Cretaceous Therian Tarsals and the Metatherian-Eutherian Dichotomy

Diverse metatherian and eutherian tarsal remains from the Late Cretaceous (middle-late Turonian) Bissekty Formation, Kyzylkum Desert, Uzbekistan (ca 90 MYA) are described. Their functional and taxonomic properties, along with those of other tarsal evidence, led to a reassessment of polarity hypotheses of therian, metatherian, and eutherian cruropedal attributes, and the consequences of this for phylogeny of taxa. There are calcaneal remains of several types of marsupials, and a single astragalus that probably belongs to one of these. This represents greater taxonomic diversity than the dental record suggests. Exceptionally large and distally extending peroneal processes, and small and steeply angled calcaneocuboid articulations facing mediodistally, as seen in the Early Cretaceous Sinodelphys and other Cretaceous and Paleogene taxa, attest not only to the metatherian status of these specimens, but also to the retention of many ancestral therian features, even more so than in both the Tiupampa and Itaboraí marsupials of the South American Paleocene (both the calcanea and the astragalus suggest therian traits that are decidedly unlike those of symmetrodonts). Calcanea allocated to the deltatherioid species at Bissekty testify unequivocally to their metatherian affinity. The morphology of the best represented sample of eutherian calcanea from Bissekty, presumably of a number of zhelestid species, appears to be more derived than that of the Late Cretaceous/Paleocene Protungulatum in having a much more reduced peroneal process and a calcaneocuboid articulation that faces distally, oriented nearly at a 90° angle to the long axis of the calcaneus. In fact, this distally facing facet, common in later eutherians (except for lineages in the Paleogene record, and various Carnivora), may not be diagnostic of either the protoeutherian, or even of the protoplacentalian, in spite of its presence in Eomaia. Many putatively “basal” lineages have derived characters, hence such outgroups should not be considered the unequivocal repositories of only ancestral character states.     

Entobia exogyrarum (Frič, 1883) from the Upper Cretaceous of the Bohemian Cretaceous Basin

The validity of the little-known ichnospecies Entobia exogyrarum (Fri?) is verified. Shells of an oyster, Rhynchostreon suborbiculatum (Lam.), represent the substrate for the sponge borings. The borings occur in many Upper Cenomanian to Middle Turonian localities of the Bohemian Cretaceous Basin. This study revealed that the sponges attacked shells of living oysters. Entobia exogyrarum (Fri?) also represents one of the shallowest occurrences of Entobia borings in the Upper Cretaceous rocks.     

Synsacrum of a primitive bird from the Upper Cretaceous of Uzbekistan

A partial synsacrum of a primitive bird from the Zenge Kurgan 3 locality in the Kyzyl Kum Desert (Uzbekistan) is described. The vertebrate fauna from this locality is similar to that of the Bissekty Formation of the Dzharakuduk locality (Central Kyzyl Kum, Uzbekistan); however, this synsacrum differs considerably from all other synsacra from Dzharakuduk. The taxonomic position of this bird remains uncertain; it is most similar to some Enantiornithes, although it could have belonged to a primitive ornithuromorph bird. The functional significance of structural differences in the synsacrum of ornithurine and enantiornithine birds is discussed.     

New Study on Dental and Skeletal Features of the Cretaceous “Symmetrodontan” Mammal Zhangheotherium

A new partial skeleton of the Cretaceous “symmetrodontan” mammal Zhangheotherium quinquecuspedens from the Yixian Formation of Liaoning, China has shed light on the dental and skeletal features of this taxon. The new fossil is a juvenile individual of late growth stage, preserved with interesting features of the premolar replacement. This fossil also provides new information on the vertebral column, the pelvis, the hindlimb and pes. Zhangheotherium has a typical diphyodont replacement of its premolars that is characterized by an alternating pattern (p1 → p3 → p2). This alternating replacement of premolars is a derived condition shared by Dryolestes, Slaughteria, and some basal eutherians, and differs from the plesiomorphic sequential replacement of anterior postcanines in eutricondontans, in most multituberculates and in stem mammaliaforms. The calcaneus and astragalus in the ankle joint of Zhangheotherium lack superposition. This shows that the trechnotherian clade, of which Zhangheotherium is a basal taxon, has retained the primitive condition of mammaliaforms in which the astragalus is in juxtaposition with the calcaneus. Coupled with recent evidence from the earliest metatherians and eutherians, this suggests that the superposition of astragalus and calcaneus evolved in parallel in metatherians and eutherians.     

Morphology of the Coracoid of Late Cretaceous Enantiornithines (Aves: Enantiornithes) from Dzharakuduk (Uzbekistan)

Based on serial material from the Turonian of Dzharakuduk (central Kyzyl Kum, Uzbekistan), the morphology of coracoid of enantiornithines is described in detail. Variants of each morphological element are considered; this allows determination of the taxonomic position of enantiornithines even using small coracoid fragments.     

Relationships of the dog-like marsupials, deltatheroidans and early tribosphenic mammals

A cladistic analysis of the major groups of early metatherian mammals shows that the Deltatheroida and South American dog-like Borhyaenoidea are closely related and are placed in a new supercohort Deltatheralia. while all other metatherians are placed in the supercohort Marsupialia. A reassessment of Early Cretaceous mammals with tribosphenic molars shows that metatherians and eutherians apparently evolved independently from a peramuran-like ancestor, and an entoconid. distinctly basined talonid and large protocone were acquired separately in each group. Consequently. the Tribosphenida ( sensu McKenna) is apparently paraphyletic. Aegialodontia are regarded as structurally ancestral to Metatheria but not Eutheria. The known fossil record supports the view that differentation of the stocks which gave rise to metatherians and eutherians apparently occurred in Late Jurassic or Early Cretaceous time.     

Sauropod teeth from the Upper Cretaceous Bissekty Formation of Uzbekistan

The isolated adult teeth of titanosaurian sauropods from the Upper Cretaceous Bissekty Formation at Dzharakuduk, Uzbekistan, differ little in overall structure but show considerable variation in enamel sculpturing and wear patterns. The crown shape of unworn juvenile teeth ranges from lanceolate to conical. Most specimens have enamel texture resembling crumpled paper or completely smooth enamel. Longitudinal grooves along the mesial and distal edges are present on only a few tooth crowns and might be developed on both the labial and lingual sides. Among 252 worn tooth crowns there are eight variants of wear patterns, all possible combinations of 0–2 apical and 0–2 lateral wear facets. The most common is wear pattern A1L0 (one apical facet, no lateral facets; 62.7%). The next most common variant has two apical and no lateral facets (A2L0, 12.3%). These apical wear facets include the primary wear facets, which are produced by an opposing functional tooth, and secondary wear facets, which are produced by a replacing upper tooth coming into contact with the functional lower tooth at a late wear stage. The relative abundance of tooth crowns with two apical wear facets possibly suggests incipient development of a tooth battery in the Bissekty titanosaur.     

On an Ornithomimid dinosaur (Saurischia, Ornithomimosauria) from the Cenomanian of Fergana

A fragmentary coracoid of Ornithomimidae indet. from the Lower Cenomanian of the Abshir River (Kyrgyzstan) is described. This is the first record of this group in a Cenomanian locality of Fergana. The coracoid from the Abshir locality is similar in the lateral deflection of the glenoid to an endemic group of Asiatic Ornithomimidae that includes Anserimimus, Gallimimus, and taxa from the Cenomanian and Turonian of western Uzbekistan. It is most similar to the unnamed ornithomimid from the Cenomanian of Karakalpakia in having a distinct vertical crest that borders anteriorly a depression for the coracobrachialis brevis muscle. Dinosaurs recorded in the Cenomanian of Fergana are reviewed.     

The rise to dominance of the angiosperm kingdom: dispersal,habitat widening and evolution during the Late Cretaceous of Europe

Coiffard, C. & Gomez, B. 2009: The rise to dominance of the angiosperm kingdom: dispersal, habitat widening and evolution during the Late Cretaceous of Europe. Lethaia, Vol. 43, pp. 164–169. The earliest fossil records of angiosperms in Europe occur in the Barremian and consist of freshwater wetland plants. From the Barremian onwards, angiosperms show a stepwise widening of their ecological range with the result that they inhabited most environments by the Cenomanian. Nevertheless, most angiosperms had still restricted habitats, while a few angiosperm trees were confined to disturbed environments, such as channel margins. A Wagner’s Parsimony Method analysis performed on a fossil plant and locality database from the Turonian to the Campanian of Europe indicates continued decrease in richness of ferns and gymnosperms compared with angiosperms, turnover between conifer and palm trees in freshwater‐related swamps at about the Cenomanian/Turonian boundary, and spreading of angiosperm trees through the floodplains. The ecological range of angiosperm trees was increased, being recorded in channel margins from the Cenomanian and spreading over floodplains (e.g. Platanaceae) and swamps (e.g. Arecaceae) by the Campanian. These new ecological ranges and successions went with innovative architectures, such as dicot trees and palm trees. Most living core angiosperm families had their earliest representatives in the Late Cretaceous, which should be considered as the dawn of modern angiosperm forests. □Core angiosperms, Europe, Late Cretaceous, palms, Wagner’s Parsimony Method.     

Biometry of Cenomanian–Turonian placoliths: a proxy for changes of fertility and surface‐water temperature?

Most publications discussing Cenomanian–Turonian calcareous nannofossils focus on abundance fluctuations across the boundary interval. So far, there have been no studies that deal with the influence of palaeoenvironmental changes on the size of common Cenomanian–Turonian nannofossil taxa. The genera Biscutum, Broinsonia, Prediscosphaera, Retecapsa and Watznaueria have therefore been analysed from 19 samples of Cenomanian–Turonian age from the Goban Spur, northeast Atlantic. The genus Biscutum shows a slight decrease of mean length from 4.14 μm in the Cenomanian to 3.94 μm in the Turonian. Broinsonia is marked by a decrease from 6.07 μm in the Cenomanian to 5.64 μm in the Turonian. On the other hand, Prediscospheara increases in size from 4.98 μm in the Cenomanian to 5.61 μm in the Turonian. Two genera (Retecapsa, Watznaueria) show no significant changes in their mean length. The mean size of Biscutum is perhaps controlled by nutrients, where larger specimens may have preferred the more fertile palaeoenvironment of the Late Cenomanian. The size decrease of Biscutum in the Turonian is probably related to reduced nutrient availability. The genus Prediscosphaera spp., may have favoured low‐fertility conditions, as its mean size increases in the Turonian. A worldwide decline of the frequency of Broinsonia spp. during the Cenomanian–Turonian transition implies that this genus is not solely controlled by the nutrient content. The size of Broinsonia spp. may have been therefore influenced by the latest Cenomanian warming event. The increase in sea‐surface temperature may have been unfavourable for Broinsonia spp. as reflected by decreasing mean size and frequency. □Calcareous nannofossils, biometry, morphometry, Oceanic Anoxic Event 2.     

A crinoid concentration Lagerstätte in the Turonian (Late Cretaceous) Conulus Bed (Miechów-Wolbrom area, Poland)

More than one hundred centrodorsals of the comatulid crinoid Glenotremites paradoxus have been found in Turonian deposits (Upper Cretaceous) in the Miechów-Wolbrom area (southern Poland). This is the first dense occurrence of the genus Glenotremites in the Upper Cretaceous of Poland. Furthermore, so many individuals of this species in one level (the so-called Conulus Bed) forming a crinoid Konzentrat-Lagerstätte are very surprising because only disarticulated remains (cirrals and brachials) are encountered in the older (Cenomanian and earliest Turonian) and younger (late Turonian) deposits. The Glenotremites individuals are accompanied by isocrinids, which prove that stalked forms remained in shallow-water settings for some time after the initiation of the Mesozoic marine revolution.     

A new mosasauroid (Squamata) from the Late Cretaceous (Turonian) of Morocco

Tethysaurus nopcsai gen. et sp. nov. is described on the basis of both cranial and postcranial material from the Late Cretaceous (Early Turonian) of the Goulmima region, southern Morocco. This new mosasauroid is mainly characterized by a parietal table ending posteriorly in two pointed pegs; jugal with a large ascending ramus; splenial with a large and notched dorsomedial process; surangular exposed medially ventral to the coronoid; large paracotylar and parazygosphenal foramina on vertebrae. A phylogenetic analysis shows that Tethysaurus is the sister-group of Mosasauridae. It fills the gap between the aigialosaurids (mainly Cenomanian) and the mosasaurids (known from the Middle-Late Turonian to the Latest Maastrichtian). To cite this article: N. Bardet et al., C. R. Palevol 2 (2003).     

Analysis of Late Jurassic to Early Cretaceous algal debris-facies of the Plassen carbonate platform in the Northern Calcareous Alps (Germany,Austria) and in the Kurbnesh area of the Mirdita zone (Albania): a tool to reconstruct tectonics and palaeogeography of eroded platforms

A characteristic microfacies of the Late Jurassic to Early Cretaceous allodapic Barmstein Limestone of the Northern Calcareous Alps are clasts of wackestones with numerous fragments of calcareous algae (“algal debris-facies”). According to dasycladale palaeocoenoses, several subtypes comprising different associations can be distinguished. One association is characterized by the debris of an unknown large dasycladalean alga reported as dasycladalean alga indet. sp. 1 from different localities in the Northern Calcareous Alps, typically forming a monospecific assemblage. Another microfacies type contains star-like calcitic bodies tentatively referred to the morphospecies Coptocampylodon pantici Ljubović-Obradović and Radoičić, originally described as being from the Turonian of NW-Serbia. Other Coptocampylodon-like bodies represent the calcified tufts of the laterals of Selliporella neocomiensis (Radoičić). The occurrence of Coptocampylodon pantici-like microfossils in the Late Tithonian to Early Berriasian, shows that obviously different species of dasycladaleans display identical to similar shaped tufts of laterals in transverse sections when becoming fragmented. Coptocampylodon pantici Ljubović-Obradović and Radoičić was observed only from different occurrences of Barmstein Limestone, but not from the autochthonous platform carbonates of the Plassen carbonate platform. The Coptocampylodon algal debris-facies is also reported from the Late Jurassic of Albania, Mirdita zone. Occurrences of different types of algal debris-facies in components of mass-flow deposits can be used as a tool to reconstruct eroded carbonate platforms and tectonics, as demonstrated in the Northern Calcareous Alps and the Albanides. Finally, the general occurrences of algal debris-facies in both settings—intra-Tethyan mostly isolated platforms (Alps, Albanides) vs. extended epeiric platforms (Middle East)—are compared and discussed.     

The Cenomanian–Turonian boundary in Jordan: Ammonite biostratigraphy and faunal turnover

Well-exposed fossiliferous Upper Cenomanian–Lower Turonian marine sedimentary rocks are present in west-central Jordan. Ammonites serve as an important faunal marker for this interval and can be used to subdivide the Cenomanian–Turonian transition into two upper Cenomanian biozones (Neolobites vibrayeanus and Vascoceras cauvini) and two lower Turonian biozones (Vascoceras proprium and Choffaticeras segne). A revised stratigraphic range of the Vascoceras cauvini Zone in the study area is proposed, consisting of the Metoicoceras geslinianum and Neocardioceras juddii zones of the standard zonation. Based on intercontinental biostratigraphic correlation, a minor unconformity appears to be present around the Cenomanian–Turonian boundary, and a part of the lower Turonian is probably missing. In addition, a faunal turnover is recorded in the uppermost Cenomanian, marked by the disappearance of most of the Cenomanian taxa, including Costagyra olisiponensis (Sharpe), Ceratostreon flabellatum (Goldfuss), Ilymatogyra africana (Lamarck), Rhynchostreon suborbiculatum (Lamarck), Harpagodes nodosus (Sowerby), and Heterodiadema libycum (Desor). This bioevent is thought to be an effect of the Oceanic Anoxic Event OAE 2; the dramatic shifts in species richness and diversity spanning the Cenomanian–Turonian boundary in the study area occurred in response to the major paleoclimatic and paleoenvironmental perturbations prevailing at that time. The stratigraphic and paleontological patterns studied in Jordan are very similar to those recorded in Egypt in terms of litho- and biostratigraphy, event stratigraphy, and macroinvertebrate content, suggesting the presence of uniform triggering mechanisms and bio-sedimentary responses in the Upper Cretaceous basins of the Middle East and providing clues for a high-resolution correlation between the two areas.     

Facies pattern and sea-level dynamics of the early Late Cretaceous transgression: a case study from the lower Danubian Cretaceous Group (Bavaria,southern Germany)

The facies development and onlap pattern of the lower Danubian Cretaceous Group (Bavaria, southern Germany) have been evaluated based on detailed logging, subdivision, and correlation of four key sections using an integrated stratigraphic approach as well as litho-, bio-, and microfacies analyses. Contrary to statements in the literature, the transgressive onlap of the Regensburg Formation started in the Regensburg–Kelheim area already in the early Early Cenomanian Mantelliceras mantelli ammonite Zone and not in the Late Cenomanian. In the Early Cenomanian, nearshore glauconitic-bioclastic sandstones prevailed (Saal Member), followed by Middle to lower Upper Cenomanian mid-shelf siliceous carbonates intercalated with fine-sandy to silty marls (Bad Abbach Member). Starting in the mid-Late Cenomanian (Metoicoceras geslinianum ammonite Zone), a considerable deepening pulse during the Cenomanian–Turonian Boundary Event (CTBE) initiated the deposition of the deeper shelf silty marls of the Eibrunn Formation, which range into the early Early Turonian. During the CTBE transgression, also the proximal Bodenwöhrer Senke (ca. 40 km NE of Regensburg) was flooded, indicated by the onlap of the Regensburg Formation onto Variscan granites of the Bohemian Massif, overlain by a thin tongue of lowermost Turonian Eibrunn Formation. A detailed record of the positive δ¹³C excursion of the global Oceanic Anoxic Event (OAE) 2 has been retrieved from this shallow-water setting. An integrated approach of bio-, event-, carbon stable isotope and sequence stratigraphy was applied to correlate the sections and to decipher the dynamics of this overall transgressive depositional system. The Cenomanian successions show five prominent unconformities, which correlate with those being known from basins in Europe and elsewhere, indicating their eustatic origin. The rate of sea-level rise during the CTBE suggests glacio-eustasy as a driving mechanism for Late Cenomanian sea-level changes. The Regensburg and Eibrunn formations of the lower Danubian Cretaceous Group are highly diachronous lithostratigraphic units. Their regional distribution and northeast-directed onlap pattern onto the southwestern margin of the Bohemian Massif can readily be explained by the lateral movements of roughly coast-parallel (i.e., NW/SE-trending) facies belts of a graded shelf system transgressing on a northeastward-rising substrate. It took the Cenomanian coastline ca. 6 Ma to transgress from southwest of Regensburg to the topographically elevated granite cliffs southeast of Roding in the Bodenwöhrer Senke (=60 km distance).     

New records of Azhdarchids (Pterosauria,Azhdarchidae) from the late Cretaceous of Russia,Kazakhstan, and Central Asia

A review of 12 azhdarchid localities in Russia, Kazakhstan, Uzbekistan, and Tajikistan is given. New records of unidentifiable azhdarchids from the Khodzhakul (Cenomanian), Tyul’keli (Turonian-Coniacian), Kansai (Santonian), Malaya Serdoba, and Beloe Ozero (Campanian) localities and a new taxon, Aralazhdarcho bostobensis gen. et sp. nov. (Shakh-Shakh, Santonian-Campanian), are described.     

Palaeobiology of tanaidaceans (Crustacea: Peracarida) from Cretaceous ambers: extending the scarce fossil record of a diverse peracarid group

Diverse assemblages of tanaidacean peracarid crustaceans from western Tethyan continental deposits suggest that the group was relatively common in or around ancient resin‐producing forests. Here we report the results of an examination of 13 tanaidacean specimens from three Cretaceous (Albian to Turonian) French amber deposits. Two new species of the fossil family Alavatanaidae are placed in the previously described Early Cretaceous genus Eurotanais: Eurotanais pyrenaensis sp. nov. from Cenomanian Pyrenean amber (Fourtou, Aude) and Eurotanais seilacheri sp. nov. from Turonian Vendean amber (La Garnache, Vendée). The remaining specimens are placed in three newly erected genera and species (but family incertae sedis): Arcantitanais turpis gen. et sp. nov. from Albian–Cenomanian Charentese amber (Archingeay, Charente‐Maritime), and Tytthotanais tenvis gen. et sp. nov. and Armadillopsis rara gen. et sp. nov. from Pyrenean amber. These are the first formally described fossils that might be related to the paratanaoidean families Nototanaidae and Paratanaidae, sharing with these some putatively derived features and providing possible evidence for the antiquity and morphological stability of these families and the suborder Tanaidomorpha. The distinctive features and character combinations of these fossil taxa are discussed in connection with possible relationships to the living lineages of tanaidaceans. Propagation phase‐contrast X‐ray synchrotron microtomography was used to obtain high‐quality 3D images for some fossils. A discussion is provided on the putative palaeobiology of tanaidaceans and the French resiniferous forest ecosystem. The discovery of these new tanaidaceans extends the palaeogeographical distribution and stratigraphical range of the family Alavatanaidae and sheds new light on the palaeoecology and diversity of tanaidaceans in pre‐angiospermous woodlands.     

Late Cretaceous (Cenomanian to Campanian) paleoenvironmental history of the Eastern Canadian margin of the Western Interior Seaway: bonebeds and anoxic events

Upper Cretaceous strata in the Pasquia Hills of the northern Manitoba Escarpment, eastern Saskatchewan, Canada provide a detailed paleoenvironmental and sea-level record of the eastern margin of the Western Interior Seaway. Sediments deposited during the Cenomanian/Turonian Greenhorn marine cycle are dominantly black mudstones deposited in a stratified water column, with bottom-water anoxia recurrently reaching into the photic zone. A middle Cenomanian sea-level lowstand event followed by transgression left a series of bonebeds within the Belle Fourche Member of the Ashville Formation, indicating a sedimentary environment starved of coarse siliciclastics. Maximum sea level resulted in the formation of limestone beds within the Favel Formation, further favoured by reduced terrigenous sediment input compared to the western margin. Limestone sedimentation was followed by a phase of increased freshwater input under lower sea level conditions, and reducing zoo- and phytoplankton diversities. During final Greenhorn regression, eastern Saskatchewan probably turned into a restricted basin severely limiting marine circulation. Poor or absent benthic foraminiferal assemblages and biomarker analysis suggest prevailing watermass stratification throughout the Cenomanian/Turonian transgressive/regressive cycle. This was caused either by a freshwater lid, stratification of Boreal and Tethyan-derived watermasses, or both, to various intensities affected by changing sea level. Basin oxygenation during Niobrara time varies between localities along the eastern margin as documented by presence/absence of benthic and planktic foraminifera.     

Cretaceous (Albian to Turonian) radiolarians from chert blocks of the Moni Mélange (Southern Cyprus)

Well-preserved Cretaceous (Albian–Turonian) radiolarians were extracted from radiolarian-bearing chert olistoliths of the Monagroulli Member within the Moni Mélange (Campanian-Maastrichtian, Southern Cyprus). Four assemblages were distinguished: Middle Albian–Lower Cenomanian (Thanarla spoletoensis Zone), Upper Albian–Lower Cenomanian (Thanarla spoletoensis Zone, Dorypyle? anisa Subzone), lowermost Turonian (base of Alievium superbum Zone) and Lower Turonian (Alievium superbum Zone). The radiolarian assemblages are diverse and have taxonomic composition similar to coeval assemblages of Italy and Spain. The sediments of the Monagroulli Member differ from coeval rocks of the Mamonia Complex (western Cyprus) by the more common presence of radiolarian cherts and may have been formed in the distal part of a continental margin with less input of clastic material. A new spicular radiolarian genus Cyprothamnus with 2 new species (C. multifurcatus and C. moniensis) is described from the Lower Turonian strata.