Teruelia diezii gen. et sp. nov.: an early polysporangiophyte from the Lower Devonian of the Iberian Peninsula

A new basal land plant, Teruelia diezii gen. et sp. nov., is described from the shallow‐water marine deposits of the Lower Devonian (Lochkovian–Pragian) Nogueras Formation of the Iberian Peninsula (north Gondwana palaeocontinent). Teruelia is preserved as a compression fossil and consists of isotomously branched, robust stems terminated in large, fusiform, twisted sporangia. This morphology suggests that Teruelia is very probably equivalent to Aglaophyton, a permineralized early polysporangiophyte known up to now only from the Lower Devonian (early Pragian to ?earliest Emsian) Rhynie Chert in Scotland (Laurussia palaeocontinent), which represents an early terrestrial hot‐spring ecosystem. Accepted phylogenies identify Aglaophyton as sister to vascular plants. Our phylogeny‐based results identify the Aglaophyton/Teruelia biological entity (i.e. Aglaophyton anatomical characters plus Teruelia external morphology) as the most direct vascular plant precursor. It shows that at least one Rhynie Chert type plant had a much wider distribution than previously known and suggests that Aglaophyton was not restricted to hydrothermal environments, unlike other Rhynie Chert plants.     

Miospore stratigraphy of Lower and early Middle Devonian deposits from Tidikelt, Central Sahara, Algeria

The Tidikelt region forms an outstanding area for subsurface Lower Devonian stratigraphy in the central Algerian Sahara. Sediments from five boreholes have revealed abundant and diverse assemblages of miospores, acritarchs, chitinozoa, scolecodont and microplant remains. The miospores are moderately well preserved. Three new miospore species (Dibolisporites saharansis nov. sp. Hassan Kermandji, Acinosporites conatus nov. sp. Hassan Kermandji and Scylaspora tidikeltense nov. sp. Hassan Kermandji) are described. Miospore assemblages vary through the regressive and transgressive sequences. Seven miospore assemblage biozones, including six new miospore assemblage biozones (Scylaspora tidikeltense-Perotrilites microbaculatus, Dictyotriletes emsiensis-Emphanisporites spinaeformis, Apiculiretusispora arenorugosa-Camptozonotriletes caperatus, Verrucosisporites polygonalis-Dictyotriletes subgranifer, Emphanisporites annulatus-Geminospora svalbardiae, Hystricosporites microancyreus-Grandispora protea, Calyptosporites velatus-Rhabdosporites langii) are proposed for the Lower and early Middle Devonian rocks of Tidikelt Plateau. The combined use of distinctive, wide distribution cosmopolitan and Gondwanan forms as biozonal and species characteristics permits accurate subdivision, dating and correlation of Tidikelt successions with other similar miospore zones of the Lower Devonian of Europe, Canada and other parts of Gondwana plate. The miospore data provide new explanations to stratigraphic relationships of regional rock units, sedimentary cycles and stratigraphic hiatus. The miospore biozones are proposed as a provincial biozonation, which may also be applied to other Palaeozoic rocks of similar miospore content.     

First report of craniide brachiopods in the Palaeozoic of Iran (Pseudocrania,Ordovician), and Early to Mid‐Ordovician biogeography of the Craniida

Abstract: Lower Ordovician faunas of Bohemia (Perunica), Baltica and North China include the oldest known representatives of the Order Craniida, but otherwise in Gondwana and associated terranes, the record of craniides is sparse. Pseudocrania insperata sp. nov. from the Lashkarak Formation of the Eastern Alborz Mountains is the first and as yet only record of the occurrence of craniides in the Middle Ordovician (Darriwilian) of Iran and temperate to high latitude peri‐Gondwana. Pseudocrania was known hitherto only from the Middle Ordovician of Baltoscandia and the Chu‐Ili terrane of Kazakhstan.     

EARLY DEVONIAN FISHES FROM GUIJIATUN AND XUJIACHONG FORMATIONS OF QUJING,YUNNAN,AND RELATED BIOSTRATIGRAPHIC PROBLEMS

Pale0nt0IogicStudiesSubclassGaleaspidaHalsteadTarl0,l9670rderHuanana8piformesJanvier,l975FamilyGantar0stra8pidaeWangetWang,l992GenusCantarostraspisWangetWang,l992CQntarostra8pisgeniWangetWang,l992(Fig.l3Pl-II,4-5)1992,Gantarostraspisgeni,Wang,J.Q.etWang,N.Z.,Fig-l,Pl.l.HolotypeAdorsalshieldV9758,LowerDevonianPosongchongF0rmation,Gu-mu,Wenshandistrict,Yunnan-ReferredspeeimensAdorsalshieldVlO494.1;arostra1pr0cessV1O494.2.Loca1ityandh0riz0nXujiach0ng,Xishan,Qujing,Yunnan,Lowe…     

A Gigantic Sarcopterygian (Tetrapodomorph Lobe-Finned Fish) from the Upper Devonian of Gondwana (Eden,New South Wales,Australia)

Edenopteron keithcrooki gen. et sp. nov. is described from the Famennian Worange Point Formation; the holotype is amongst the largest tristichopterids and sarcopterygians documented by semi-articulated remains from the Devonian Period. The new taxon has dentary fangs and premaxillary tusks, features assumed to be derived for large Northern Hemisphere tristichopterids (Eusthenodon, Hyneria, Langlieria). It resembles Eusthenodon in ornament, but is distinguished by longer proportions of the parietal compared to the post-parietal shield, and numerous differences in shape and proportions of other bones. Several characters (accessory vomers in the palate, submandibulars overlapping ventral jaw margin, scales ornamented with widely-spaced deep grooves) are recorded only in tristichopterids from East Gondwana (Australia-Antarctica). On this evidence Edenopteron gen. nov. is placed in an endemic Gondwanan subfamily Mandageriinae within the Tristichopteridae; it differs from the nominal genotype Mandageria in its larger size, less pointed skull, shape of the orbits and other skull characters. The hypothesis that tristichopterids evolved in Laurussia and later dispersed into Gondwana, and a derived subgroup of large Late Devonian genera dispersed from Gondwana, is inconsistent with the evidence of the new taxon. Using oldest fossil and most primitive clade criteria the most recent phylogeny resolves South China and Gondwana as areas of origin for all tetrapodomorphs. The immediate outgroup to tristichopterids remains unresolved – either Spodichthys from Greenland as recently proposed, or Marsdenichthys from Gondwana, earlier suggested to be the sister group to all tristichopterids. Both taxa combine two characters that do not co-occur in other tetrapodomorphs (extratemporal bone in the skull; non-cosmoid round scales with an internal boss). Recently both ‘primitive’ and ‘derived’ tristichopterids have been discovered in the late Middle Devonian of both hemispheres, implying extensive ghost lineages within the group. Resolving their phylogeny and biogeography will depend on a comprehensive new phylogenetic analysis.     

Callistophytalean pteridosperms from Permian aged floras of China

Abstract: Recent investigations into Permian aged floras from China have highlighted the widespread occurrence of callistophytalean pteridosperms that challenge previous understanding of their spatial and temporal distribution and diversity. In China, the group spans the Permian period and constitutes a distinctive but rare component in many peat‐forming environments. The stratigraphically earliest callistophytalean occurs in the Asselian‐Sakmarian stages with fossils from the Taiyuan Formation of northern China including ovules of Callospermarion undulatum in coal ball assemblages, and ovulate fronds of Norinosperma shanxiensis and synangiate fronds of Norinotheca shanxiensis in adpression assemblages. More abundant in the fossil records are adpression remains from the Roadian‐Wordian stages with the Lower Shihhotse Formation preserving abundant vegetative and ovulate remains of Emplectopteris triangularis that is now considered to represent a callistophytalean. The youngest callistophytalean recognised is from the Wuchaipingian‐Changhsingian stages with the Xuanwei Formation of southern China containing a single stem of Callistophyton boyssetii that provides indisputable evidence of the group in the lead up to the end‐Permian mass extinction. These accounts are augmented by analysis of pollen records that demonstrate the callistophytalean pollen genus Vesicaspora to be widespread through palynological assemblages from the Permian period in both North and South China, including the Upper Shihhotse Formation, Shihchienfeng Group, Xuanwei Formation, and possibly also in the mid‐Pennsylvanian Benxi Formation. Although macrofossil specimens are uncommon elements in the assemblages that contain them, they demonstrate the continuity of callistophytalean pteridosperms from the Pennsylvanian sub‐period into the early Guadalupian epoch of the Permian in North China and into the Lopingian epoch of the Permian in South China. Of the species present, both Callistophyton boyssetii and Callospermarion undulatum are known from the Pennsylvanian–earliest Permian age floras of Euramerica, whereas Norinosperma, Norinotheca and Emplectopteris appear to represent endemic Cathaysian elements. Results imply that callistophytalean pteridosperms can no longer be excluded from theories of post‐Carboniferous plant evolution and floristics, appearing to have played an important role in both Permian and Carboniferous aged plant communities. The presence of Vesicaspora in several formations from which macro‐remains have not been identified is a hopeful indicator that further callistophytalean pteridosperms are yet to be found.     

Thelodont Scales from the Lower Devonian Red Bay Group, Spitsbergen

Scales of nine thelodont taxa are described from the Red Bay Group of Spitsbergen, also characterized by its assemblage of other typical Early Devonian vertebrates. The four thelodont–bearing horizons of the lower Fraenkelryggen Formation yield typical Lochkovian thelodont assemblages, containing Boreania minima , Canonia grossi , Nikolivia depressa sp. nov., Nikolivia elongata , Nikolivia gutta and Turinia pagei . One horizon from the younger Ben Nevis Formation has a similar thelodont assemblage, except for two new taxa; Apalolepis angelica sp. nov. and Turinia barentsia sp. nov. The appearance of Turinia polita and the disappearance of Boreania minima is also characteristic for this assemblage. Comparison with similar faunas from the Northern Hemisphere supports this faunal change and allows further elaboration of Early Devonian thelodont biostratigraphy.     

A new ammonoid fauna from the Carnian (Upper Triassic) Kasimlar Formation of the Taurus Mountains (Anatolia,Turkey)

We describe a new ammonoid fauna from the Taurus Mountains of southern Turkey. The Carnian ammonoid fauna from A?a?iyaylabel is presented for the first time. Ammonoids were obtained from limestone to marl beds of an approximately 35‐m‐thick section, which presents the rare opportunity to investigate ammonoid faunas across the Lower–Upper Carnian boundary. Intense sampling near the village of A?a?iyaylabel led to the recognition of a new Lower Carnian (Julian 2) to Upper Carnian (Tuvalian 1) ammonoid fauna from the Kasimlar Formation. The genus Kasimlarceltites gen. nov. is reported for the first time from the Taurus Mountains, which represents the main faunal element and occurs as huge mass occurrence (n ? 1 million). Kasimlarceltites krystyni gen. et sp. nov., Klipsteinia disciformis sp. nov. and Anasirenites crassicrenulatus sp. nov. occur within the Lower Carnian Carbonate member (Units A–B) of the Kasimlar Formation from the Taurus Platform Units. Ammonoids described from the marls of the Tuvalian Marlstone member were deposited during a major, Tethyan‐wide climate crisis – the so‐called Carnian crisis – characterized by a demise of carbonate platforms. Based on the biostratigraphic relevance of certain ammonoid taxa described herein, the age of the analysed parts of the Kasimlar Formation is Julian 2 to Tuvalian 1. The discovery of the new ammonoid assemblages from A?a?iyaylabel substantiates the significance of Upper Triassic faunas within the Taurus Mountains and facilitates the correlation with faunal assemblages from other regions in the Tethyan Realm. The ammonoid fauna and facies indicate a general deepening from open‐platform margins, over deeper shelf settings down to an open marine‐influenced basinal environment. The tentative habitat for Kasimlarceltites gen. nov. is a shallow platform environment to upper mid‐ramp.     

黔南独山地区泥盆系法门阶–石炭系杜内阶革老河组 和汤粑沟组长身贝类新认知

本文描述了在黔南独山县其林寨剖面和黎家寨剖面上泥盆统革老河组和下石炭统汤粑沟组中发现的长身贝类化石共8属10种(含未定种)。其中, 在革老河组中发现有3属4种: Productella? sp.、Yanguania dushanensis、Xinshaoproductus xinshaoensis和X. quadrata; 汤粑沟组中有8属9种: Productella? sp.、Hunanoproductus hunanensis、Spinocarinifera qilinzhaiensis sp. nov.、Yanguania dushanensis、Y.? sp.、Xinshaoproductus xinshaoensis、Tomiproductus sp.、Ozora? sp.和Ericiatia kiangsuensis。总体而言, 革老河组和汤粑沟组中的长身贝类在组成上主要以Productoidea超科中Productidae科的分子为主, Productellidae科和Echinoconchoidea超科中Sentosiidae科的分子各有1种。本文记述的Hunanoproductus和Xinshaoproductus系在黔南地区首次发现, 并且Xinshaoproductus在黔南地区的首现层位低于湘中地区。常见于欧亚地区的中泥盆统吉维特阶至上泥盆统弗拉斯阶及湘中地区上泥盆统的Productella属, 在黔南地区上延到了下石炭统的汤耙沟组中, 为该属最晚的化石记录。Spinocarinifera和Tomiproductus是欧亚、北美、西澳和北非等地杜内阶或杜内阶-维宪阶中的常见分子, 汤粑沟组中发现的S. qilinzhaiensis sp. nov.则是目前已知该属在华南地区的代表。该种以近方圆形的轮廓、铰合线短、耳翼小、放射状壳线连续但微弱为特征, 区别于该属的其他种。而Tomiproductus在华南地区见于湘中刘家塘组的上部和下扬子地区的金陵组。Ozora属于美国密苏里地区维宪阶下部的代表分子, 亦见于墨西哥南部的下石炭统。如果鉴定无误, 黔南地区的发现则有可能代表了该属最早的化石记录。革老河组的长身贝类组成属种单调, 以Yanguania、Xinshaoproductus属等地方性分子为特征; 随着杜内期的海侵, 汤耙沟组中的长身贝类组成与湘中地区的更为相近, 生物多样性显著增加, 既继承了革老河组中的地方性分子, 同时在属级分类单元中也出现了一些在欧美、澳大利亚等地区的类型, 如Spinocarinifera、Tomiproductus和Ozora属。反映了从杜内期开始的海侵, 使华南地区的腕足动物群与欧美地区同时期的腕足动物群之间开始发生较为密切的生物地理区系联系。     

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.     

滇东泥盆系几个含鱼层的时代和对比

从鱼化石的角度对滇东泥盆系下述含鱼地层进行了分析,并认为:1)曲靖地区下泥盆统徐家冲组与坡松冲组或四川的平驿铺组、广西的那高岭组层位相当,时代为布拉格期;其层位低于含 duyunolepids-Kueichowlepis鱼类组合的贵州乌当组或舒家坪组,后者时代为埃姆斯期。2)滇东中泥盆统含两个不同时期的含鱼层,其中含Yinosteus-Wudinolepis鱼类组合的武定地区旧城组属中泥盆统早期,该鱼类组合现仅知分布于武定地区。而中泥盆统晚期含鱼地层,在曲靖地区由下而上包括穿洞组、上双河组和海口组三个岩石组;其中穿洞组＋上双河组与武定的鱼子甸组对比,而海口组与曲靖组为同时异相;武定茶花箐组与曲靖组对比。3)文中时上述地层中的某些鱼化石的层位或鉴定作了评述或厘定。     

Some observations on Zosterophyllutn australianum Lang & Gookson from the Lower Devonian of Yunnan, China

SHOU-GANG, H., 1992. Some observations on Zosterophyllutn australianum Lang & Cookson from the Lower Devonian of Yunnan, China. Recent collections of Zosterophyllum australianum have been made from the Xujiachong Formation which outcrops near Xujiachong village in the Qujing district of Yunnan province. The unit is a non-marine facies of the type section of the Lower Devonian of China. Sediments containing the fossil material are probably Siegenian-Emsian in age. Details of the morphology of fertile spike of Z. australianum are recorded. The spikes bear short-stalked, rather large, transversely elliptical sporangia, each with a wide, thickened, distal border. Sporangia are borne close together in a dextrorse helix, and usually in four or five longitudinal rows. Morphological variation of in-situ spores is also described. Comparison of the new specimens with others of reported Z. australianum has led to some nomenclatural changes.     

High‐latitude Hirnantian (latest Ordovician) brachiopods from the Eusebio Ayala Formation of Paraguay,Paraná Basin

Abstract: Examination of newly collected brachiopods from the Eusebio Ayala Formation of Paraguay reveals the occurrence of Arenorthis paranaensis sp. nov., Plectothyrella? itacurubiensis sp. nov., Hindella sp. and Eostropheodonta conradii (Harrington). Associated graptolites of the N. persculptus Zone indicate that the age of the fossiliferous beds is Hirnantian. The overall generic composition of the fauna is similar to that of the atypical Hirnantia Fauna of the Bani Province. The record in Paraguay of Arenorthis, hitherto only known from North Africa, together with species of Plectothyrella? and Eostropheodonta different from those recorded in the Kosov Province, emphasizing the affinities between the Paraguayan fauna and the low‐diversity African assemblages. Stratigraphic and faunal evidence indicates that biogeographical links between South America and Africa already existed by the end of the Ordovician when most of the intra‐cratonic basins of Gondwana were flooded during the postglacial sea level rise.     

Stenolaemate bryozoans from the Geirud Formation (Upper Devonian/Lower Carboniferous) of Central Alborz (Iran)

Two bryozoan species are described from the Geirud Formation (Upper Devonian/Lower Carboniferous) of Central Alborz (Iran). Trepostome Schulgina mutabilis Troizkaya, 1975 is known from the Upper Devonian (Famennian) of Central Kazakhstan. The new species Ascopora geirudensis n. sp. is the earliest known representative of the rhabdomesine genus Ascopora Trautschold, 1876.     

New fossil insects (Diaphanopterodea: Martynoviidae) from the Lower Permian of the Boskovice Basin, southern Moravia

Two new genera and new species assigned to the family Martynoviidae (Diaphanopterodea), Boskovicia celtai nov. gen., nov. sp. and Moravoptera hladilovae nov. gen., nov. sp. are described and illustrated from the Lower Permian (Artinskian/Sakmarian) locality of Obora, southern Moravia (Czech Republic). Discovery of the species Martynovia insignis Tillyard, originally described from the Lower Permian of Kansas (USA), supports close relationship of both faunas across Euramerica.     

Mesozoic Trachypachidae (Insecta: Coleoptera) from China

Abstract: Four well‐preserved beetles, attributed to Eodromeinae within Trachypachidae, are described from the Mesozoic of Inner Mongolia, China. Eodromeus robustus sp. nov., E. daohugouensis sp. nov. and Unda chifengensis sp. nov. are from the Middle Jurassic of Daohugou. The diagnostic characters for the two genera are revised, and all species of these genera are keyed. A new genus and species, Sinodromeus liutiaogouensis gen. nov., sp. nov., is described from the Lower Cretaceous Yixian Formation of Liutiaogou. The current fossil records of Trachypachidae from China are reviewed, and only five species (four described here) can be convincingly attributed to this family. The morphological disparity of Mesozoic Eodromeinae suggests that they evolved a broad spectrum of locomotory lifestyles.     

Snipe flies (Diptera: Rhagionidae) from the Daohugou Formation (Jurassic), Inner Mongolia,and the systematic position of related records in China

Abstract: Four new species referred to three new genera and one known genus of fossil snipe flies (Family Rhagionidae) from the Daohugou Formation of Chifeng City in Inner Mongolia, north‐eastern China, are described as Daohugorhagio elongatus gen. et sp. nov., Parachrysopilus jurassicus gen. et sp. nov., Sinorhagio sinuatus sp. nov. and Trichorhagio gregarius gen. et sp. nov. These taxa represent only a fraction of the rhagionid assemblage in the Daohugou biota. They contribute towards the high diversity and abundance of snipe flies during the latest Middle Jurassic – earliest Late Jurassic. The diagnosis of Sinorhagio K. Zhang, Yang and Ren, 2006 is revised based on the new results. Many genera and species from the Lower Cretaceous of China have been assigned to Rhagionidae. Most of these do not actually belong to this family: specifically Mesorhagiophryne incerta Hong and Wang, 1990, M. robusta Hong and Wang, 1990, Mesostratiomyia laiyangensis Hong and Wang, 1990, Stratiomyopsis robusta Hong and Wang, 1990, Longhuaia orientalis Hong, Wang and Sun, 1992, Basilorhagio venusius Ren, 1995. Pauromyia oresbia Ren, 1998 is transferred to an archisargid genus as Sharasargus oresbius (Ren, 1998) comb. nov.     

Late Emsian Rutoceratoidea (Nautiloidea) from the Prague Basin,Czech Republic: morphology,diversity and palaeoecology

Abstract: Nautiloids of the superfamily Rutoceratoidea from the late Emsian (late Early Devonian) of the Prague Basin (Czech Republic) are commented upon. Species recognized include the hercoceratids Hercoceras mirum, H.? transiens, Ptenoceras proximum, P. nudum, P. minusculum and Anomaloceras anomalum, as well as the rutoceratids Adelphoceras bohemicum, Homoadelphoceras devonicans, Pseudorutoceras bolli and Goldringia? devonicans. In addition, four new species are described: Parauloceras regulare sp. nov., Roussanoffoceras chlupaci sp. nov., Otomaroceras sp. nov. and Goldringia sp. nov. Morphology and distribution patterns of Pragian and late Emsian rutoceratoid faunas from the Prague Basin are compared. They show that an increased diversity was accompanied by a higher level of specialization of rutoceratoids, which manifested itself in low abundance, increased facies dependence and greater variation in shell size during the Early Devonian. The evolution of sculpture and a contracted aperture, both regarded as protective adaptive features, was also examined, but no adaptive trend towards more pronounced sculpture and constriction of the aperture was found to have occurred in the Early Devonian. A more distinctive sculpture was, however, observed in shallow‐water assemblages of P. proximum in comparison with deeper‐water faunules, and two additional cephalopod species were examined in order to obtain comparative data. The presence of distinct sculpture patterns in coeval shallow‐ and deeper‐water assemblages suggests limited migration between them and consequently reflects some degree of territoriality in Devonian nautiloids. New data on early shell development in P. proximum are presented. During the Chote? Event, rutoceratoid generic diversity dropped dramatically, one family became extinct and the Early Devonian diversification of the group came to an end. The recovery of nautiloid faunas was slower than that of other cephalopods and associated, unrelated invertebrates. The absence of change in abundance patterns between Pragian and late Emsian rutoceratoid faunas, i.e. prior to and subsequent to ammonoid radiation, suggests that the appearance and radiation of the latter group in the early Emsian did not affect the structure of nautiloid assemblages, i.e. these two clades did not occupy the same niches.     

Further evidence for the presence of holoptychiid porolepiforms (Sarcopterygii,Dipnomorpha) from the Frasnian of Colombia

The Devonian (Frasnian) of Colombia has provided important information on the taxonomical diversity and palaeobiogeographic relationships of the Late Devonian marginal marine vertebrate faunas of South America (northwestern margin of Gondwana). This fauna is mostly composed of Gondwanan endemics, but includes two taxa also known in Euramerica: the antiarchan placoderm Asterolepis and the porolepiform sarcopterygian Holoptychius. The occurrence of holoptychiid porolepiforms in Colombia was previously suggested based solely on the presence of scales attributed to Holoptychius, although with caution. Here, we describe further holoptychiid remains that include a large isolated tooth with dendrodont microstructure, typical of porolepiforms, and additional Holoptychius scales. These new findings increase the record of holoptychiid porolepiform occurrences in Gondwana and suggest that dispersion of fish faunas between Euramerica and Gondwana by the beginning of the Late Devonian was possible through South America.