Patterns of evolution and extinction in proetid trilobites during the late Devonian mass extinction event,Canning Basin,Western Australia

Abstract: In the early Late Devonian, terminal Frasnian proetid trilobites have previously only been known from Europe and North Africa. For the first time, a rich fauna of late Frasnian proetids is described from the Virgin Hills Formation, Canning Basin, Western Australia. Seventeen species in six genera are described, of which three are new: Rudybole gen. nov., Palpebralina gen. nov. and Canningbole gen. nov. A new subgenus, Chlupaciparia (Australoparia) subgen. nov. is also described. Fourteen of the species are new: Palpebralia initialis sp. nov., P. pustulata sp. nov., ?P. sp. nov. A, Rudybole depressa sp. nov., Palpebralina pseudopalpebralis sp. nov. (comprising the subspecies P. pseudopalpebralis pseudopalpebralis subsp. nov. and P. pseudopalpebralis ultima subsp. nov.), P. minor sp. nov., P. ocellifer sp. nov., Canningbole latimargo sp. nov., C. henwoodorum sp. nov., C. macromma sp. nov., Pteroparia extrema sp. nov., Chlupaciparia (Chlupaciparia) planiops sp. nov., Chlupaciparia (Australoparia) australis sp. nov. and C. (Australoparia) lata sp. nov. The subspecies Rudybole adorfensis angusta subsp. nov. is also described. The proetids range through conodont Zones 11–13b and terminate at the Upper Kellwasser Event, which marks the terminal Frasnian mass extinction event. Three of the six proetid lineages, Palpebralia, Canningbole and Pteroparia, show evolutionary trends of eye reduction. Two of the remaining lineages, Rudybole and Palpebralina, consist exclusively of blind taxa. The last, Chlupaciparia, also comprises forms with reduced eyes. The proetids show a stepped pattern of extinction during the late Frasnian, which correlate with two Kellwasser biocrises documented in European/North African Frasnian sections. The highest diversity preceded the Lower Kellwasser event that occurred at the end of conodont Zone 12 and saw the extinction of all species present in that zone. However, only one genus, Pteroparia, locally became extinct. A major higher‐level taxonomic mass extinction at the top of Zone 13b initiated the Upper Kellwasser extinction event. This included extinction at the generic level, with all five remaining genera becoming extinct, and at the family level, with the loss of the Tropidocoryphidae.     

The fate of the homoctenids (Tentaculitoidea) during the Frasnian–Famennian mass extinction (Late Devonian)

The homoctenids (Tentaculitoidea) are small, conical‐shelled marine animals that are among the most abundant and widespread of all Late Devonian fossils. They were a principal casualty of the Frasnian–Famennian (F‐F, Late Devonian) mass extinction, and thus provide an insight into the extinction dynamics. Despite their abundance during the Late Devonian, they have been largely neglected by extinction studies. A number of Frasnian–Famennian boundary sections have been studied, in Poland, Germany, France, and the USA. These sections have yielded homoctenids, which allow precise recognition of the timing of the mass extinction. It is clear that the homoctenids almost disappear from the fossil record during the latest Frasnian ‘Upper Kellwasser Event’. The coincident extinction of this pelagic group, and the widespread development of intense marine anoxia within the water column, provides a causal link between anoxia and the F‐F extinction. Most notable is the sudden demise of a group, which had been present in rock‐forming densities, during this anoxic event. One new species, belonging to Homoctenus is described, but is not formally named here.     

Trilobites Asteropyginae dévoniens d’Iran

Ten species of trilobites belonging to 4 genera of the sub-family Asteropyginae DELO are described from the Devonian (Givetian-Frasnian) of Iran. Three of which are new: Neocalmonia lutensis nov. sp., Neocalmonia yazdii nov. sp. and Radiopyge sardarensis nov. sp. The study of this material have required the revision of Heliopyge sharudensis(PILLET, 1974) from the Givetian of the Eastern Alborz Mountains and of Neocalmonia quadricosta PILLET, 1969, type-species of the genus Neocalmonia, from the Frasnian of Afghanistan. The recognition of genera and species in common with the Givetian and the Frasnian of Afghanistan emphasizes the close relations between these two regions. The genera Bradocryphaeus, Heliopyge, Neocalmonia, Radiopyge, represented by related species, are likewise present in Iran, Afghanistan and Western Europe. Their presence confirms that these regions belong, during the Devonian, to the North Gondwanian domain.     

First selenosteid placoderms from the eastern Anti‐Atlas of Morocco; osteology,phylogeny and palaeogeographical implications

Abstract: A new placoderm assemblage is reported from the Kellwasser facies of the eastern Anti‐Atlas, Morocco. This is the first record of an abundant Frasnian placoderm assemblage from Africa. The following new selenosteid taxa are described: Draconichthys elegans gen. et sp. nov., Enseosteus marocanensis sp. nov. and Walterosteus lelievrei sp. nov. In addition, material of Rhinosteus cf. parvulus is figured and described for the first time from Gondwana. The genera Enseosteus, Rhinosteus and Walterosteus are reviewed. ‘Wildungenichthys’ is regarded as a junior subjective synonym of Walterosteus. The new material and a revision of the taxa from Bad Wildungen, Germany, are the basis for a new phylogenetic analysis of the selenosteid arthrodires. The monophyly of the selenosteids is confirmed. The North American selenosteids are a sister group of Moroccan and European selenosteids, excluding Pachyosteus. The Moroccan vertebrate assemblage resembles that of Bad Wildungen. This supports the close palaeogeographical positions of Gondwana and Laurussia during the Late Devonian and indicates that biogeographical barriers for placoderms that could have separated the two assemblages were absent.     

Environmentally controlled phyletic evolution, blindness and extinction in Late Devonian tropidocoryphine trilobites

The Middle and Upper Devonian carbonate succession of the Montagne Noire, Southern France has been precisely zoned by an unbroken sequence of conodont zones. Stratigraphic control is excellent, and has allowed evolutionary changes in tropidocoryphine trilobites, which occur throughout the succession, to be directly established. The tropidocoryphine had been a stable group for some 40 million years prior to the Middle Givetian. In their last few million years, however, they underwent rapid evolution and exhibit some striking transformations of the cephalon and the regression and virtual disappearance of the eye within a relatively short space of time. They also show a marked diminution in size, and lose their original relief so that the glabella becomes virtually flush with the surface. There are two separate lineages, both of which show eye-reduction and subsequent blindness. In the earlier lineage Tropidocoryphe (Longicoryphe)-Erbenicoryphe, the main features of the ancestral rootstock are conserved and the stable, strongly divergent anterior sutural pattern of the cephalon remains the same. The eye, however, became reduced to a slightly convex surface lacking lenses, only indistinctly defined. Erbenicorphe is confined to well-oxygenated facies, and probably lived as a shallow burrower within the sediment; it became extinct in the early Frasnian. The second lineage T. (Longicoryphe)-Pterocoryphe- Pteroparia shows a remarkable backward migration of the suture, which progressively swings posteriorly in successive species spanning four Frasnian conodont zones (about three million years duration). At the same time the eye progressively degenerates so that the last forms are blind. Sutural migration and eye reduction are not genetically linked, however; the unusual form of the cephalon and suture probably resulted from an adaptation to the euxinic environment in which Pterocoryphe originated. The loss of the eye resulted from the adoption of an endobenthic habit in Pteroparia which descended from the ancestral Pterocoryphe but which had migrated to an oxygenated facies. Eye-reduction is therefore parallel in the two lineages, but superimposed upon a different original cephalic configuration. The last Pteroparia became extinct when the late Frasnian fauna became overwhelmed by the first pulse of the ‘Kellwasser Event’ (probably an anoxic overturn). The evolving characters, through virtually all observed steps, show progressive unidirectional change without sudden breaks or saltations. Such unidirectional evolution is an adaptive response to constant long-lasting environmental influences. □Evolution, Devonian trilobites, France, gradualism, eye reduction.     

New conodonts of the genus <Emphasis Type="Italic">Polygnathus</Emphasis> from the Evlanovian and Livnian (Upper Devonian) of the Voronezh Anteclise (central Devonian Field)

New conodont species of the genus Polygnathus (P. krutoensis sp. nov., P. makhlinae sp. nov., P. menneri sp. nov., P. obruchevae sp. nov.) are described from the Evlanovian-Livnian (Upper Devonian) deposits of the Voronezh Anteclise (central regions of the Rassian platform). The ontogenetic series of the new species are presented.     

Les vertébrés dévoniens de l'Iran central: IV: Arthrodires et ptyctodontes

The Upper Devonian localities of the Kerman and Tabas area yielded numerous brachythoracid arthrodire remains, referred here mainly to Golshanichthys asiatica nov. gen., nov. sp., and Holonema cf. radiatumObrouchev. Some specimens are suggestive of species belonging to the genera Plourdosteus, Eastmanosteus and Aspidichthys. G. asiatica is a primitive dinichthyid characterized by e.g. very deep posterior and anterior embayments of the central plates.Some ptyctodontid tooth plates of «Rhynchodus and «Ptyctodus type have been found in the Frasnian and Famennian.     

Continuing of the revision of Spriferid (Brachiopoda) zonal species from the Frasnian Stage (Upper Devonian) of the Russian Platform

Adolfia siratschoica (Ljaschenko) and Adolfia krestovnikovi (Ljaschenko) are accepted as zonal species of the Petino and Voronezh Horizons of the Frasnian Stage of the Russian Platform. However it was established that these species do not belong to the genus Adolfia. The former species was recognized as the nomenclatural type species of Ljaschenkovia gen. nov. and the latter species was recognized as the type species of Tokmospirifer gen. nov. The comparative analysis provided of the shell structure, microornamentation, and shell interior of the type species of Ljaschenkovia gen. nov., Tokmospirifer brevis gen. et sp. nov., and Adolfia solita Ljaschenko is presented.     

New Late Devonian strophomenids (Brachiopoda) from Mongolia

Two new strophomenid species, Leptagonia barunkhuraica sp. nov. (Famennian) and Floweria mongolica sp. nov. (Frasnian), from the Upper Devonian of the Barunkhurai Depression of southern Mongolia are described.     

SVALBARDIA BANKSII SP. NOV. FROM THE UPPER DEVONIAN (FRASNIAN) OF NEW YORK STATE

Svalbardia banksii sp. nov., is described from the Upper Devonian (Frasnian) Fish Cabin Creek locality near Pond Eddy, New York. The compression consists of ultimate branches bearing spirally arranged, unwebbed leaves. The unwebbed leaves are up to 3.2 cm long, are somewhat flexuous, and dichotomize in more than one plane. The new species is similar to the reconstruction of Actinoxylon banksii from the Middle Devonian (Givetian) of New York and strengthens the supposed relationship between Svalbardia polymorpha and Actinopodium nathorstii from Mimerdalen, Spitzbergen.     

Upper Devonian Sponges from the Holy Cross Mountains, Central Poland

The rich fauna of Late Devonian (Late Frasnian) siliceous sponges from the Holy Cross Mountains, Poland is composed of 15 species and 11 genera. Both astylospongid demosponges (lithistids) and hexactinosan hexactinellids are present. The following new genera and/or species are proposed: D regulara Rigby and Pisera sp. nov., Jazwicella media Rigby and Pisera gen. et sp. nov., Astyloscyphia irregularia Rigby and Pisera gen. et sp. nov., A. turbinata Rigby and Pisera gen. et sp. nov., Astylotuba modica Rigby and Pisera gen. et sp. nov., Paleoregulara cupula Rigby and Pisera gen. et sp. nov., Paleoramospongia bifurcata Rigby and Pisera gen. et sp. nov., Cordiospongia conica Rigby and Pisera gen. et sp. nov., Paleocraticularia elongata Rigby and Pisera gen. et sp. nov., P gigantia Rigby and Pisera gen. et sp. nov., Polonospongiadevonica Rigby and Pisera gen. et sp. nov., P fistulata Rigby and Pisera gen. et sp. nov., Urnospongia modica Rigby and Pisera gen. et sp. nov., and Conicospongia annulata Rigby and Pisera gen. et sp. nov. The investigated fauna contains the youngest astylospongiids known and the oldest well-preserved, and most diversified Palaeozoic hexactinosans. The sponge fauna constituted a significant element of a brachiopod-coral-sponge assemblage that inhabited a deep slope of the local Dyminy Reef structure, during its final phase of growth, in a clearly hemipelagic setting. This fauna is limited to the intrashelf depression within an incipiently drowned carbonate platform.     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

Brachiopods from the family Ambocoeliidae George (Order Spiriferida) from the Emsian (Lower Devonian) of northeastern Russia

Brachiopods from the family Ambocoeliidae George from the Emsian (Lower Devonian) of northeastern Russia are described: Bisinocoelia despecta sp. nov.; Yakutospirifer gen. nov. with the type species Y. krivensis sp. nov.; Micospirifer gen. nov. with the type species M. simplex sp. nov.; Kolymospirifer gen. nov. with the type species K. optatus sp. nov. (subfamily Ambocoeliinae); Datnospirifer gen. nov. with the type species D. alius sp. nov.; Khalimospirifer gen. nov. with the type species Kh. microscopicus sp. nov.; Gerospirifer gen. nov. with the type species G. normalis sp. nov., and G. gerensis sp. nov.; Dogdospirifer gen. nov. with the type species D. yolkini sp. nov. (subfamily Rhynchospiriferinae).     

Evolution of Devonian biodiversity of radiolarians with two porous spheres and one main spine

Features and major distinctions in the appearance and distribution of radiolarians with two porous spheres and one main spine in the Devonian basins are considered. Four solitary population waves and expansion scenario of radiolarians of this morphotype have been recognized in the Devonian. A new species, Radiobisphaera rozanovi sp. nov., from the Upper Emsian, Upper Eifelian, and Middle Frasnian of the southern Ural Mountains, from the Upper Eifelian of the Prague Basin (Barrandian), from the Middle-Upper Frasnian of the Rudnyi Altai, and from the Lower Famennian of the Timan-Pechora Basin and the Polar Ural Mountains is described.     

New Devonian brachiopods from northeastern Russia

Brachiopods from the Devonian of northeastern Russia are described: Eoprokopia gen. nov., with the type species E. aequalis sp. nov. (subfamily Prokopiinae, order Orthida); Davoustia settedabanica sp. nov. and D. verkhojanica sp. nov. (family Anopliidae, order Chonetida); and Alkhovikovia gen. nov. with the type species A. libera sp. nov. and A. importuna sp. nov. and Tikhyspirifer gen. nov. with the type species T. globosus sp. nov. (subfamily Rhynchospiriferinae, order Spiriferida).     

New Emsian foraminifers from of the Karpinsky Horizon limestones Severouralsk Bauxite Mine

Four new foraminiferal species, Parathuramminites mutilatus sp. nov., P. stelliformis sp. nov., P. subrus sp. nov., and P. minutus sp. nov., and a new genus and species, Algaeformis porosus gen. et sp. nov. from clayey limestones and marls of the basal Karpinsky Horizon (Emsian Stage, Lower Devonian) of the Severouralsk Bauxite Mine are described.     

A new find of the placoderm genus <Emphasis Type="Italic">Bothriolepis</Emphasis> Eichwald in the Upper Devonian of Uzbekistan

A new placoderm species, Bothriolepis sanzarensis sp. nov., from the Upper Frasnian (Upper Devonian) of the western marginal area of the Turkestan Mountain Ridge (Samarkand Region, Uzbekistan) is described. The new species is distinguished by the isometric anterior medio-dorsal plate, with a broad posterior region, and the undulating sutures between this and other trunk plates. This is the second find of Bothriolepididae in Uzbekistan.     

New gall midges of the tribe Leptosynini (Diptera,Cecidomyiidae) from the Late Eocene ambers and the classification of the supertribe Heteropezidi

Three new genera and 27 new species of gall midges are described from the Late Eocene ambers: Henria baltica sp. nov., Frirenia manca sp. nov., F. musicata sp. nov., Leptosyna samlandica sp. nov., L. fastosa sp. nov. from Baltic amber and H. xystica sp. nov., H. liquida sp. nov., Stellasegna vlaskini gen. et sp. nov., S. vaporea sp. nov., S. nexa sp. nov., Rasnitsia verticosa gen. et sp. nov., F. rohdendorfi sp. nov., F. schevchenkoi sp. nov., F. melica sp. nov., F. lukashevichae sp. nov., F. leporidis sp. nov., F. marmarygma sp. nov., F. vesana sp. nov., Vincinescia alisae gen. et sp. nov., L. margarita sp. nov., L. munifera sp. nov., L. sukachevae sp. nov., L. assa sp. nov., L. larga sp. nov., L. vegeta sp. nov., L. vaticina sp. nov., and L. shcherbakovi sp. nov. from Rovno amber. Strobliella capitata Fedotova is redescribed as Henria capitata (Fedotova, 2004) (comb. nov.). Diagnoses of Henria (= Electroxylomyia Nel et Prokop, syn. nov.), Frirenia, and Leptosyna are revised. As a result, Henria comprises 3 extant and 5 extinct (Late Eocene) species, including H. eocenica (Nel et Prokop), comb. nov. (= Electroxylomyia eocenica), Frirenia comprises 1 extant and 10 Late Eocene species, and Leptosyna comprises 3 extant and 11 Late Eocene species. The tribe Heteropezini is elevated to the supertribal rank (Heteropezidi) and included in the subfamily Lasiopterinae. Leptosynini is treated as a separate tribe, and Lasiopterinae is considered as part of Cecidomyiidae s. str. (i.e., excluding Lestremiidae). Keys to the tribes and genera of Heteropezidi and to species of Henria, Stellasegna, Frirenia, and Leptosyna are provided. The gall midge faunas of the Rovno and Baltic ambers are compared. Phylogenetic relationships within the supertribe are hypothesized.     

Orthid brachiopods from the Lower and Middle Devonian of northeastern Russia

Brachiopods of the order Orthida from the Lower and Middle Devonian of northeastern Russia: Skenidium diversus Baranov (family Skenidiidae); Datnia gen. nov., with the type species D. asiatica sp. nov. (subfamily Prokopiinae); Janzhinshinia gen. nov., with the type species J. datnensis sp. nov. (subfamily Isorthinae); Cortezorthis chobotchalensis (Alekseeva) (subfamily Cortezorthinae); Elenia gen. nov., with the type species E. gerensis sp. nov. (family Dicoelosiidae); Seimtchania communis (family Seimtchaniidae); Simakovia gen. nov., with the type species S. rara sp. nov. (family Draboviidae); Schizophoria distenta sp. nov., S. grande Baranov, S. striatula (Schlotheim) (family Schizophoriinae); and Hypsomionia sinsera Baranov (family Hypsomioniidae) are described.