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.     

Evolution in Monograptus priodon

Biometric analysis of the Silurian graptolite Monograptus priodon (Bronn) from Arctic Canada and the British Isles shows that the length of the sicula and the rhabdosome width at the level of theca 5 undergo a progressive increase from latest Llandoverian to late Wenlockian time. This change is attributed to sympatric evolution within the species. Monograptus flemingii (Salter) may have evolved elsewhere from priodon stock and migrated later into the two regions. Monograptus riccartonensis Lapworth probably ranges from latest Llandoverian through the Wenlockian, and may have been ancestral to the Ludlovian to Devonian Monograptus lineages.     

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.     

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     

Ludlovian,Pridolian and Lochkovianin la Meignanne (Massif Armoricain): Biostratigraphy and correlations based on Bivalvia and Chitinozoa

Black carbonate rocks («ampelitic limestones)exposed in la Meignanne, NW of Angers, Massif Armoricain, as tectonic lenses within a complex tectonic structure of grey Pragian limestones are assigned from Bivalvia and Chitinozoa to the Upper Silurian and Lowermost Devonian. Biostratigraphical and paleoecological analyses show that the Silurian-Devonian boundary in this area is developed in offshore facies and that it may be correlated directly with the international stratotype of the Prague Basin (Barrandian, Bohemia). Paleogeographical communication between the two areas is indicated by 28 species of bivalves in common from two Pridolian assemblages (with Cheiopteria bridgei and with Snoopyia insolita) and one Lochkovian (with Antipleura bohemica). Stratigraphical correlation with the Prague Basin is also based on the chitinozoan zonal index species for the Uppermost Pridolian (Urnochitina urna) and Lowermost Lochkovian (Eisenackitina bohemica). A Lochkovian age is further supported by the discovery of Monograptus uniformis uniformis in the assemblage with Antipleura bohemica. A Ludlovian age is indicated by the occurence of «Hemicadium elevatum and an abundance of the ostracode Entomozoe (Richteria) migrans, which is an index species for the Middle Ludlovian in Bohemia.     

THE THECAL STRUCTURE OF MONOCLIMACIS? GALAENSIS

Monoclimacis? galaensis (Lapworth), a biform Llandoverian graptolite, is interpreted as having proximal thecal ‘hooks’ composed of paired lateral lappets, and lacking retroversion of the dorsal thecal wall. A parallel is drawn between M? galaensis and members of the Ludlovian genus Lobograptus Urbanek. The significance of the thecal structure is discussed with reference to the principles involved in the classification of the monograptids.     

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.     

新疆巴里坤三塘湖志留纪三叶虫

新疆巴里坤三塘湖晚志留世阿苏山组三叶虫包括Ananaspis fecunda Barrande,Cheirurus barkolensis sp.nov.和Pacificurus sp.,与准噶尔盆地西北部乌图布拉格,哈萨克斯坦及捷克波西米亚的晚志留世三叶虫关系密切。另外对中国5个地区晚志留世三叶虫地层作了对比。Ananaspis,Pacificurus二属在中国是初次发现。     

A Wenlock ancestor of the micropoma phylogenetic lineage

A new graptolite species, Monoclimacis praemicropoma , from the Upper Wenlock of Poland is described. It is an ancestor of the Lower Ludlow M. micropoma (Jaeckel). The phytogeny of the praemicropoma-micropoma lineage is discussed. M. micropoma nannopoma (Jaeger) has been assigned to a separate evolutionary lineage. Graptolites, Wenlock, phytogeny .     

Enantiomorphous dimorphism in Silurian and Devonian bivalves; Maminka Barrande, 1881 (Lunulacardiidae, Silurian) - the oldest known example

The genus Maminka Barrande, 1881 (Lunulacardiidae) from the Wenlock and Ludlow (Silurian, Lower Paleozoic) of Europe is the oldest known bivalve with enantiomorphous dimorphism which is common among the Silurian and Devonian Lunulacardiidae and Antipleuridae. It represents the result of a special adaptation to the reclining mode of life combined with byssal attachment when early in ontogeny either the left or right valve became the lower valve; later, the bivalve develops as two enantiomorphous sets of shells. Distribution, ontogeny and evolution of this interesting feature in the genera Maminka Barrande, 1881, Stolidotus Hede, 1915, Mila Barrande, 1881, Dualina Barrande, 1881, Antipleura Barrande, 1881, Silurina Barrande, 1881 and Hercynella Kayser, 1878 are discussed.     

THE DEVELOPMENT OF THE LUDLOVIAN GRAPTOLITE SAETOGRAPTUS VARIANS

The Ludlovian graptolite Saetograptus varians (Wood 1900) has been isolated from a limestone matrix. The thecal structure indicates that the species is closer to the genus Saetograptus Přibyl 1942, emend. Urbanek 1958, than to Colonograptus Přibyl 1942, emend. Urbanek 1958; but it requires that the definition of the former be amended to include Colonograptus as a junior synonym.     

Le genre leptaenopyxis (brachiopoda,leptaenidae) dans le Devonien du Massif Armoricain (France)

The study of genus LeptaenopyxisHavlicek, 1963 in the Lower Devonian from Armorican Massif allows to recognize four species: Leptaenopyxis kerfornei n. sp., Leptaenopyxis cf. bouei (Barrande), Leptaenopyxis sp. 1 and Leptaenopyxis sp. 2. These species were described, until now, under specific name boueiBarrande, 1848. From this study it follows some biostratigraphical and palaeobiogeographical results for the Armorican Massif and its relations with surrounding areas during Lower Devonian time.     

广西早泥盆世节甲鱼类及胴甲鱼类化石及华南早泥盆世脊椎动物生物地层对比

描述了早泥盆世的3个属种,其中节甲鱼类Yujiangolepisliujingensisgen．etsp.nov.采自广西六景,Buchanosteussp．及胴甲鱼类？Chuchinolepissp.采自广西玉林。该3件标本均产自海相地层中,与海相无脊椎动物化石共生。还讨论了华南区几个主要产脊椎动物化石（包括脊椎动物微体化石）剖面的生物地层对比。     

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.     

LOWER DEVONIAN GRAPTOLITES FROM SOUTHEASTERN ALASKA

The youngest zones of Monograptus occur in Devonian black shale and slate along the western shore of Prince of Wales Island and on Noyes Island, southeastern Alaska. The following five species of Monograptus, in close association with vascular plants and marine shelly faunas, indicate a Praguian, (Siegenian to Early Emsian), late Lower Devonian and possibly early Middle Devonian age: M. yukonensis Jackson & Lenz, M. n. sp. aff. yukonensis ? Jaeger & Stein, M. aeqimbilis notoaequakilis Jaeger & Stein, M. pacificus Jaeger n. sp. and M. craigensis Jaeger n. sp. The yukonensis zone containing M. craigensis in its upper part is followed stratigraphically higher by the M. pacificus zone – the youngest occurrence of Monograptus. The new species belong to the M. uncinatus group and, like most Praguian monograptids, have long dorsal tongues on their siculae. M. craigensis and M. pacificus increase to seven the number of species of Monograptus known from the Praguian. On the basis of the Alaskan material, the geographical distribution of M. aequabilis notoaequabilis now includes North America in addition to previously recorded occurrences in Australia, Southeast Asia, and Europe. This broad geographic distribution suggests that the last survivors of Monograptus had worldwide distribution and a rather uniform assemblage of species.     

The oldest Gondwanan cephalopod mandibles (Hangenberg Black Shale,Late Devonian) and the mid‐Palaeozoic rise of jaws

It is widely accepted that the effects of global sea‐level changes at the transition from the Devonian to the Carboniferous are recorded in deposits on the shelf of northern Gondwana. These latest Devonian strata had been thought to be poor in fossils due to the Hangenberg mass extinction. In the Ma'der (eastern Anti‐Atlas), however, the Hangenberg Black Shale claystones (latest Famennian) are rich in exceptionally preserved fossils displaying the remains of non‐mineralized structures. The diversity in animal species of these strata is, however, low. Remarkably, the organic‐rich claystones have yielded abundant remains of Ammonoidea preserved with their jaws, both in situ and isolated. This is important because previously, the jaws of only one of the main Devonian ammonoid clades had been found (Frasnian Gephuroceratina). Here, we describe four types of jaws of which two could be assigned confidently to the Order Clymeniida and to the Suborder Tornoceratina. These findings imply that chitinous normal‐type jaws were likely to have already been present at the origin of the whole clade Ammonoidea, i.e. in the early Emsian (or earlier). Vertebrate jaws evolved prior to the Early Devonian origin of ammonoids. The temporal succession of evolutionary events suggests that it could have been the indirect positive selection pressure towards strong (and thus preservable) jaws since defensive structures of potential prey animals would otherwise have made them inaccessible to jawless predators in the course of the mid‐Palaeozoic marine revolution. In this respect, our findings reflect the macroecological changes that occurred in the Devonian. [Correction added on 28 July 2016 after first online publication: In the Abstract, the sentence “Vertebrate jaws probably … in the Early Devonian” was amended]     

400 million years on six legs: On the origin and early evolution of Hexapoda

Identifying the unambiguous sister group to the hexapods has been elusive. Traditional concepts include the Myriapoda (the Tracheata/Atelocerata hypothesis), but recent molecular studies consistently indicate it is the Crustacea, either in part or entirety (the Pancrustacea/Tetraconata hypothesis). The morphological evidence in support of Tracheata is reviewed, and most features are found to be ambiguous (i.e., losses, poorly known and surveyed structures, and probable convergences), though some appear to be synapomorphic, such as tentorial structure and the presence of styli and eversible vesicles. Other morphological features, particularly the structure of the eyes and nervous system, support Pancrustacea, as does consistent molecular evidence (which is reviewed and critiqued). Suggestions are made regarding hexapod–crustacean limb homologies. Relationships among basal (apterygote) hexapods are reviewed, and critical Paleozoic fossils are discussed. Despite the scarceness of Devonian hexapods, major lineages like Collembola and even dicondylic Insecta appeared in the Early Devonian; stem-group and putative Archaeognatha are known from the Carboniferous through Permian and the Late Devonian, respectively. Thus, the earliest divergences of hexapods were perhaps Late Silurian, which is considerably younger than several estimates made using molecular data.