin. Palynological analysis of the samples has revealed that the palynomorph assemblage is characterized by high incidence of Densipollenites spp. in association with dominance of striate bisaccates chiefly Striatopodocarpites spp. and Faunipollenites spp. The other stratigraphically significant taxa recorded in the section are Guttulapollenites hannonicus, Klausipollenites schaubergeri, Alisporites spp. Falcisporites spp. and Lunatisporites spp. On the basis of the palynocomposition the section has been assigned a latest Permian age. Age determination for the section has been inferred by comparison with coeval assemblages from previous Gondwana studies from India and across Gondwana which were compared to the biostratigraphically dated palynological records from Australia (oldest APP6 Zone) as it is one of the few areas in Gondwana where the palynological records have been calibrated against adequately dated marine invertebrate zones. However, comparison with the westernmost portion of Gondwana was difficult due to its mid latitudinal position (c. 20°S) during the Lopingian (late Permian) which lead to an early transition from coal-bearing strata to red-bed and evaporitic deposits in some parts.