New bethylid and chrysidid wasps (Hymenoptera: Chrysidoidea) from Canadian Late Cretaceous amber

A new bethylid species, Celonophamia granama, and two new chrysidid species, Procleptes eoliami, and P. hopejohnsonae, are described from Late Cretaceous (Campanian) amber collected at the Grassy Lake locality in Alberta, Canada. Within the deposit these taxa constitute the first bethylid, and the second and third chrysidid species to be described, respectively. The new taxa expand the sparse fossil record of Chrysidoidea, particularly that of Chrysididae—a group that was previously represented by only three described species in the Mesozoic. The presence of Celonophamia species in both Canadian amber and Siberian (Taimyr) amber further emphasizes faunal similarities between these two northern Late Cretaceous amber deposits. Given the prevalence of metallic coloration in Chrysididae, the specimens described here also provide evidence for the taphonomic alteration of perceived insect colors in Cretaceous amber inclusions.     

A neotaphonomic experiment in pollen oxidation and its implications for archaeopalynology

Oxidation is considered to be a major taphonomic process involved in differential pollen preservation. An experimental protocol has been developed in the laboratory to test the impact of oxidation on pollen concentrations. Seven modern pollen taxa were selected among common trees, shrubs and herbaceous plants from the European flora, with regard to their typical pollen morphology (Pinus, Alnus, deciduous Quercus, Cichorioideae, Caryophyllaceae, Ericaceae and Poaceae). An aqueous solution of potassium permanganate (KMnO₄) was used to oxidise the pollen grains. The impact of oxidation was measured as a function of increasing oxidation intensity and increasing exposure time. For each taxon, the pollen concentration (grains µl⁻¹) was measured following each oxidation stage. The results show that all pollen taxa are affected by oxidation. Both low oxidant concentrations and short oxidation exposure times led to a fast decrease in pollen concentrations. The thickness of the exine and a specific morphology appear to be the deciding factors controlling the ability to withstand oxidation. A simulation of the impact of oxidation was performed on two model pollen spectra simulating a forested and an open environment. Significant differences were recorded between the initial and final pollen rates. The AP/NAP ratio is modified to a greater extent in the spectrum associated with the open environment. However, over-representation of a specific taxon in a pollen spectrum cannot be attributed to the impact of oxidation alone. For instance, when Cichorioideae are recorded with high frequencies in pollen analyses carried out in an archaeological context, oxidation should not be regarded as the only taphonomic process responsible for this over-representation. The differential conservation of pollen is a complex process in which oxidation represents but one of a variety of interacting taphonomic factors (such as pollen transport, bacterial activity, temperature, pH, hygrometry).     

Late Ordovician vertebrates from the Bighorn Mountains of Wyoming, USA

Abstract:  Late Ordovician vertebrate faunas occur in clastic sedimentary units along the length of the Rocky Mountains from Colorado to Montana, and across the border into Canada. Most research has, however, been conducted on localities in the southern part of the outcrop belt, particularly the Harding Sandstone Formation of Colorado. Micropalaeontological sampling of the coeval South Piney Member (Winnipeg Formation) in the Bighorn Mountains of Wyoming has revealed an abundant vertebrate fauna. The vertebrate assemblage includes a low-abundance fauna of 13 conodont taxa that together indicate an undatus Chronozone age (mid-Mohawkian; mid-Caradoc; Late Ordovician). The pteraspidomorphs Astraspis desiderata Walcott and Eriptychius americanus Walcott are also present together with one new taxon, Eleochera glossa gen. et sp. nov., which is interpreted as a derived stem-gnathostome on the basis of its scale histology and morphology. The fauna bears a strong similarity to that of the Harding Sandstone but is of lower diversity. In particular, it lacks the fine-grained, deeper water component of the Harding Sandstone that contains, inter alia , thelodonts and stem-chondrichthyans.     

A new abelisaurid dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical implications

The Abelisauridae are a family of mainly Cretaceous theropod dinosaurs with a wide distribution across the Gondwanan land masses. Although their presence in Europe was reported twenty-five years ago, it has often been considered as controversial largely because of the incompleteness of the available specimens. We report here the discovery of well-preserved abelisaurid material, including a highly diagnostic braincase, at a Late Cretaceous (late Campanian) locality in the Aix-en-Provence Basin, near the eponym city in south-eastern France. A new abelisaurid taxon is erected, Arcovenator escotae gen. nov., sp. nov., on the basis of cranial and postcranial material. A phylogenetic analysis reveals that the new Abelisauridae from Provence is more closely related to taxa from India and Madagascar than to South American forms. Moreover, Genusaurus, Tarascosaurus and the previous Late Cretaceous discoveries are identified as basal abelisaurids. Contrary to previously proposed palaeobiogeographical models of abelisaurid evolution, the presence of the new taxon in Europe suggests that Europe and Africa may have played a major role in abelisaurid dispersal, which apparently involved crossing marine barriers.     

Faunal and taphonomic analyses of a Late Pleistocene bird-bone assemblage from a cave deposit in north-west Hungary

The paper presents a faunal and taphonomic discussion of the Late Pleistocene bird-bone assemblages excavated in Kálvária Cave n^o 4 near the city of Tatabánya in north-west Hungary. A total of 1150 complete and fragmentary avian remains were recovered from two locations on two separate occasions. Of these, 873 bones could be identified on the species-, family- or order-level. Fifteen of the identified 19 bird taxa could be determined on the species level and they indicate a habitat typical of forest margins. Several species yielded juvenile or subadult specimens, which in view of the breeding period means that these birds died during the summer, most probably between July and September.Each of the two assemblages had distinct characteristics. The high number of kestrel (Falco tinnunculus) long bones in Assemblage I generally included complete skeletal parts from subadult birds, suggesting a colony which bred in the cave, whose chicks died of natural causes. Even though the bones were exposed to a certain degree of water transport and soil corrosion, they were generally well preserved in the cave sediment. Size-characteristic analyses demonstrated that, in spite of the undeveloped morphological features, the sizes of limb bones usually corresponded to those of adult birds.The greater part of the material in Assemblage II originated from a stratified deposit and was dominated by small and fragmented remains of various taxa. The taxa which could be identified and the taphonomic features of the remains suggest that owl pellets formed the core of this bone assemblage.     

Cranial growth and variation in edmontosaurs (Dinosauria: Hadrosauridae): implications for latest Cretaceous megaherbivore diversity in North America

The well-sampled Late Cretaceous fossil record of North America remains the only high-resolution dataset for evaluating patterns of dinosaur diversity leading up to the terminal Cretaceous extinction event. Hadrosaurine hadrosaurids (Dinosauria: Ornithopoda) closely related to Edmontosaurus are among the most common megaherbivores in latest Campanian and Maastrichtian deposits of western North America. However, interpretations of edmontosaur species richness and biostratigraphy have been in constant flux for almost three decades, although the clade is generally thought to have undergone a radiation in the late Maastrichtian. We address the issue of edmontosaur diversity for the first time using rigorous morphometric analyses of virtually all known complete edmontosaur skulls. Results suggest only two valid species, Edmontosaurus regalis from the late Campanian, and E. annectens from the late Maastrichtian, with previously named taxa, including the controversial Anatotitan copei, erected on hypothesized transitional morphologies associated with ontogenetic size increase and allometric growth. A revision of North American hadrosaurid taxa suggests a decrease in both hadrosaurid diversity and disparity from the early to late Maastrichtian, a pattern likely also present in ceratopsid dinosaurs. A decline in the disparity of dominant megaherbivores in the latest Maastrichtian interval supports the hypothesis that dinosaur diversity decreased immediately preceding the end Cretaceous extinction event.     

New discoveries of tetrapods (ichthyostegid‐like and whatcheeriid‐like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium)

The origin of tetrapods is one of the key events in vertebrate history. The oldest tetrapod body fossils are Late Devonian (Frasnian–Famennian) in age, most of them consisting of rare isolated bone elements. Here we describe tetrapod remains from two Famennian localities from Belgium: Strud, in the Province of Namur, and Becco, in the Province of Liège. The newly collected material consists of an isolated complete postorbital, fragments of two maxillae, and one putative partial cleithrum, all from Strud, and an almost complete maxilla from Becco. The two incomplete maxillae and cleithrum from Strud, together with the lower jaw previously recorded from this site, closely resemble the genus Ichthyostega, initially described from East Greenland. The postorbital from Strud and the maxilla from Becco do not resemble the genus Ichthyostega. They show several derived anatomical characters allowing their tentative assignment to a whatcheeriid‐grade group. The new tetrapod records show that there are at least two tetrapod taxa in Belgium and almost certainly two different tetrapod taxa at Strud. This locality joins the group of Devonian tetrapod‐bearing localities yielding more than one tetrapod taxon, confirming that environments favourable to early tetrapod life were often colonized by several tetrapod taxa.     

Small Theropod Teeth from the Late Cretaceous of the San Juan Basin,Northwestern New Mexico and Their Implications for Understanding Latest Cretaceous Dinosaur Evolution

Studying the evolution and biogeographic distribution of dinosaurs during the latest Cretaceous is critical for better understanding the end-Cretaceous extinction event that killed off all non-avian dinosaurs. Western North America contains among the best records of Late Cretaceous terrestrial vertebrates in the world, but is biased against small-bodied dinosaurs. Isolated teeth are the primary evidence for understanding the diversity and evolution of small-bodied theropod dinosaurs during the Late Cretaceous, but few such specimens have been well documented from outside of the northern Rockies, making it difficult to assess Late Cretaceous dinosaur diversity and biogeographic patterns. We describe small theropod teeth from the San Juan Basin of northwestern New Mexico. These specimens were collected from strata spanning Santonian – Maastrichtian. We grouped isolated theropod teeth into several morphotypes, which we assigned to higher-level theropod clades based on possession of phylogenetic synapomorphies. We then used principal components analysis and discriminant function analyses to gauge whether the San Juan Basin teeth overlap with, or are quantitatively distinct from, similar tooth morphotypes from other geographic areas. The San Juan Basin contains a diverse record of small theropods. Late Campanian assemblages differ from approximately co-eval assemblages of the northern Rockies in being less diverse with only rare representatives of troodontids and a Dromaeosaurus-like taxon. We also provide evidence that erect and recurved morphs of a Richardoestesia-like taxon represent a single heterodont species. A late Maastrichtian assemblage is dominated by a distinct troodontid. The differences between northern and southern faunas based on isolated theropod teeth provide evidence for provinciality in the late Campanian and the late Maastrichtian of North America. However, there is no indication that major components of small-bodied theropod diversity were lost during the Maastrichtian in New Mexico. The same pattern seen in northern faunas, which may provide evidence for an abrupt dinosaur extinction.     

Unusual assemblages of Late Cretaceous silicoflagellates from the Canadian Archipelago

Devon and Ellef Ringnes islands in the Canadian Arctic Archipelago contain well-preserved Santonian through early Campanian silicoflagellates. These occur in three unusual assemblages, which comprise three biostratigraphic zones. The oldest, the Vallacerta tumidula Partial Range Zone of Santonian age, contains an extremely variable microflora that includes species of the poorly-known genus Variramus, two new species of Cornua, and a new genus Umpiocha. The Schulzyocha ruppelii Range Zone of early Campanian age includes several species of the new genus Schulzyocha, reflecting quadrate silicoflagellate skeletal morphologies that lack a basal structure. The youngest assemblage includes the poorly-known species Cornua trifurcata, the nominative taxon of the C. trifurcata Partial Range Zone (proposed by McCartney et al., in press c; emended herein), which allows a correlation with a silicoflagellate-bearing sequence from the Horton River area, Northwest Territories of Canada. This report presents information on 24 silicoflagellate taxa, proposes 2 new genera, Schulzyocha and Umpiocha, and describes 13 new species and 1 new combination. All of these new taxa disappear by the top of the Campanian Cornua trifurcata Partial Range Zone. These assemblages reveal a new history of silicoflagellate evolution prior to the late Campanian, the period from which most of our prior knowledge about Cretaceous silicoflagellates was derived. The assemblages show progressive change from skeletons with highly variable morphologies to more stable silicoflagellate skeletal geometry. Two important Corbisema species, C. apiculata and C. archangelskiana, appear to be derived from separate lines within Cornua in the latest Santonian and early Campanian. The important Cretaceous genus Lyramula makes its first appearance at the beginning of the Campanian.     

Taphonomy of Sheridan College Quarry 1, Buffalo, Wyoming: Implications for reconstructing historic dinosaur localities including Utterback's 1902-1910 Morrison dinosaur expeditions

The Upper Jurassic Morrison Formation has yielded some of the most important, voluminous and diverse dinosaur bonebeds in western North America, yet many of its historic sites were excavated during the celebrated period of vertebrate paleontology in western North America referred to as the first and second “Great Dinosaur Rush” (1870-1910s). Because of the large quantity of fossils collected during this era, a considerable amount of data pertaining to patterns of sedimentation, preservation, and paleoecology across broad portions of the Morrison Formation (and indeed many other Mesozoic and Cenozoic units) is still poorly understood. This paper critically re-evaluates the Sheridan College Quarry 1 dinosaur bonebed which lies along the western rim of the Powder River Basin in the region of localities excavated during Utterback's expeditions in the 1900s. Sedimentologically, the bonebed is interpreted as having been formed by episodic flooding events affecting the proximal floodplain depocenter of a meandering river system. Limited evidence of bone abrasion or rounding and progressive upsection changes in bone orientations suggest that minimal transport occurred, but that at least four episodes of overbank flooding resulted in the concentration and burial of attritional, time averaged vertebrate skeletal material that accumulated in topographic lows on the floodplain. Taphonomic analysis indicates that multiple unassociated to partially associated fossil elements excavated represent at least three taxa of sauropod dinosaurs, whereas isolated elements from the site indicate the presence of several other small vertebrate taxa. This work provides significant new information not only about the Sheridan College Quarry 1, but also about local sedimentary and taphonomic conditions that were likely influential to burial and preservation of other nearby Morrison dinosaur localities in the Big Horn Mountains, most notably those excavated during the famous Utterback expeditions. This study highlights the research potential in reconstructing lost data for historic dinosaur localities.     

A remarkable short-snouted horned dinosaur from the Late Cretaceous (late Campanian) of southern Laramidia

The fossil record of centrosaurine ceratopsids is largely restricted to the northern region of western North America (Alberta, Montana and Alaska). Exceptions consist of single taxa from Utah (Diabloceratops) and China (Sinoceratops), plus otherwise fragmentary remains from the southern Western Interior of North America. Here, we describe a remarkable new taxon, Nasutoceratops titusi n. gen. et sp., from the late Campanian Kaiparowits Formation of Utah, represented by multiple specimens, including a nearly complete skull and partial postcranial skeleton. Autapomorphies include an enlarged narial region, pneumatic nasal ornamentation, abbreviated snout and elongate, rostrolaterally directed supraorbital horncores. The subrectangular parietosquamosal frill is relatively unadorned and broadest in the mid-region. A phylogenetic analysis indicates that Nasutoceratops is the sister taxon to Avaceratops, and that a previously unknown subclade of centrosaurines branched off early in the group''s history and persisted for several million years during the late Campanian. As the first well-represented southern centrosaurine comparable in age to the bulk of northern forms, Nasutoceratops provides strong support for the provincialism hypothesis, which posits that Laramidia—the western landmass formed by inundation of the central region of North America by the Western Interior Seaway—hosted at least two coeval dinosaur communities for over a million years of late Campanian time.     

Selachian teeth from Campanian sediments (Upper Cretaceous) of the Münsterland Cretaceous Basin (NW-Germany)

Selachian teeth obtained from trial samples of the Bosenberg quarry near Vorhelm (Beckumer Beds; lower Upper Campanian) and from outcrops along the Hamelbach brook near Stromberg/Oelde (Stromberger Beds; upper Lower Campanian) are presented. Both sites are located in the central southeast of the Münsterland Cretaceous Basin. Fifteen different taxa were identified but only seven could be assigned to a known species. Compared with other taxa the fauna is clearly dominated by squaliform selachians in terms of the number of species and number of specimens. The mixture of selachians presumed to inhabit both deep and shallow waters corresponds with typical taphonomic assemblages of turbiditic deposits and fits very well with the geological setting of the sampled locations.     

Late Devonian - Early Carboniferous vertebrate microremains from the Carnic Alps, northern Italy

Vertebrate microremains from the Late Devonian-Early Carboniferous of the Carnic Alps are predominantly chondrichthyan, with minor placoderm and actinopterygian remains. The faunas are sparse and, with very few exceptions, occur only in conodont-rich pelagic limestones (Pramosio Limestone) representative of the palmatolepid-bispathodid conodont biofacies. Phoebodont and jalodont chondrichthyans, also reflecting open-ocean environments, predominated during the Famennian, and eventually symmoriids seem to predominate during the Early Carboniferous. The presence of Siamodus in this assemblage gives a new locality for this genus known from few regions in the world and allows confirming its stratigraphical range (limpidus Zone) and its relation to deep-water environments. The Late Devonian vertebrate faunas are tropical and cosmopolitan, having much in common with coeval taxa from the North-Gondwanan margins and Asian terranes. Composition of the vertebrate faunas is consistent with the Carnic Alps terrane having occupied a position intermediate between Gondwana and Laurussia, as hypothesized by various authors, but because of sparsity of the taxa represented and the pronounced cosmopolitan nature of both the conodont and vertebrate faunas, the data are not compelling.     

Taxonomic Composition and Trophic Structure of the Continental Bony Fish Assemblage from the Early Late Cretaceous of Southeastern Morocco

The mid-Cretaceous vertebrate assemblage from south-eastern Morocco is one of the most diversified continental vertebrate assemblages of this time worldwide. The bony fish component (coelacanths, lungfishes and ray-finned fishes) is represented by relatively complete specimens and, mostly, by fragmentary elements scattered along 250 kilometres of outcrops. Here we revisit the bony fish assemblage by studying both isolated remains collected during several fieldtrips and more complete material kept in public collections. The assemblage comprises several lungfish taxa, with the first mention of the occurrence of Arganodus tiguidiensis, and possibly two mawsoniid coelacanths. A large bichir cf. Bawitius, is recorded and corresponds to cranial elements initially referred to ‘Stromerichthys’ from coeval deposits in Egypt. The ginglymodians were diversified with a large ‘Lepidotes’ plus two obaichthyids and a gar. We confirm here that this gar belongs to a genus distinctive from Recent gars, contrary to what was suggested recently. Teleosteans comprise a poorly known ichthyodectiform, a notopterid, a probable osteoglossomorph and a large tselfatiiform, whose cranial anatomy is detailed. The body size and trophic level for each taxon are estimated on the basis of comparison with extant closely related taxa. We plotted the average body size versus average trophic level for the Kem Kem assemblage, together with extant marine and freshwater assemblages. The Kem Kem assemblage is characterized by taxa of proportionally large body size, and by a higher average trophic level than the trophic level of the extant compared freshwater ecosystems, but lower than for the extant marine ecosystems. These results should be regarded with caution because they rest on a reconstructed assemblage known mostly by fragmentary remains. They reinforce, however, the ecological oddities already noticed for this mid-Cretaceous vertebrate ecosystem in North Africa.     

Can incomplete taxa rescue phylogenetic analyses from long-branch attraction?

Taxon sampling may be critically important for phylogenetic accuracy because adding taxa can help to subdivide misleading long branches. Although the idea that added taxa can break up long branches was exemplified by a study of "incomplete" fossil taxa, the issue of taxon completeness (i.e., proportion of missing data) has been largely ignored in most subsequent discussions of taxon sampling and long-branch attraction. In this article, I use simulations to test the ability of incomplete taxa to subdivide long branches and improve phylogenetic accuracy in situations of potential long-branch attraction. The results show that for most methods and conditions examined, adding taxa that are only 50% complete may provide similar benefits to adding the same number of complete taxa (suggesting that the advantages of increased taxon sampling may be obtained with less data than previously considered). For parsimony, taxa that are less complete (5% to 25% complete) may often have limited ability to rescue analyses from long-branch attraction. In contrast, highly incomplete taxa can be surprisingly beneficial when using model-based methods. The results also suggest the importance of model-based methods in phylogenetic analyses that combine molecular and fossil data.     

Palynological investigations on some Hypericum taxa (Hypericaceae) growing naturally in Turkey

This study examined the detailed pollen morphological structures of sixteen Hypericum taxa (four endemic, twelve non endemic) including eight sections showing the natural distribution in Turkey: H. sect. Ascyreia (H. calycinum L.), H. sect. Heterophyllum (H. heterophyllum Vent.), H. Sect. Taeniocarpium (Hypericum confertum Choisy subsp. confertum, H. venustum Fenzl, H. linaroides Bosse), H. sect. Drosocarpium (H. montbretii Spach, H. bithynicum Boiss.), H. sect. Crossophyllum (H. adenotrichum Spach, H. orientale L.), H. sect. Olympia (H. olympicum L. subsp. olympicum), H. sect. Origanifolia (H. origanifolium Willd., H. avicularifolium Jaub and Spach subsp. depilatum (Freyn and Bornm.) Robson var. depilatum, H. avicularifolium Jaub. and Spach subsp. byzantinum) and H. sect. Hypericum (H. tetrapterum Fries, H. perforatum L., H. triquetrifolium Tura). These taxa were studied under light microscope and scanning electron microscope for the first time. Of the taxa studied, H. tetrapterum has the smallest pollen grains (on average 15.85–17.20 × 15.45–16.05 μm); and H. olympicum subsp. olympicum the largest grains (on average 22.90–23.10 × 21.40–22.10 μm). The pollen grains of 15 taxa are subprolate and one taxon is prolate-spheroidal. The regular pollen grains of all 16 taxa are 3-zonocolporate. Ornamentation is microreticulate; lumina regularly spaced in eight taxa, tectum perforatum; tectal perforations regularly spaced in seven taxa and tectum perforatum; and tectal perforations grouped together in one taxon. Endoaperture is cruciform porus, with short lateral and meridional extensions in ten taxa, cruciform porus, transversally elongated, with very small lateral extensions in one taxon and lalongate colpus in five taxa. Basic pollen types are ten taxa in type X, five taxa in type IV and one taxon in type II.     

Genesis and environmental significance of Upper Cretaceous shell concentrations from the Cauvery Basin,southern India

Upper Cretaceous shallow marine calcarenites of the Upper Campanian–Maastrichtian Kallankurichchi Formation from the Cauvery Basin of southeastern India are rich in shell concentrations dominated by large bivalves with minor admixtures of brachiopods, bryozoans, and larger foraminifera. Rarely, these concentrations occur as lenses or pavements, but usually as beds 10 to 200 cm thick. Depending on the dominant skeletal elements, five types of shell concentrations can be distinguished: Isognomon, Pycnodonte, Ceratostreon, inoceramid (?Platyceramus, Cataceramus), and polyspecific concentrations. These shell concentrations occur in various preservational states which reflect the gradual mixing and mechanical destruction of former communities. Analysis of the taphonomic signatures of their components and of the biofabric suggests that storm-induced waves and currents, reduced sediment input, the settling behaviour of taxa, and a high rate of production of biogenic hardparts were responsible, in varying degrees, for the formation of the various shell concentrations. Accordingly they can be classified as storm wave, proximal tempestite, and primary biogenic concentrations. The shell concentrations formed below fair weather wave base in shallow, subtropical, low-stress environments.     

New sabre-toothed cats in the Late Miocene of Toros Menalla (Chad)

The Toros Menalla Late Miocene Formation (7 Ma), Djurab desert, Chad, has yielded a large number of vertebrate remains, and among them around twenty taxa of carnivores. Recent collections allow us to describe two sabre-toothed felids assigned to the genera Lokotunjailurus and cf. Megantereon. The former does exist in the Nawata Formation, Lothagam, northern Kenya with a slightly derived species and the TM specimens are considered as belonging to a new species. The latter cannot be identified to the species level, but it could be one of the earliest specimens of Megantereon known in the world and it would reinforce the hypothesis of an Old World origin for this taxon. Both allow us to put forward some biogeographical conclusions.