Proharpoceras Chao: a new ammonoid lineage surviving the end-Permian mass extinction

Based on new, bed-rock controlled material from northwestern Guangxi and Oman, the Early Triassic genus Proharpoceras Chao is shown to be a representative of Otocerataceae. Character analysis excludes a direct link with the Griesbachian Otoceratidae and favours a derivation of Proharpoceras from the late Permian Anderssonoceratidae. The biostratigraphic range of Proharpoceras is restricted to the Smithian and its biogeographic distribution comprises Oman, South China, and Primorye, thus indicating an essentially low palaeolatitudinal distribution. Proharpoceras has no apparent relatives among other Early and Middle Triassic Ceratitida and is thus considered to be the last representative of Otocerataceae. This offshoot of the late Permian Anderssonoceratidae implies that an additional ammonoid lineage survived the end Permian extinction and that it dwindled away for some 2 Myr before going extinct.     

Palaeoecological significance of a new Griesbachian (Early Triassic) gastropod assemblage from Oman

A new Early Triassic (Griesbachian) gastropod fauna from the Al Jil Formation of Oman is described. Early Triassic faunas from elsewhere (e.g. the Italian Dolomites and the western USA) are typically of low diversity and high dominance, usually attributed to environmental stress in the immediate aftermath of the end-Permian mass extinction event. The new Oman fauna has, by contrast, relatively high diversity, low dominance and a more even spread of individuals between taxa. It is the most diverse Griesbachian fauna known to date. This is attributed to the favourable (i.e. well-oxygenated) conditions under which the fauna lived. This uncharacteristic Griesbachian gastropod fauna demonstrates that, in the absence of oceanic anoxia, biotic recovery after the end-Permian extinction event may occur surprisingly quickly (within one conodont zone). The fauna is also partially silicified, which has increased its preservation potential relative to other Griesbachian gastropod assemblages. However, only one reappearing Lazarus genus is present in the Oman fauna. This suggests that there was some other control on the abundance of Lazarus genera at this time, other than the absence of silicified faunas as previously suggested.     

Diversity variation and tempo-spatial distribution of Otozamites (Bennettitales) in the Mesozoic of China

Otozamites is a representative fossil leaf morphogenus of the extinct Bennettitales, with an extensive distribution during the Mesozoic, especially in China. Understanding the fossil diversity variation and distribution pattern of Otozamites in China will provide information on biodiversity of bennettitalean plants as well as for reconstruction of palaeogeography and palaeoclimate conditions during the Mesozoic. So far, 46 species of this genus have been described in China, excluding unspecified species. The results show that the fossils of Otozamites are extensively recorded in the Late Triassic, and then reach their maximum development in the Early Jurassic, followed by a reduction in diversity in the Middle and Late Jurassic, and finally become extinct at the end of Early Cretaceous. Geographically, they occur in both Northern and Southern Floristic Provinces in the Mesozoic of China, with a relatively higher abundance in the Southern Floristic Province. It implies that the diversity variation and distribution of Otozamites are closely related to the change of the palaeoclimatic conditions. The warm and humid climate prevailed in the Late Triassic and Early Jurassic in South China, propitious to the development of Otozamites. After the Middle Jurassic, dry and hot climate may have caused the lower diversity level and blocked the development of Otozamites; finally at the end of the Early Cretaceous, the frequent arid climate may be a major cause for the extinction of Otozamites.     

An unusually diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic recovery after the end-Permian biotic crisis

A single carbonate coquinoid lens from the Griesbachian (Early Triassic) of Shanggan, South China, yielded 11 bivalve species described in this study in addition to four gastropod and one ammonoid species reported elsewhere. This makes the Shanggan fauna one of the richest mollusc faunas from the early post-extinction interval after the end-Permian mass extinction event. Four of the present genera are long-term survivors, five are holdovers that went extinct at the end of the Griesbachian or later in the Early Triassic, and seven first appear in the Griesbachian. Three new bivalve species are described: Myalinella newelli nov. sp., Scythentolium scutigerulus nov. sp., and Eumorphotis shajingengi nov. sp. The genus Astartella, previously assumed to have vanished at the end of the Permian, is reported for the first time from the Early Triassic, which also removes Astartidae from Early Triassic Lazarus taxa. The small growth size of the Astartella specimens supports an earlier hypothesis that many of the Early Triassic Lazarus taxa did not survive in unknown refuges but were simply overlooked due to the scarcity of easily observable large-sized specimens. Ecologically, a comparatively high proportion of infaunal bivalve species (4/11) is remarkable for the early post-extinction interval, supporting the impression of a relatively advanced recovery state. Moreover, abundance-data of the bivalve-gastropod community reveal a remarkably low dominance index (D = 0.17) that is suggestive for advanced recovery and stable environmental conditions. It is proposed that the Shanggan fauna represents a late Griesbachian benthic recovery event that coincided with the appearance of similarly diverse benthic faunas in Oman and Primorye. A high proportion of genera that have previously not been reported from the Early Triassic indicate that the prevalence of poor preservation conditions is a major obstacle in identifying early phases of recovery from the greatest crisis in the history of metazoan life. The early recovery of benthic faunas reported in this study questions previous claims of a prolonged lag phase as a consequence of the extraordinary extinction magnitude or the persistence of adverse environmental conditions.     

Triassic temnospondyl biostratigraphy, biochronology and correlation of the German Buntsandstein and North American Moenkopi Formation

Based on temnospondyl distributions, we propose a correlation of the non-marine Lower and Middle Triassic strata of Germany and the western United States. This correlation relies principally on the presence of two genera, Nonesian (late Early Triassic) Parotosuchus and Perovkan (early Middle Triassic) Eocyclotosaurus . The distribution of Parotosuchus helgolandicus indicates that the Torrey Member of the Moenkopi Formation in Utah, USA is equivalent to the Volpriehausen Formation of the German Buntsandstein of Nonesian age. Magnetostratigraphy and the distribution of Eocyclotosaurus indicate that the Perovkan Holbrook Member of the Moenkopi Formation in Arizona corresponds to a short interval of Anisian time, as does the German Upper Buntsandstein. The Buntsandstein and Moenkopi biostratigraphic schemes agree with a recently proposed biochronology of Triassic terrestrial tetrapods.     

A RHEXOXYLON-LIKE STEM FROM THE MORRISON FORMATION OF UTAH

Arnold , Chester A. (U. Michigan, Ann Arbor.) A Rhexoxylon-like stem from the Morrison formation of Utah. Amer. Jour. Bot. 49(8): 883–886. Illus. 1962.—A silicified stem 11 ft long from the Upper Jurassic of the Henry Mountains region of southern Utah resembles the South African and South American Upper Triassic genus Rhexoxylon, except for the absence of anomalously developed vascular tissue in the pith. It is described as Hermanophyton kirkbyorum gen. et sp. nov.     

A new species of basal rhynchosaur (Diapsida: Archosauromorpha) from the early Middle Triassic of South Africa,and the early evolution of Rhynchosauria

Rhynchosauria was an important clade of herbivorous archosauromorph reptiles during the Triassic, with a worldwide distribution. We describe a new genus and species of early rhynchosaur, E ohyosaurus wolvaardti gen. et sp. nov. , from the early Middle Triassic (early Anisian) Cynognathus Assemblage Zone (Subzone B) of the Karoo Supergroup, South Africa. Eohyosaurus wolvaardti is known from a single skull, and is recovered as the sister taxon of Rhynchosauridae in a new phylogenetic analysis. Cynognathus Subzone B has previously yielded the stratigraphically oldest well‐understood rhynchosaur species, Mesosuchus browni and Howesia browni. Eohyosaurus wolvaardti increases the rhynchosaur diversity within this stratigraphical horizon to three species. Intriguingly, all currently confirmed rhynchosaur occurrences from the Early Triassic to earliest Middle Triassic are from South Africa. This may suggest a relatively restricted palaeogeographical distribution for early rhynchosaurs, followed by a global dispersal of rhynchosaurids during the Middle Triassic. © 2015 The Linnean Society of London     

Diversity variation and tempo-spatial distributions of the Dipteridaceae ferns in the Mesozoic of China

The extant family Dipteridaceae is a remarkable leptosporangiate fern because it includes only one genus with a restricted distribution to tropical regions. The fossil record of this family has been widely reported from the Mesozoic strata in Eurasia, America, Australia, and Greenland. In China, numerous fossils of the Dipteridaceae have been documented, in total, about 74 species of 6 genera. Geographically, they are distributed both in the Southern and Northern Floristic Provinces, and were particularly well developed in the Southern Floristic Province during the Late Triassic and the Early Jurassic intervals. Fossil diversity of Dipteridaceae varies in the different episodes of the Mesozoic in China. It is shown that Dipteridaceae has undergone a diversity development process and a distinct turnover during the Mesozoic. They appear to have diversified in the warm and humid Late Triassic–Early Jurassic, but declined sharply as aridity developed in the Middle Jurassic, and became extinct at the end of the Early Cretaceous. The diversity variation and tempo-spatial distribution pattern is suggested to be linked with paleoclimatic variations during the Mesozoic.     

A New Genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for Timing the Recovery of Brachiopoda After the End-Permian Mass Extinction

A new genus, Meishanorhynchia , is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic–Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1 ± 0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3 ± 0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4 ± 0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.     

Plants of Leptostrobus Heer （Czekanowskiales） from the Early Cretaceous and Late Triassic of China, with Discussion of the Genus

Investigation of the Mesozoic seed plant Leptostrobus Heer from the Yangcaogou Formation of the Late Triassic and the Yixian Formation of the Early Cretaceous, Liaoning Province, China, provides new Insight Into Its general morphology and geographical distribution. The materials of L. cancer from the Yixian Formation described herein are later than all the past findings of this species and add to the record of L. cancer during the Early Cretaceous. Based on well-preserved specimens, the specific diagnosis Is slightly emended and the reconstructlon of L. cancer Is perfected. The materials from the Yangcaogou Formation of the Late Triassic are placed in L. spheericus, in addition, we review the history of investigation of the genus Leptostrobus since its establishment in 1876 and discuss the main characteristics of each species.     

Earliest Triassic microbialites in the South China block and other areas: controls on their growth and distribution

Earliest Triassic microbialites (ETMs) and inorganic carbonate crystal fans formed after the end-Permian mass extinction (ca. 251.4 Ma) within the basal Triassic Hindeodus parvus conodont zone. ETMs are distinguished from rarer, and more regional, subsequent Triassic microbialites. Large differences in ETMs between northern and southern areas of the South China block suggest geographic provinces, and ETMs are most abundant throughout the equatorial Tethys Ocean with further geographic variation. ETMs occur in shallow-marine shelves in a superanoxic stratified ocean and form the only widespread Phanerozoic microbialites with structures similar to those of the Cambro-Ordovician, and briefly after the latest Ordovician, Late Silurian and Late Devonian extinctions. ETMs disappeared long before the mid-Triassic biotic recovery, but it is not clear why, if they are interpreted as disaster taxa. In general, ETM occurrence suggests that microbially mediated calcification occurred where upwelled carbonate-rich anoxic waters mixed with warm aerated surface waters, forming regional dysoxia, so that extreme carbonate supersaturation and dysoxic conditions were both required for their growth. Long-term oceanic and atmospheric changes may have contributed to a trigger for ETM formation. In equatorial western Pangea, the earliest microbialites are late Early Triassic, but it is possible that ETMs could exist in western Pangea, if well-preserved earliest Triassic facies are discovered in future work.     

黔西滇东早三叠世早期Annalepis(脊囊属)的出现及其地层意义

为解决早三叠世卡以头组地质时代和植物组合问题,对采于黔西滇东地区两条二叠系-三叠系海陆交互相界线剖面卡以头组中、下部的植物化石Annalepis(脊囊属)进行深入研究。与Annalepis同层或上部层位见有早三叠世早期的海相双壳类、介形类、腕足类、菊石类及大羽羊齿植物群残余分子,因此可以确定Annalepis在该地区出现于早三叠世早期的印度期(Induan),说明卡以头组(至少中、下部)地质时代属早三叠世早期。推测我国南方的Annalepis在早三叠世始于黔西滇东,随后向北迁移、中三叠世广布于长江中下游地区。建立由早三叠世An-nalepis和大羽羊齿植物群残存分子共同构成的我国南方早三叠世早期植物组合,填补了早三叠世晚期岭文组与晚二叠世B期Gigantonoclea guizhouensis-Ullmannia cf.bronnia-Annularia pinglouensis组合之间的空白。     

Jurassic evolution of Southern Hemisphere marine palaeobiogeographic units based on benthonic bivalves

The distribution of benthonic Jurassic bivalve genera in the Southern Hemisphere is analysed here. For this region, palaeobiogeographic units (biochoremas) are quantitatively characterized according to their biologic contents (mainly levels of endemism). Their evolution through time is followed from the latest Triassic to the earliest Cretaceous. The Tethyan Realm is undoubtedly the most mature and persistent through time, with three subordinate units: an Australian unit restricted to the Late Triassic, a North Andean unit, which appears sporadically as an endemic centre, and an East African unit which is recognisable from Bajocian times onwards. From Late Triassic times, a South Pacific Realm has been recognised, with a Maorian Province mostly based on the distribution of monotoid genera. A South Andean unit is also recognisable through most of the Jurassic, and its reference either to the South Pacific unit or to the Tethyan Realm is a matter of debate. Being a transitional biogeographic setting between Tethyan and South Pacific first-order units, it is included in the South Pacific unit due to the common presence of antitropical (didemic) genera. The East African unit is included within the Tethyan Realm during the Jurassic, but during Early Cretaceous times, it splits into two units, one of which was regarded as part of the “South Temperate Realm” by Kauffman. The rank of all these units changed with time. Throughout the Jurassic, the ecotone between South Pacific and Tethyan palaeobiogeographic units fluctuated in position with time. The approximate latitudinal location of the ecotonal boundary area and its shift through time are recognised on the basis of faunal composition along the Andean region.     

Lystrosaurus species composition across the Permo–Triassic boundary in the Karoo Basin of South Africa

Lystrosaurus is one of the few therapsid genera that survived the end-Permian mass extinction, and the only genus to have done so in abundance. This study identifies which species of Lystrosaurus have been recovered from Permian and Triassic strata to determine changes in the species composition across the Permo–Triassic (P–T) boundary in the Karoo Basin of South Africa. Data generated from museum collections and recent fieldwork were used to stratigraphically arrange a total of 189 Lystrosaurus specimens to determine which species survived the extinction event. Results reveal that L. curvatus and L. maccaigi lived together on the Karoo floodplains immediately before the extinction event. L. maccaigi did not survive into the Triassic in South Africa. L. curvatus survived, but did not flourish and soon became extinct. Two new species of Lystrosaurus , L. murrayi and L. declivis , appeared in the Early Triassic. It is possible that L. murrayi and L. declivis occupied different niches to L. maccaigi and L. curvatus , and had special adaptations that were advantageous in an Early Triassic environment. We suggest that L. maccaigi may be used as a biostratigraphic marker to indicate latest Permian strata in South Africa and that, in support of previous proposals, the genus Lystrosaurus should not be used as a sole indicator of Triassic-aged strata. Our field data also show that L. curvatus may be regarded as a biostratigraphic indicator of the P–T boundary interval.     

New data from Oman indicate benthic high biomass productivity coupled with low taxonomic diversity in the aftermath of the Permian–Triassic Boundary mass extinction

A new Early Triassic marine fauna is described from an exotic block (olistolith) from the Ad Daffah conglomerate in eastern Oman (Batain), which provides new insights into the ecology and diversity during the early aftermath of the Permian–Triassic Boundary mass extinction. Based on conodont quantitative biochronology, we assign a middle Griesbachian age to the upper part of this boulder. It was derived from an offshore seamount and yielded both nektonic and benthic faunas, including conodonts, ammonoids, gastropods and crinoid ossicles in mass abundance. This demonstrates that despite the stratigraphically near extinction at the Permian–Triassic Boundary, Crinoidea produced enough biomass to form crinoidal limestone as early as middle Griesbachian time. Baudicrinus, previously placed in Dadocrinidae, is now placed in Holocrinidae; therefore, Dadocrinidae are absent in the Early Triassic, and Holocrinidae remains the most basal crown‐group articulates, originating during the middle Griesbachian in the Tethyan Realm. Abundant gastropods assigned to Naticopsis reached a shell size larger than 20 mm and provide another example against any generalized Lilliput effect during the Griesbachian. Whereas the benthic biomass was as high as to allow the resumption of small carbonate factories, the taxonomic diversity of the benthos remained low compared to post‐Early Triassic times. This slow benthic taxonomic recovery is here attributed to low competition within impoverished post‐extinction faunas.     

Some Fossil Plants of Thinnfeldia from the Jiuligang Formation in Jingmen-Dangyang Basin,Western Hubei

The genus Thinnfeldia was seldom reported in former geological literature from Hubei. In the present paper, the following fossil plants of Thinnfeldia from Hubei are collected for the first time: Thinnfeldia elegans (sp. nov.), T. nanzhangensis (sp. nov.), T. yuananensis (sp. nov.), from Late Triassic Jiuligang Formation of the Jingmen-Dangyang Basin. It is interesting to point that the Jiuligang flora has transational characters between Yanchang flora of North China and Anyun flora of South China of Late Triassic.     

贵州关岭上三叠统的楯齿龙类化石

记述的新铺中国豆齿龙（Sinocyamodus xinpuensis gen. et sp. nov.）代表了齿龙目的一新属。这是齿龙类化石在我国的首次发现,也是这类特殊的海生爬行动物在欧洲、北非和中东以外地区的首次记录。标本产自贵州关岭晚三叠世瓦窑组一段的泥晶灰岩中,与鱼龙类、幻龙类及海龙类等多种海生爬行类同时产出。这一化石组合表明,贵州关岭三叠纪海生爬行动物群的性质与西特提斯动物群（wstern-Tethyan fauna）的性质极为相似,这两个地区在晚三叠世早期有广泛的动物交流。     

Battenizyga, a new Early Triassic gastropod genus with a discussion of the caenogastropod evolution at the Permian/Triassic boundary

Battenizyga, a new Early Triassic gastropod genus from the Moenkopi Formation of Utah, is described and the speciesAnoptychia eotriassica Batten & Stokes, 1986 is placed in it. The new genus has an axially ribbed planktonic larval shell and a teleoconch with an angulated periphery. This character combination is unknown from the Palaeozoic. Therefore,Battenizyga represents additional evidence that recovery from the end-Permian mass extinction was connected with a faunal turnover. Additionally, the extinction of diverse Palaeozoic groups of the Caenogastropoda in the Permian (e.g., the Pseudozygopleuridae) suggest a turnover. All caenogastropod genera that hold Early Triassic species, have post-Palaeozoic type species and most were not reported from the Palaeozoic. This corroborates the view that there was an intense faunal turnover within the Caenogastropoda.Battenizyga is probably a caenogastropod that is closely related to the superfamily Zygopleuroidea which is abundant in the late Palaeozoic and early Mesozoic.      

New Middle Jurassic record of fern genus Eboracia (Dicksoniaceae,Filicales) and its spatio-temporal distribution in China

As an important component of the Mesozoic flora, the extinct fern genus Eboracia Thomas (Dicksoniaceae, Filicales) is widely reported in China with diverse fossil records. New material of Eboracia lobifolia, represented by a nearly intactly preserved fossil frond, is described herein from the Middle Jurassic Haifanggou Formation in Beipiao of western Liaoning, Northeast China. The frond is lanceolate in gross outline, at least 38.0 cm long, and can be divided into the basal sterile part, the upper fertile part and a transitional part in between. The new discovery confirms for the first time that the frond of E. lobifolia is hemidimorphic rather than holodimorphic. Many in-situ spores were detached from the sori of the fertile pinnae, which are characterized by rounded-tetrahedral shape, smooth surface, distinct border, and a wide, long triradiate crack almost reaching the equator. Comparing with dispersed spores in the same horizon, these in-situ spores seem to be most similar to Cyathidites minor Couper in morphology. A spatio-temporal analysis of Eboracia in China shows that Eboracia with totally four species occurred in a time interval ranging from the Late Triassic to the Early Cretaceous in China, and mostly flourished in the Middle Jurassic; the genus was widely distributed in both the Northern and Southern Phytofloristic Provinces of China, particularly in southern China during the Late Triassic to Early Jurassic, while more abundant and diverse in northern China during the Middle Jurassic to Early Cretaceous.     

New early Jurassic decapod crustacean from Patagonia (Chubut province), Argentina

A well-preserved decapod specimen was found in early Toarcian deposits cropping out on the western slope of Meseta Catreleo, central Chubut province, Argentina. It is a nearly complete exoskeleton preserved in lateral view, slightly crushed, in fine-grained sandstones. The skeleton is mostly articulated, though some pieces are disarticulated or missing. Taphonomic features indicate a relatively rapid burial after death, with little or no transport. This specimen is here described as a new species of the genus Mecochirus Germar (Decapoda: Glypheoidea). Mecochirus robbianoi n. sp. is characterized by a very long, achelate, first pair of pereiopods with a narrow, long, straight dactylus, a thin carapace ornamented by low tubercules, with a slightly oblique cervical groove and a short acute rostrum. The family Mecochiridae arose in the Triassic, but the group diversified during the Early Jurassic. Mecochirus had a wide geographical distribution and is known for sure from Lower Jurassic to Upper Cretaceous deposits. The new record from Chubut is one of the oldest for the genus, the oldest for South America, and the first for Argentina, thus considerably extending the known distribution of the genus during the Early Jurassic.