The structure and phylogenetic significance of the conifer Pseudohirmerella delawarensis nov. comb. from the Upper Triassic of North America

The conifer genus Pseudohirmerella Arndt is based on ovulate cone scales of the type species, Pseudohirmerella platysperma (Mägdefrau) Arndt, from the Upper Triassic of Germany, which bear five distal lobes and two paired structures represented by bulges on the adaxial surface. The paired structures were first described as seeds, but have recently been re-interpreted as arils by Arndt. Similar scales from the Passaic Formation at Milford, NJ, USA were described as Glyptolepis delawarensis Bock, and the new combination Pseudohirmerella delawarensis is proposed. A relatively complete concept of Pseudohirmerella is presented, which includes ovulate cones, possible associated pollen cones, associated shoots with Pagiophyllum-Brachyphyllum morphotype leaves and associated, anatomically preserved wood. Based on the ovulate cone morphology and the presence of organic matter lining the concavities on the scale rather than the corresponding bulges on the counterparts, it is likely that the supposed seeds or arils of Pseudohirmerella are actually casts of empty, seed-bearing depressions. Cheirolepidiaceous affinities are likely based on ovulate cone scale morphology, persistent pollen cones, foliage type and details of wood anatomy. The derived ovulate cone scale morphology of Pseudohirmerella indicates a substantial, but mostly undocumented, Triassic diversification of the Cheirolepidiaceae.     

Implications of fossil conifers for the phylogenetic relationships of living families

Fossils have played a central role in our understanding of the evolution of conifers. Interpretation of the seed cone as a compound strobilus and the homologies of the ovuliferous scales of modern conifers with the axillary dwarf shoot of Pennsylvanian forms are based on fossils. Similarly, early evolutionary trends involving the reduction, fusion, and planation of the fertile and sterile elements of the axillary dwarf shoot, leading to structures characteristic of modern families, are documented in Late Permian and Triassic conifers. However, a phylogeny elucidating the derivation of modern families from fossil forms based on shared derived features has been elusive. The present cladistic treatment using 11 characters of ovulate cones and one of pollen grains suggests three phylogenetic groups of Late Paleozoic conifers, represented loosely by the Emporicaceae, Utrechtiaceae, and Majonicaceae of Mapes and Rothwell. The Taxaceae appears to have diverged from ancestors within the Utrechtiaceae, whereas the other modern families owe their origins to the Majonicaceae. The origin of the Taxodiaceae appears to have been biphyletic.Taxodium, Cupressus andSciadopitys are strongly linked toDolmitia of the Majonicaceae, butCryptomeria, Cunninghamia andAraucaria are grouped together and diverge basal to the former taxa.Pinus branches from a position basal to the known genera of the Majonicaceae and all modern families except the Taxaceae.Podocarpus also diverges basal toMajonica but may share an ancestor with this genus;Cepahalotaxus diverges basal to theDolmitiaPseudovoltzia subclade but distal toMajonica. Similarly, the Cheirolepidiaceae originated from basal members of the Majonicaceae and shows no close phylogenetic relationship with any modern family. Except for a strong linkage betweenCycadocarpidium and theAraucariaCunninghamia subclade, genera of the Voltziaceae appear to have branched more or less independently from within the Majonicaceae and show no strong affinity with modern conifers. Thus differences between modern conifer families are due mainly to their divergence from different Paleozoic ancestors.     

THE SYSTEMATIC POSITION OF EOGINKGOITES

Eoginkgoites is a fan-shaped, imparipinnate leaf with a short rachis and long petiole. It was first described from the Upper Triassic Newark Group of Pennsylvania by Bock who assigned the fossil to the Ginkgoales. The fossil has also been found in the Upper Triassic Chinle Formation in Utah and Arizona and in the Newark Group in North Carolina. Investigation of the well-preserved specimens found in the Chinle Formation shows that the leaf has anastomosing venation, a marginal vein and paracytic (syndetocheilic) stomata. These characters indicate that the leaf is bennettitalean and Eoginkgoites is reassigned to the Bennettitales although its shape is perplexing. Eoginkgoites may be an important index fossil to the lower Upper Triassic (middle Carnian) rocks of North America.     

Some Upper Triassic species of Dipteridaceae from Japan and Borneo

Three Upper Triassic species of Dipteridaceae are described. By new finds of twisted arms and fertile pinnae Dictyophyllum japonicum Yokoyama (1891) from the Yamanoi Formation (Upper Carnic to the Lowest Noric) is transferred to Camptopterisjaponica (Yokoyama) Kon'no, comb. nov. Morphological and taxonomical relation of Dictyophyllum falcatum Naito (MS.) from the Middle Carnic formation with other genera and species is discussed with additional data of fertile pinnae. CUxthropteris meniscoides Brongn. is described from near the south-western border of Sarawak; this gives evidence of the presence of the southernmost outpost of the Dipteridaceae flora of south-eastern Asia in Borneo during the Upper Triassic Period.     

Dasycladalean green algae from the Upper Triassic of Mt. Rotonda (Verbicaro Unit, Calabria-Lucania Border, Southern Italy)

Summary A rich and diverse dasycladalean algae association is described from the Upper Triassic succession of Mt. Rotonda (Calabria-Lucania border, Southern Italy). This association consists of:Neoteutloporella rajkae n.sp.,Griphoporella bechst?dti n.sp.,Physoporella zamparelliae n.sp.,Spinaporella andalusica Flügel & Flügel-Kahler, 1984,S.? granadaensis Flügel & Flügel-Kahler, 1984,Chinianella? sp.,Gyroporella sp.,Griphoporella? sp. andPhysoporella aff.leptotheca. Neoteutloporella rajkae n.sp. is characterised by an undulated calcareous skeleton with short acrophore primary laterals bearing a tuft of 4–6 elongate, segmented, trichophore secondary laterals. This species allows to extend back to the Upper Triassic the stratigraphic range of the genusNeoteutloporella, previously known only from Upper Jurassic levels. Griphoporella bechst?dti n.sp. has a cylindrical calcareous skeleton and primary laterals only, consisting of a thin proximal part followed by a swollen portion that pinches out distally and finally opens outward with a cup-like swelling. Physoporella zamparelliae n.sp. is characterised by a calcareous skeleton made by partly welded thin individual sheaths enclosing the laterals. The laterals are piriferous, vertically compressed, roughly triangular both in vertical and in verticillar section. In some specimens they end with a spine-like thin apophysis. This species confirms that the typical Middle Triassic genusPhysoporella survived up into the Norian. The dasycladalean algal association of the Norian of Mt. Rotonda shows some similarities with the algal association found in the Upper Triassic of the Betic Cordillera whereas it is markedly different from the rich association occurring in the Upper Triassic of Sicily and of the Northern Calcareous Alps. This pattern is coupled with a different composition of the platform margin communities: microbial/serpulids bioconstructions in the Upper Triassic of the Calabria-Lucania border and of Alpujarridevs. Dachstein-type reefs in Sicily and the Northern Calcareous Alps. This indicates that the palaeoceanographic and palaeogeographic conditions controlled both the development of the different platform margin and of the different algal assemblages.     

Reinvestigation of Nystroemia pectiniformis Halle, an enigmatic seed plant from the Upper Permian of China

Reinvestigation of Nystroemia pectiniformis Halle from the Upper Shihhotze Formation of Shanxi Province, China, has led to the identification of new and important features of this enigmatic Late Permian seed plant, permitting its typification and diagnosis. After reassembling several of the previously studied specimens to form a single articulated branching system comprising at least four orders of branching, previously unknown features of its branching pattern and morphology have been characterized. First–order axes are wide and branch to one side only, bearing second–order branches either singly or in pairs and of two kinds: one fertile and bearing characteristic ovulate branching systems and the other presumably vegetative. Ovulate second–order axes are narrow and branch to one side only, producing numerous, closely spaced lateral branches in two alternate to sub–opposite rows. Lateral branches are slender and produce numerous ovulate branching systems to one side of the axis only. Ovulate branching systems divide unequally to produce 3–15 ultimate axes of different lengths that are planated. Each ultimate axis bears a single terminal ovule with 180 degree rotational symmetry and two horn–like integumentary projections distally. The other kind of second–order axes are distinct from those bearing ovules; they are wider and longer and branches occur on both sides of the secondary axis, lacking divisions in close proximity to the first–order axis. These have only been observed incomplete although their distinct morphology indicates they are unlikely to be ovulate branches from which ovules/seeds have been shed. Additional organs of the Nystroemia plant are considered, including pollen organs previously assigned by Halle to the same species (displaying its characteristic branching style), and also leaves of Chiropteris reniformis Halle that were probably borne on the larger kind of second–order branches. Implications of Nystroemia on seed plant evolution and distribution are discussed, and it is concluded that this most likely represents a late stratigraphic occurrence of a plesiomorphic hydrasperman–type seed plant with affinities closely allied to members of the Lyginopteridales.     

Two new species of the pinnate microsporophyll Pramelreuthia from the upper Triassic of the southwestern United States

Two new species of the enigmatic gymnosperm microsporophyll Pramelreuthia, found in the Upper Triassic Chinle Formation at five localities in the southwestern United States, provide significant new evidence on key morphological characters of the genus and extend its known geographical range. These new fossils also demonstrate that the genus was polytypic and reveal the plant megafossil sources for several common and geographically widespread dispersed Upper Triassic microfossil taxa. The genus Pramelreuthia, which until this study was known only from a single specimen from the Upper Triassic of Austria, is a planar pinnate structure consisting of a slender naked axis bearing stalked synangia in opposite to subopposite pairs. Synangia of all three species of Pramelreuthia are oval to subrectangular in lateral view and are composed of two adpressed flattened valves each of which contains up to 20 or more elongate, subcylindrical, tapered sporangia that bear nonstriate bisaccate pollen. Pramelreuthia yazzi sp. nov. is slightly smaller than the type species P. haberfelneri, and its synangia contain pollen generally similar in morphology and size to several species of the dispersed pollen taxon Pityosporites, including P. chinleana, P. oldhamensis, and P. devolvens. Pramelreuthia dubielii sp. nov. is much larger than the other two species; its synangia contain pollen similar to the dispersed pollen species Protodiploxypinus americus.     

Evidence for Prosauropod Dinosaur Gastroliths in the Bull Run Formation (Upper Triassic,Norian) of Virginia

Definitive criteria for distinguishing gastroliths from sedimentary clasts are lacking for many depositional settings, and many reported occurrences of gastroliths either cannot be verified or have been refuted. We discuss four occurrences of gastrolith-like stones (category 6 exoliths) not found within skeletal remains from the Upper Triassic Bull Run Formation of northern Virginia, USA. Despite their lack of obvious skeletal association, the most parsimonious explanation for several characteristics of these stones is their prolonged residence in the gastric mills of large animals. These characteristics include 1) typical gastrolith microscopic surface texture, 2) evidence of pervasive surface wear on many of these stones that has secondarily removed variable amounts of thick weathering rinds typically found on these stones, and 3) a width/length-ratio modal peak for these stones that is more strongly developed than in any population of fluvial or fanglomerate stones of any age found in this region. When taken together, these properties of the stones can be explained most parsimoniously by animal ingestion and gastric-mill abrasion. The size of these stones indicates the animals that swallowed them were large, and the best candidate is a prosauropod dinosaur, possibly an ancestor of the Early Jurassic gastrolith-producing prosauropod Massospondylus or Ammosaurus.

Skeletal evidence for Upper Triassic prosauropods is lacking in the Newark Supergroup basins; footprints (Agrestipus hottoni and Eubrontes isp.) from the Bull Run Formation in the Culpeper basin previously ascribed to prosauropods are now known to be underprints (Brachychirotherium parvum) of an aetosaur and underprints (Kayentapus minor) of a ceratosaur. The absence of prosauropod skeletal remains or footprints in all but the uppermost (upper Rhaetian) Triassic rocks of the Newark Supergroup is puzzling because prosauropod remains are abundant elsewhere in the world in Upper Triassic (Carnian, Norian, and lower Rhaetian) continental strata. The apparent scarcity of prosauropods in Upper Triassic strata of the Newark Supergroup is interpreted as an artifact of ecological partitioning, created by the habitat range and dietary preferences of phytosaurs and by the preservational biases at that time within the lithofacies of the Newark Supergroup basins.     

THE MORPHOLOGY OF STIGMARIA STELLATA

Stigmaria stellata Goeppert is a lycopod underground system occurring in Upper Mississippian and equivalent age rocks of Europe and North America. This taxon has previously been based on impressions exhibiting radiating ridges and furrows around each lateral appendage scar and numerous polyhedral projections on the remainder of the axis. Anatomically preserved specimens from the Chester Series (Upper Mississippian) of Illinois reveal that the distinctive surface pattern of this species results from polyhedral wedges of thick-walled cells in the outer cortex. Decortication produces a smooth outer surface that is indistinguishable from that of the much more abundant Stigmaria ficoides. The structure of S. stellata is quite similar to the structure of other petrified stigmarians, but the following are some of the anatomical characters that distinguish it: (1) presence of abundant polyhedral wedges of thick-walled cells in the outer cortex; (2) absence of secondary cortex; (3) very tall rays associated with appendage traces that remain confluent with the secondary xylem to its outer margin; (4) the absence of a connective in the lateral appendages. The anatomical characters of Stigmaria stellata confirm it as a taxon of at least specific rank.     

Integrated conodont and radiolarian biostratigraphy of the upper Norian in Baoshan Block,Southwestern China

Two significant stratigraphical microfossils, conodonts and radiolarians, are usually used for the Upper Triassic chronostratigraphy. The Baoshan Block was located in eastern Tethys during the Late Triassic where the biostratigraphical data of Upper Triassic are still poorly known. We collected new samples from the Hongyan section (HY) for biostratigraphical study. This 24-m-thick section in Dabaozi Village, Baoshan City, is mainly composed of thin-layered limestones, sandstone and siltstone. The conodont fauna is referred to Sevatian 1 (late Norian), in which the species Mockina englandi, Mockina carinata and Mockina mosheri morphotype B are first recognized in the Baoshan Block, and thus eastern Tethys. The Norian radiolarian associations are first reported in the Baoshan Block, which correlate with the biozonation of North America and also that proposed for Central Japan. The radiolarian assemblages found in the analysed samples in HY section can be referred to the Sevatian Betraccium deweveri Zone. The Baoshan Block is a key area for conodont and radiolarian-based correlations between the Tethys, Japan and North American domains.     

A traversodont cynodont from the Middle Triassic (Ladinian) of Baden-Württemberg (Germany)

Nanogomphodon wildi n. gen., n. sp. is based on a tiny lower postcanine tooth from the lower Lettenkeuper (Lower Keuper or Erfurt Formation; Ladinian) of Michelbach an der Bilz (Baden-Württemberg). It represents the first record of a traversodont cynodont from the Middle Triassic of Europe and exhibits a distinctive combination of dental features. Along with recent discoveries of other traversodont taxa from the Upper Triassic of eastern North America,Nanogomphodon indicates the existence of a distinct lineage of these cynodonts in the Northern Hemisphere.      

A comparison of crayfish burrow morphologies: Triassic and Holocene fossil,paleo‐ and neo‐ichnological evidence,and the identification of their burrowing signatures

The architectural and surficial morphologies of crayfish burrows from the Upper Triassic Chinle Formation and Holocene sediments were compared in order to determine: 1) if Triassic burrows could truly be attributed to crayfish activity; 2) how comparable the burrowing mechanisms are; and 3) whether or not a common set of burrowing signatures could be identified for both ancient and modern freshwater crayfish. Materials used in this study include burrows from the members of the Upper Triassic Chinle Formation, casts of modern burrows constructed by Procambarus clarkii Hobbs and Procambarus acutus acutus (Girard) in the laboratory, and casts of naturally constructed modern burrows of Cambarus diogenes di‐ogenes (Girard).

Triassic and Holocene crayfish burrow morphologies exhibit simple to complex architectures, varying degrees of branching, chamber, and chimney development. They also exhibit relatively textured surficial morphologies (bioglyphs) such as scrape and scratch marks, mud‐ and lag‐liners, knobby and hummocky surfaces, pleopod striae, and body impressions. Holocene crayfish construct distinctive burrows due to their conservative limb arrangement, functional morphology, and behavior with respect to environmental stimuli. Similarities between Holocene and Triassic crayfish burrows suggest that extant and Triassic crayfish employed identical burrowing mechanisms. Features of the surficial and architectural morphologies impart a distinctive signature to burrows of both ancient and modern freshwater burrowing crayfish.

Burrowing signatures of crayfish can be used to identify new and previously misinterpreted continental trace fossils. These are useful in studies of the paleohydrogeology, paleoclimatology and paleoecol‐ogy of burrow‐bearing deposits.     

Halimeda: paleontological record and palaeoenvironmental significance

Udoteacean algae, identical or related to Halimeda, have been recorded in shallow-marine carbonate rocks since Upper Triassic. About 30 species have been described, most of which occur in Lower Cretaceous shelf carbonates. These species are conventionally attributed to four “genera” (Arabicodium Elliott, Boueina Toula, Halimeda Lamouroux, Leckhamptonella Elliott), but the validity of these taxa is a matter of discussion (generic or subgeneric position, or synonyma of Halimeda?) owing to wide discrepancies in the classification of fossil and recent species of halimediform algae. The paleoenvironmental setting of the Mesozoic and the Tertiary is comparable with that of recent Halimeda: lagoonal as well as reefal environments are already known from Upper Triassic occurrences. A reinvestigation of Boueina limestones described from Norian-Rhaetian lagoonal carbonates of Western Thailand indicates the important role of the alga (Boueina marondei n. sp.) in sediment accumulation from its very beginnings.

     

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.     

Limb bone histology and growth in Placerias hesternus (Therapsida: Anomodontia) from the Upper Triassic of North America

Abstract: Patterns of bone deposition are reported and deduced from mid‐shaft sections of 21 limb bones of the dicynodont Placerias hesternus from the Placerias Quarry (Upper Triassic), Arizona, USA. All sampled elements of P. hesternus have a large medullary cavity completely filled with bony trabeculae surrounded by dense cortical bone. Dense Haversian bone extends from the perimedullary region to at least the mid‐cortex in all sampled bones. Primary bone in the outer cortex of limb elements of P. hesternus is generally zonal fibrolamellar with a peripheral layer of parallel‐fibred bone. These data suggest periodic rapid osteogenesis followed by slower growth. Among dicynodonts, this strategy is most similar to growth previously reported in other Triassic (Lystrosaurus, Wadiasaurus) and some Permian taxa (Oudenodon, Tropidostoma). An external fundamental system (EFS), suggesting complete or near complete cessation of appositional growth, is present in the largest tibia. This is the first report of EFS in dicynodonts and may represent the attainment of maximum size in P. hesternus. Slow‐growing peripheral bone was observed in elements of varying size in our sample and may support a differential growth pattern between P. hesternus individuals from this locality. A complete growth series of P. hesternus, analysis of Placerias specimens from other localities, and further sampling of other Upper Triassic dicynodonts are needed to better understand a more complete picture of the growth and remodelling patterns that we have initially investigated.     

SPERMOPTERIS,A NEW GENUS OF PTERIDOSPERMS FROM THE UPPER PENNSYLVANIAN SERIES OF KANSAS

Cridland , Arthur A., and John E. Morris . (Kansas U., Lawrence.) Spermopteris, a new genus of pteridosperms from the Upper Pennsylvanian Series of Kansas. Amer. Jour. Bot. 47(10) : 855–859. Illus. 1960.— Spermopteris gen. nov., based upon seed-bearing specimens of the formgenus Taeniopteris, is described from the Lawrence Shale, Pennsylvanian System, of Kansas. The single species S. coriacea (Göppert) comb. nov. is known. Comparison is made with other fertile and supposedly fertile Paleozoic specimens of Taeniopteris and with specimens of T. spatulata from the Rhaetic of Tonkin.     

New and little known blattogryllidae (Insecta: Grylloblattida) from the Permian and Triassic of Russia,Kazakhstan, and Kyrgyzstan

New members of the family Blattogryllidae are described, including Permoblattogryllus praecox gen. et sp. nov. from the Middle Permian Soyana locality in Russia, Protoblattogryllus nedubrovensis from the Lower Triassic Nedubrovo locality in Russia, Madygenocephalus micropteron gen. et sp. nov. (the second known case of brachyptery in grylloblattids), and Costatoviblatta similis sp. nov. from the Middle Triassic Madygen locality in Kyrgyzstan. Protoblattogryllus zajsanicus Storozhenko, 1990 from the Upper Permian Karaungir II locality in Kazakhstan is redescribed. The genus Microblattogryllus Storozhenko, 1990 from Madygen is considered to be a synonym of Protoblattogryllus Storozhenko, 1990. Protoblattogryllus abruptus Storozhenko, 1990 from the Madygen locality is transferred to the genus Mesoblattogryllus Storozhenko, 1990.