A living fossil tale of Pangaean biogeography

The current distributions of widespread groups of terrestrial animals and plants are supposedly the result of a mixture of either vicariance owing to continental split or more recent trans-oceanic dispersal. For organisms exhibiting a vicariant biogeographic pattern—achieving their current distribution by riding on the plates of former supercontinents—this view is largely inspired by the belief that Pangaea lacked geographical or ecological barriers, or that extinctions and dispersal would have erased any biogeographic signal since the early Mesozoic. We here present a time-calibrated molecular phylogeny of Onychophora (velvet worms), an ancient and exclusively terrestrial panarthropod group distributed throughout former Pangaean landmasses. Our data not only demonstrate that trans-oceanic dispersal does not need be invoked to explain contemporary distributions, but also reveal that the early diversification of the group pre-dates the break-up of Pangaea, maintaining regionalization even in landmasses that have remained contiguous throughout the history of the group. These results corroborate a growing body of evidence from palaeontology, palaeogeography and palaeoclimatic modelling depicting ancient biogeographic regionalization over the continuous landmass of Pangaea.     

A new non-mammalian eucynodont from the Chinle Formation (Triassic: Norian), and implications for the early Mesozoic equatorial cynodont record

The Upper Triassic tetrapod fossil record of North America features a pronounced discrepancy between the assemblages of present-day Virginia and North Carolina relative to those of the American Southwest. While both are typified by large-bodied archosaurian reptiles like phytosaurs and aetosaurs, the latter notably lacks substantial representation of mammal relatives, including cynodonts. Recently collected non-mammalian eucynodontian jaws from the middle Norian Blue Mesa Member of the Chinle Formation in northeastern Arizona shed light on the Triassic cynodont record from western equatorial Pangaea. Importantly, they reveal new biogeographic connections to eastern equatorial Pangaea as well as southern portions of the supercontinent. This discovery indicates that the faunal dissimilarity previously recognized between the western and eastern portions of equatorial Pangaea is overstated and possibly reflects longstanding sampling biases, rather than a true biogeographic pattern.     

Bored Bivalves in Upper Triassic (Norian) Event Beds,Northeastern British Columbia,Canada

Skeletobionts are important components of most shallow marine ecosystems. Prior to the fossils reported herein, evidence of skeletobionts was absent from Upper Triassic successions on the northwestern margin of Pangaea. The boring Talpina ramosa is reported from bivalve body fossils from the Upper Triassic (Lower Norian) lower Pardonet Formation at Pink Mountain in northeastern British Columbia. This Ichnotaxon penetrates through both the outer and inner surface of articulated and disarticulated bivalve shells preserved within sharp-based event beds. The occurrence of these trace fossils underscores the paucity of borings, bioerosional structures, and encrusting taxa from Triassic successions in the western Pangaean realm. Due to erosional removal of shallow water strata by a post-Triassic unconformity, these event beds provide the only available information regarding the ecological health of Late Triassic depositional systems in the study area.     

Wildfires in the Early Triassic of northeastern Pangaea: Evidence from fossil charcoal in the Bogda Mountains,northwestern China

Fires are an integral part of modern and ancient ecosystems, serving as friends for renewal or foes for complete destruction and extinction. Indicators of palaeowildfire were so far absent from the Lower Triassic. Lack of plants in the Early Triassic due to the end-Permian mass extinction event and low atmospheric oxygen levels were proposed as the major reasons for the scarcity of wildfires. We present macroscopic charcoals from the Olenekian (Lower Triassic) in northwestern China, indicating probable ground/smoldering fires occurred on landscapes in mid-latitudes of northeastern Pangaea. Atmospheric oxygen concentration during the Olenekian would have been above 18.5%. These findings demonstrate that wildfires continued to be a source of disturbance of terrestrial ecosystems in Bogda Mountains after the end-Permian marine biotic crisis. There were adequate supplies of fuels and oxygen during this critical time period in the Earth history.     

Ecological succession among late palaeozoic and mesozoic tetrapods

Natural selection and the development of new taxa are associated with ecological replacement and the increase in number of niches with time. Continental faunal interchange was possible globally because of the existence of the super-continent Pangaea during much of the Upper Palaeozoic and Mesozoic. Figures of tetrapod niches vs. time and discussion of this concept for that period are presented for the first time. Four habitat divisions are used, namely marine, fresh-water, lowland and upland.The marine habitat was colonised rather late by tetrapods and these may have been the first predators on the early bony fishes which had diversified in the Permian. The radiation of bony fishes in the Jurassic was followed by a further increase in variety of their reptilian predators. Predators seem to develop some time after the radiation of a new potential prey group.Most early amphibians occupied fresh-water habitats in “crocodile” or “frog” niches, but from the Triassic tetrapods moved from fresh-waters and lowlands into the uplands also.In terrestrial habitats, the replacement of mammal-like reptiles by dinosaurs is tentatively explained in terms of palaeoclimatology and thermoregulatory physiology. Ornithischians capable of dealing with tough vegetation evolved to occupy the new niches produced by the radiation of conifers in the Jurassic. The extinction of dinosaurs appears to have been connected with temperature and habitat changes.Conclusions are supported by a summary of published opinions on the palaeoecological roles of early tetrapods.     

Bioeroders in fossil reefs

Summary Boring algae, fungi and bacteria have been the most constant factor in bioerosion through earth history. Their record reaches back into the middle Precambrian. The only fossil reefs specifically researched for these microendoliths are of Triassic and Upper Jurassic age. Boring worms appear in reefs in the Lower Cambrian. Boring sponges and bivalves first appear also in the lower Paleozoic, but do not become abundant in reefs until the Triassic. Effective substrate excavating grazers are relatively young geologically: Patellids and substrate excavating Echinoids evolved in the Triassic but did not become important bioeroders until the Jurassic or Cretaceous. Scarid fishes are even younger, the oldest representatives having been found in the Miocene. Thus, it seems that the intensity of bioerosion changed significantly during earth history. This may have had consequences for diversity of reef organisms, quality and quantity of reef debris, for diagenesis and record of reef rock.     

The origins of Dinosauria: much ado about nothing

Research this century has greatly improved our knowledge of the origin and early radiation of dinosaurs. The unearthing of several new dinosaurs and close outgroups from Triassic rocks from various parts of the world, coupled with improved phylogenetic analyses, has set a basic framework in terms of timing of events and macroevolutionary patterns. However, important parts of the early dinosauromorph evolutionary history are still poorly understood, rendering uncertain the phylogenetic position of silesaurids as either non‐dinosaur Dinosauriformes or ornithischians, as well as that of various early saurischians, such as Eoraptor lunensis and herrerasaurs, as either noneusaurischians or members of the sauropodomorph or theropod lineages. This lack of agreement in part derives from a patchy distribution of traits among early members of the main dinosauromorph lineages and requires a more meticulous assessment of characters and homologies than those recently conducted. Presently, the oldest uncontroversial dinosaur records come from Late Triassic (Carnian) rocks of South America, southern Africa and India, hinting at a south‐western Pangaea origin of the group. Besides, macroevolutionary approaches suggest that the rise of dinosaurs was a more gradual process than previously understood. Obviously, these tentative scenarios need to be tested by new fossil finds, which should also help close the major gaps recognized in the fossil record of Triassic dinosauromorphs.     

Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed ( Alternanthera philoxeroides ) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.     

广西凤山、西林等地中三叠统上部双壳类分带研究*

讨论广西西部中三叠统上部的双壳类生物地层序列,建立5个化石带;并附述黔南和滇东南晚三叠世1个双壳类带.这些带自下而上为:(1) Daonella (Longidaonella) producta-D. americana 带,属晚安尼早中期;(2) D. moussoni 高峰带,属晚安尼晚期;(3) D. rieberi-D. indica 带,属早拉丁期;(4) D. kotoi-D. jilongensis 带,属晚拉丁早期;(5) Halobia subcomata-D. varifurcata带,属晚拉丁晚期;(6) H. rugosoides-H. bifurcata 带,属早卡尼期.文内简要描述了各带的主要标志化石23种.     

Climate and vegetational regime shifts in the late Paleozoic ice age earth

The late Paleozoic earth experienced alternation between glacial and non-glacial climates at multiple temporal scales, accompanied by atmospheric CO₂ fluctuations and global warming intervals, often attended by significant vegetational changes in equatorial latitudes of Pangaea. We assess the nature of climate–vegetation interaction during two time intervals: middle–late Pennsylvanian transition and Pennsylvanian–Permian transition, each marked by tropical warming and drying. In case study 1, there is a catastrophic intra-biomic reorganization of dominance and diversity in wetland, evergreen vegetation growing under humid climates. This represents a threshold-type change, possibly a regime shift to an alternative stable state. Case study 2 is an inter-biome dominance change in western and central Pangaea from humid wetland and seasonally dry to semi-arid vegetation. Shifts between these vegetation types had been occurring in Euramerican portions of the equatorial region throughout the late middle and late Pennsylvanian, the drier vegetation reaching persistent dominance by Early Permian. The oscillatory transition between humid and seasonally dry vegetation appears to demonstrate a threshold-like behavior but probably not repeated transitions between alternative stable states. Rather, changes in dominance in lowland equatorial regions were driven by long-term, repetitive climatic oscillations, occurring with increasing intensity, within overall shift to seasonal dryness through time. In neither case study are there clear biotic or abiotic warning signs of looming changes in vegetational composition or geographic distribution, nor is it clear that there are specific, absolute values or rates of environmental change in temperature, rainfall distribution and amount, or atmospheric composition, approach to which might indicate proximity to a terrestrial biotic-change threshold.     

Shell bone histology indicates terrestrial palaeoecology of basal turtles

The palaeoecology of basal turtles from the Late Triassic was classically viewed as being semi-aquatic, similar to the lifestyle of modern snapping turtles. Lately, this view was questioned based on limb bone proportions, and a terrestrial palaeoecology was suggested for the turtle stem. Here, we present independent shell bone microstructural evidence for a terrestrial habitat of the oldest and basal most well-known turtles, i.e. the Upper Triassic Proterochersis robusta and Proganochelys quenstedti. Comparison of their shell bone histology with that of extant turtles preferring either aquatic habitats or terrestrial habitats clearly reveals congruence with terrestrial turtle taxa. Similarities in the shell bones of these turtles are a diploe structure with well-developed external and internal cortices, weak vascularization of the compact bone layers and a dense nature of the interior cancellous bone with overall short trabeculae. On the other hand, 'aquatic' turtles tend to reduce cortical bone layers, while increasing overall vascularization of the bone tissue. In contrast to the study of limb bone proportions, the present study is independent from the uncommon preservation of appendicular skeletal elements in fossil turtles, enabling the palaeoecological study of a much broader range of incompletely known turtle taxa in the fossil record.     

Triassic nannoplankton limestones of deep basin origin in the central mediterranean region

Samples of Triassic pelagic limestones collected in several regions of the Central Mediterranean area were examined with the electron microscope. They show calcareous nannoplankton existing already from Middle Triassic time. The Middle Triassic nannofossils are contained in nodular limestones such as the “Ammonitico Rosso” of Epidaurus (Greece); the Upper Triassic ones are contained in nodular limestones like the Ammonitico Rosso of Hallstatt (Austria), as well as in Halobia cherty limestones of the Pindos basin in Italy, Yugoslavia and Greece. In the Halobia limestones it is possible, in spite of the recrystallization effects, to recognize such an abundance of organic forms — even if in fragments — that we can suppose the original sediment to be a kind of nannoplankton ooze.Because of the very low accumulation rate (few metres/m.y.) of the Middle Triassic Ammonitico Rosso limestones, the coccoliths cannot be considered a significant factor of carbonate pelagic sedimentation at this time; the very high accumulation rate (up to 25 metres/m.y.) of the Upper Triassic cherty limestones in the Pindos basin, on the contrary, allows the nannoplankton to be considered a very important agent of calcareous pelagic sedimentation. The authors hypothesize that the Dogger-Malm, mainly represented by radiolarites, was only a temporary interlude in the Mesozoic pelagic carbonate sedimentation.     

A NEW SPECIES OF CLEVOSAURUS (LEPIDOSAURIA: RHYNCHOCEPHALIA) FROM THE UPPER TRIASSIC OF RIO GRANDE DO SUL, BRAZIL

Abstract:  Well-preserved cranial remains of a small sphenodontian lepidosaur from the Upper Triassic Caturrita Formation of Rio Grande do Sul, Brazil, are the first record of the genus Clevosaurus Swinton, 1939 from South America. They represent a new species, Clevosaurus brasiliensis , which is distinguished by a very short antorbital region of the skull (corresponding to about 20 per cent of skull length) and the presence of teeth in addition to two longitudinal rows on the pterygoid. C. brasiliensis most closely resembles C. bairdi from the Lower Jurassic of Nova Scotia (Canada) and C. mcgilli from the Lower Jurassic of Yunnan (China). The discovery of Clevosaurus in the Upper Triassic of southern Brazil provides a significant range extension of this widely distributed sphenodontian genus. Along with other recent finds, it also suggests that there may have been less biotic provincialism among terrestrial vertebrates during the Late Triassic than has previously been assumed.     

Evolution of the Indian Ocean and the drift of India: A vicariant event

The contemporary hypothesis of plate tectonics allows for the dismembering of a Mesozoic megacontinental land mass, Pangaea, into two supercontinents, Laurasia and Gondwana. Encompassing the current position of the Indo-West Pacific Oceans, the Tethys Ocean was circumglobal from the Jurassic to the Miocene and separated these two supercontinents. Fragmentation of Gondwana allowed for the northward track of India across the Tethys to collide with Asia, closing the Indo-Mediterranean-Atlanto seaway by early Miocene in conjunction with the suturing of Africa and Europe. It is hypothesized that the evolution of the Indian Ocean and the drift of the Indian Plate represent vicariant events which explain patterns of faunal endemism recognized in the region today.     

Biogeography of limno-terrestrial Tardigrada, with particular reference to the Antarctic fauna

The Tardigrada is a cosmopolitan phylum of pre-Pangaean origin, yet tardigrade families and genera show distinct biogeographic components isolated by two major geological events. Separate Laurasian and Gondwanan familial clusters correlate with the Triassic disintegration of Pangaea, while discrete Antarctic, Australian and New Zealand familial/generic clusters relate to the subsequent Jurassic/Cretaceous disintegration of Gondwana.     

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.     

Linkichnus Terebrans New Ichnogenus et Ichnospecies,an Insect Boring from the Late Triassic of the Germanic Basin,Southern Germany

The proposed new ichnotaxon Linckichnus terebrans n. ig. et isp., is a small test-tube shaped boring produced by insects in a sphenophyte stem and occurs in Upper Triassic (Keuper) terrestrial deposits.     

Panbiogeography of Nothofagus (Nothofagaceae): analysis of the main species massings

Aim The aim of this paper is to analyse the biogeography of Nothofagus and its subgenera in the light of molecular phylogenies and revisions of fossil taxa. Location Cooler parts of the South Pacific: Australia, Tasmania, New Zealand, montane New Guinea and New Caledonia, and southern South America. Methods Panbiogeographical analysis is used. This involves comparative study of the geographic distributions of the Nothofagus taxa and other organisms in the region, and correlation of the main patterns with historical geology. Results The four subgenera of Nothofagus have their main massings of extant species in the same localities as the main massings of all (fossil plus extant) species. These main massings are vicariant, with subgen. Lophozonia most diverse in southern South America (north of Chiloé I.), subgen. Fuscospora in New Zealand, subgen. Nothofagus in southern South America (south of Valdivia), and subgen. Brassospora in New Guinea and New Caledonia. The main massings of subgen. Brassospora and of the clade subgen. Brassospora/subgen. Nothofagus (New Guinea–New Caledonia–southern South America) conform to standard biogeographical patterns. Main conclusions The vicariant main massings of the four subgenera are compatible with largely allopatric differentiation and no substantial dispersal since at least the Upper Cretaceous (Upper Campanian), by which time the fossil record shows that the four subgenera had evolved. The New Guinea–New Caledonia distribution of subgenus Brassospora is equivalent to its total main massing through geological time and is explained by different respective relationships of different component terranes of the two countries. Global vicariance at family level suggests that Nothofagaceae/Nothofagus evolved largely as the South Pacific/Antarctic vicariant in the breakup of a world‐wide Fagales ancestor.