On the Turonian origin of the Goniocamax-Belemnitella stock (Cephalopoda,Coleoidea)

A new subgenus within the genus Goniocamax, Progoniocamax nov. subgen., is described. It includes the Late Turonian-Early Coniacian species “Actinocamax” surensis Naidin and Actinocamax intermedius Arkhangelsky. The new subgenus represents a transition between the genera Praeactinocamax and Goniocamax. It occurs in the Central Russian Subprovince. From a geographical point of view, it indicates the origin of the Goniocamax-Belemnitella stock to be situated in the Russian Platform (Central Russian Subprovince, East European Province) during the Late Turonian. The origin of the Upper Cretaceous belemnitellid Goniocamax Naidin, and consequently the Belemnitella-stock, is explained herein by the deepening of the pseudoalveolus and the gradual calcification of the anterior part of the rostrum. The occurrence of the so-called “bottom [base] of ventral fissure”, a major morphological feature known in the later belemnitellids (Goniocamax, Gonioteuthis, Belemnitella), the enlarging of the ventral fissure, and the calcification of its surrounding area, enabled a gradualistic increase of the rostra within the Progoniocamax-Goniocamax-Belemnitella evolutionary lineage.     

Ordovician conodont biogeography – reconsidered

Review of the traditional separation of global Ordovician conodont distribution into the North American Midcontinent Province (NAMP) and North Atlantic Province (NAP) reveals a confusing variety of concepts and definitions that hinder biogeographic analysis. Use of this twofold scheme and its subsequent variants should be­discontinued in favour of the more detailed divisions proposed here. Major biogeographical entities of the Shallow-Sea and Open-Sea Realms, separated by the shelf-slope break, are both further subdivisible into Tropical, Temperate and Cold Domains. In the Cold domains, faunal differences between the two Realms and their subdivisions are not easily discernible, since biofacies zones and different habitats were highly condensed. Faunal differences are amplified in the tropical regions, where the North American Midcontinent Province and North Atlantic Province were originally defined. Recognition of endemic taxa is essential for finer classification within domains of the Shallow-Sea Realm (SSR). Our preliminary analysis of Early Ordovician conodont distribution identifies the Laurentian Province (in the Tropical Domain), Australian­(Tropical Domain), North China (Tropical Domain), South China (Temperate Domain), Argentine Precordillera (Temperate Domain) and Balto-Scandian Province (in the Cold Domain). The Open-Sea Realm (OSR) is dominated by cosmopolitan and widespread taxa, and formal subdivision at provincial level is yet to be achieved. The North Atlantic Province encompasses both the Open-Sea Realm and the Temperate and Cold Domains of the Shallow-Sea Realm. The North American Midcontinent Province sensu stricto is more or less equivalent to the Laurentian Province, representing shallow-water regions fringing Laurentia; in a broader sense the North American Midcontinent Province includes all provinces of the Tropical Domain within the Shallow-Sea Realm.     

The Early Cretaceous of North-East Greenland: A crossroads of belemnite migration

Recent field work in Lower Cretaceous successions of Traill Ø and Wollaston Forland, North-East Greenland, have resulted in c. 2350 belemnite guards collected bed-by-bed from the upper Ryazanian – Hauterivian. The most common belemnite genera observed, Acroteuthis, Pachyteuthis, and Cylindroteuthis are of boreal-arctic affinities and closely related to NW European and Siberian faunas. Other taxa, including Hibolithes (common), Pseudobelus (relatively common) and Duvalia (rare), show faunal links to both NW European and Mediterranean faunas. This paper describes and discusses these findings in their taxonomic, biostratigraphic, palaeobiogeographic, palaeoecologic and palaeoceanographic context. In particular, the occurrence of Pseudobelus which is common in the circum Mediterranean area, is remarkable since it is the first observation of this Tethyan genus in the entire Boreal Realm. The palaeoecological interpretation of these observations result in the recognition of four different palaeobiogeographic belemnite assemblages for the Boreal Realm: 1) North-East Greenland, 2) Spitsbergen, 3) NW Europe and 4) Siberia. In contrast to the other assemblages, the belemnite faunas of North-East Greenland consist of a) Boreal-Arctic elements, b) Boreal-European taxa, c) endemic belemnites of Tethyan ancestry, and d) Tethyan species. These findings make North-East Greenland part of an immigration route from the Tethyan Realm via the north Atlantic to the high Boreal. This allowed Tethyan species, which are otherwise unknown from the Boreal Realm, to reach North-East Greenland. The occurrence of the Tethyan genus Pseudobelus in North-East Greenland also supports the interpretation of this taxon as a hemipelagic dweller, capable of crossing major distances. The belemnite patterns further suggest the existence of a proto Gulf-stream, documenting a south-to-north flow of warm surface waters as far north as Greenland already in the earliest Cretaceous (Valanginian). This has substantial implications for the interpretation of Early Cretaceous climate and oceanic current systems, as well as for the palaeobiology of belemnites.     

A Ceratopsian Dinosaur from the Lower Cretaceous of Western North America,and the Biogeography of Neoceratopsia

The fossil record for neoceratopsian (horned) dinosaurs in the Lower Cretaceous of North America primarily comprises isolated teeth and postcrania of limited taxonomic resolution, hampering previous efforts to reconstruct the early evolution of this group in North America. An associated cranium and lower jaw from the Cloverly Formation (?middle–late Albian, between 104 and 109 million years old) of southern Montana is designated as the holotype for Aquilops americanus gen. et sp. nov. Aquilops americanus is distinguished by several autapomorphies, including a strongly hooked rostral bone with a midline boss and an elongate and sharply pointed antorbital fossa. The skull in the only known specimen is comparatively small, measuring 84 mm between the tips of the rostral and jugal. The taxon is interpreted as a basal neoceratopsian closely related to Early Cretaceous Asian taxa, such as Liaoceratops and Auroraceratops. Biogeographically, A. americanus probably originated via a dispersal from Asia into North America; the exact route of this dispersal is ambiguous, although a Beringian rather than European route seems more likely in light of the absence of ceratopsians in the Early Cretaceous of Europe. Other amniote clades show similar biogeographic patterns, supporting an intercontinental migratory event between Asia and North America during the late Early Cretaceous. The temporal and geographic distribution of Upper Cretaceous neoceratopsians (leptoceratopsids and ceratopsoids) suggests at least intermittent connections between North America and Asia through the early Late Cretaceous, likely followed by an interval of isolation and finally reconnection during the latest Cretaceous.     

A New Hadrosauroid Dinosaur from the Early Late Cretaceous of Shanxi Province,China

Background

The origin of hadrosaurid dinosaurs is far from clear, mainly due to the paucity of their early Late Cretaceous close relatives. Compared to numerous Early Cretaceous basal hadrosauroids, which are mainly from Eastern Asia, only six early Late Cretaceous (pre-Campanian) basal hadrosauroids have been found: three from Asia and three from North America.

Methodology/Principal Findings

Here we describe a new hadrosauroid dinosaur, Yunganglong datongensis gen. et sp. nov., from the early Late Cretaceous Zhumapu Formation of Shanxi Province in northern China. The new taxon is represented by an associated but disarticulated partial adult skeleton including the caudodorsal part of the skull. Cladistic analysis and comparative studies show that Yunganglong represents one of the most basal Late Cretaceous hadrosauroids and is diagnosed by a unique combination of features in its skull and femur.

Conclusions/Significance

The discovery of Yunganglong adds another record of basal Hadrosauroidea in the early Late Cretaceous, and helps to elucidate the origin and evolution of Hadrosauridae.     

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

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

Cenomanian belemnites from the teutoburger Wald,NW Germany

A small fauna of belemnites collected recently from the Cenomanian deposits of the Teutoburger Wald, NW Germany, is described. It includes the belemnopseidNeohibolites ultimus (d’Orbigny) and the belemnitellidsBelemnocamax boweri Crick,Actinocamax primus Arkhangelsky, andA. plenus (Blainville). The Teutoburger Wald is thus one of the few regions in NW Europe, where these four species occur. The Cenomanian belemnite events, theultimus eustato-event, theprimus eco-event, andme plenus eustato-event, are discussed.     

A new hadrosaurid dentary from the latest Maastrichtian of the Pyrenees (north Spain) and the high diversity of the duck-billed dinosaurs of the Ibero-Armorican Realm at the very end of the Cretaceous

In the latest Maastrichtian, the European hadrosauroid fauna was more diverse than those of North America and Asia. The European record of hadrosaurid dentaries is an example of this diversity, and most of the sites with mandibular remains are located in the Ibero-Armorican Realm. Within the Iberian Peninsula, most of the remains are located in the Tremp Basin (South Central Pyrenees). Two of the three valid hadrosaurid taxa defined in this basin are from the Blasi sites (Arén, Huesca province): Arenysaurus ardevoli (Blasi-3) and Blasisaurus canudoi (Blasi-1). A new locality in Blasi (Blasi 3.4) has provided a new dentary from an indeterminate euhadrosaurid. This dentary presents some characters intermediate between Arenysaurus and Blasisaurus, some characters similar to Pararhabdodon isonensis (from the nearby province of Lleida), and some characters of its own. Nevertheless, due to its fragmentary character, without dentition or its edentulous anterior part, it cannot be determined above the level of Euhadrosauria. It thus represents a fourth Iberian euhadrosaurian taxon in the Ibero-Armorican Realm, different from Arenysaurus, Blasisaurus and Pararhabdodon, increasing the diversity of hadrosauroids in this realm at the very end of the Cretaceous.     

A second hadrosauroid dinosaur from the early Late Cretaceous of Zuoyun,Shanxi Province,China

A second basal hadrosauroid dinosaur, Zuoyunlong huangi gen. et sp. nov., is reported from the early Late Cretaceous Zhumapu Formation in Zuoyun County, Shanxi Province, northern China. Zuoyunlong preserves a partial right ilium and ischium and is unique in having a very short postacetabular process 50% as long as the iliac central plate. Our cladistic analysis recovers Zuoyunlong as the most basal Late Cretaceous hadrosauroid, with a sister taxon relationship with Probactrosaurus from the late Early Cretaceous of Inner Mongolia. Including Zuoyunlong, four Cenomanian basal hadrosauroids have been recorded, and the two taxa in North America (Eolambia and Protohadros) represent the earliest known hadrosauroids outside of Asia. In the light of the proposed phylogenetic topology and biogeographic data, the discovery of Zuoyunlong indicates that the first dispersal of hadrosauroids from Asia to North America probably happened around the boundary between the Early and Late Cretaceous.     

A new delthyridoid spiriferid genus from the Dra Valley, Morocco, and its phylogenetic affinities (Brachiopoda, Lower Devonian)

Multiplicatispirifer, a new delthyridoid brachiopod genus with a gently divergent capillate micro-ornamentation and bifurcating costae on flanks, sulcus and fold, has been identified in the Mdâouer-el-Kbîr Formation of the Dra Valley (southern Anti-Atlas, Morocco). Its type speciesMultiplicatispirifer foumzguidensis n. gen., n. sp., at present the only species of this genus, is described. This form has hitherto been determined asFimbrispirifer orStruveina by various authors. In this work, the new genus is compared with the multiplicate and capillate spiriferid genera with bifurcating costae which also have costae in the sulcus and on the fold:Costispirifer andPerryspirifer from North America,Elymospirifer from South China,Borealispirifer from Asia andMultispirifer from Central Europe. The relation ofMultiplicatispirifer to the fimbriateFimbrispirifer, Struveina andVandercammenina, from which it is distinguished by its capillate micro-ornamentation, is discussed. The new genus is hitherto only known from the Moroccan southern Anti-Atlas. Its colsest relatives, taxa ofCostispirifer, occur in the Eastern American Realm (Gaspé, Canada and the eastern United States) and the Nevada Province (western United States).Multiplicatispirifer occurrences are restricted to the Lower/Upper Emsian boundary interval (upper Lower Devonian). The reasons for a possible migration corridor between North America and North Africa before Emsian time are discussed.     

Global historical biogeography of hadrosaurid dinosaurs

Hadrosaurids were the most derived ornithopods and amongst the most diverse herbivore dinosaurs during the Late Cretaceous of Europe, Asia, and the two Americas. Here, their biogeographical history is reconstructed using dispersal‐vicariance analysis (DIVA). The results showed that Hadrosauridae originated in North America and soon after dispersed to Asia no later than the Late Santonian. The most recent common ancestor of Saurolophidae (= Saurolophinae + Lambeosaurinae) is inferred to have been widespread in North America and Asia. The split between saurolophines and lambeosaurines occurred in response to vicariance no later than the Late Santonian: the former clade originated in North America, whereas the latter did so in Asia. Saurolophine biogeographical history included a minimum of five dispersal events followed by vicariance. Four of these dispersals were inferred to have occurred from North America to Asia during the Campanian and Early Maastrichtian, whereas a fifth event represented a southward dispersal from North to South America no later than the Late Campanian. The historical biogeography of lambeosaurines was characterized by an early evolution in Asia, with a Campanian dispersal to the European archipelago followed by vicariance. Reconstruction of the ancestral areas for the deepest nodes uniting the more derived lambeosaurines clades (‘hypacrosaurs’, ‘corythosaurs’, and ‘parasaurolophs’) is ambiguous. The split between North American and Asian clades of ‘hypacrosaurs’ and ‘parasaurolophs’ occurred in response to vicariance during the Campanian. The evolutionary history of North American ‘hypacrosaurs’ and ‘parasaurolophs’ was characterized by duplication events. The latter also characterized the Late Campanian ‘corythosaurs’, which remained restricted to North America. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 503–525.     

A new crocodylian from the late Maastrichtian of Spain: implications for the initial radiation of crocodyloids

Background

The earliest crocodylians are known primarily from the Late Cretaceous of North America and Europe. The representatives of Gavialoidea and Alligatoroidea are known in the Late Cretaceous of both continents, yet the biogeographic origins of Crocodyloidea are poorly understood. Up to now, only one representative of this clade has been known from the Late Cretaceous, the basal crocodyloid Prodiplocynodon from the Maastrichtian of North America.

Methodology/Principal Findings

The fossil studied is a skull collected from sandstones in the lower part of the Tremp Formation, in Chron C30n, dated at −67.6 to 65.5 Ma (late Maastrichtian), in Arén (Huesca, Spain). It is located in a continuous section that contains the K/P boundary, in which the dinosaur faunas closest to the K/P boundary in Europe have been described, including Arenysaurus ardevoli and Blasisaurus canudoi. Phylogenetic analysis places the new taxon, Arenysuchus gascabadiolorum, at the base of Crocodyloidea.

Conclusions/Significance

The new taxon is the oldest crocodyloid representative in Eurasia. Crocodyloidea had previously only been known from the Palaeogene onwards in this part of Laurasia. Phylogenetically, Arenysuchus gascabadiolorum is situated at the base of the first radiation of crocodyloids that occurred in the late Maastrichtian, shedding light on this part of the cladogram. The presence of basal crocodyloids at the end of the Cretaceous both in North America and Europe provides new evidence of the faunal exchange via the Thulean Land Bridge during the Maastrichtian.     

Designation of lectotypes forGonioteuthis westfalicagranulata andg. granulataquadrata

Lectotypes are designated forGonioteuthis westfalicagranulata (Stolley, 1897) andG. granulataquadrata (Stolley, 1897). They are chosen from the preserved syntypes of Stolley, deposited in the Geologisch-Paläontologisches Institut und Museum of the University of Kiel. In order to select representative lectotypes the syntypes of Stolley are compared with populations ofG. westfalicagranulata from Bülten and ofG. granulataquadrata from Weinberg. In addition, specimes of the two taxa bought from Stolley in 1923 and housed in the Mineralogical-Geological Museum of Copenhagen are included in the study.     

Les Brachiopodes de la craie blanche de Meudon (Campanien supérieur) de la collection dˈOrbigny (MNHN,Paris)

Brachiopods from the White Chalk of Meudon (Upper Campanian), listed in the catalogue of dˈOrbigny collection, are presented following the revised classification. This is not without difficulty, due to the few specimens found and/or to the heterogeneity of sets listed under the same number and labelled with the same name. The Brachiopod fauna in the Belemnitella mucronata Zone is well represented, considering the Invertebrate fauna as a whole. Two representatives of Craniiformea and about ten species of Rhynchonelliformea, from several superfamilies shared by two orders, are critically described, considering the recent observations and the last works after dˈOrbigny.     

A new basal hadrosauroid dinosaur from the Late Cretaceous of Uzbekistan and the early radiation of duck-billed dinosaurs

Levnesovia transoxiana gen. et sp. nov., from the Late Cretaceous (Middle–Late Turonian) of Uzbekistan, is the oldest well-documented taxon referable to Hadrosauroidea sensu Godefroit et al. It differs from a somewhat younger and closely related Bactrosaurus from Inner Mongolia (China) by a tall sagittal crest on the parietals and the absence of club-shaped dorsal neural spines in adult specimens. Levnesovia, Bactrosaurus and possibly Gilmoreosaurus represent the earliest radiation of Hadrosauroidea, which took place during the Cenomanian–Turonian and possibly in North America. The second, Santonian-age radiation of Hadrosauroidea included Aralosaurus, Hadrosauridae and lineages leading to Tanius (Campanian) and Telmatosaurus (Maastrichtian). Hadrosauridae appears to be monophyletic, but Hadrosaurinae and Lambeosaurinae originated in North America and Asia, respectively.     

The capsule: an organic skeletal structure in the Late Cretaceous belemnite Gonioteuthis from north‐west Germany

Abstract: An unusual, bilaterally symmetrical black structure that embraces the protoconch and the phragmocone and is overlain by a rostrum has been studied in the Santonian–early Campanian (Late Cretaceous) belemnite genus Gonioteuthis from Braunschweig, north‐west Germany. The structure is here named the capsule. Energy dispersed spectrometry analyses of the capsule show a co‐occurrence of sulphur with zinc, barium, iron, lead and titanium, suggesting their chemical association. The capsule was originally made of organic material that was diagenetically transformed into sulphur‐containing matter. The material of the capsule differs from the chitin of the connecting rings in the same specimens. The capsule has a complex morphology: (1) ventral and dorsal wing‐like projections that are repeated in a breviconic shape of the alveolus, (2) an aperture with lateral lobes and ventral and dorsal sinuses copied by growth lines and (3) a ventral ridge that fits with the position of the fissure in the rostrum. The alveolus in the most anterior part of the rostrum is crater‐like. It is lined with thin, pyritized, laminated material, which appears to be the outermost portion of the capsule attached to the inner surface of the rostrum. A flare along the periphery of the alveolus marks a region where the rostrum was not yet formed, suggesting that the capsule extended beyond the rostrum. Modification of the skeleton in Gonioteuthis comprises a set of supposedly interrelated changes, such as innovation of the organic capsule, partial elimination of the calcareous rostrum and a diminishing of the pro‐ostracum, resulting in the appearance of a new type of pro‐ostracum that became narrower and shorter and lost the spatula‐like shape and gently curved growth lines of a median field that are typical for the majority of Jurassic and Cretaceous belemnites. The partial replacement of a calcareous rostrum with an organic capsule in belemnitellids may have been an adaptive reaction to an unfavourable environmental condition, perhaps related to difficulties in calcium carbonate secretion during the Late Cretaceous that forced animals to reduce carbonate production and to secret an organic capsule around the protoconch and the phragmocone.     

Late devonian facies inter-relationships in bordering areas of the North Atlantic and their palaeogeographic implications

Late Devonian faunal and facies relationships are examined in seven around the North Atlantic — in eastern North America, Greenland, western Europe and northwest Africa. A shallow marine (“littoral”) environment, characterized by the genus Cyrtospirifer, is distinguished from a deeper water (“bathyal”) goniatite-conodont milieu on the one hand, and from the “Old Red Sandstone” terrestial facies bearing plant an d fresh-water fish remains on the other.Current or source directions indicate that an “Acadian Divide” existed, separating west-flowing drainage systems in North America from those flowing to the east on the Afro-European side. All species of the osteolepid Latvius, and the majority of species of Glyptopomus are found on the eastern flank. Conversely, the earliest amphibian, Ichthyostega, may have been confined to the western side of the divide.Palaeogeographic reconstruction places northwest Africa fairly close to the Catskill Delta in the Late Devonian, thus accounting for the presence of an “American fauna” in the former. North—south migration of littoral faunas along the Afro-European shores was, however, apparently inhibited.     

Brachiopod community paleoecology, paleobiogeography, and depositional topography of the Devonian Onondaga Limestone and correlative strata in eastern North America

The Emsian? through early Eifelian Onondaga Limestone of the Appalachian Basin was deposited in a topographic basin and on the carbonate platform which surrounded the basin on the west, north, and northeast. Onondaga strata thin from the platform into the basin. Two sedimentary cycles are present in the sub-Tioga Onondaga of eastern North America. The Edgecliff-Amherstburg represents an interval of transgression, in which epeiric seas spread over much of eastern North America. During the Nedrow-Lucas regression, the interior of the carbonate platform became restricted, resulting in the deposition of evaporites. The Moorehouse-Anderdon transgression continued through the deposition of the Tioga Bentonite, followed by the pre-Speeds-Dundee regression from the craton. Early Eifelian Appalachian Basin Onondaga brachiopod communities, arranged from nearshore to offshore, include the Atrypid-Megakozlowskiella, Atrypid-Levenea, Chonetid, Atlanticocoelia, Ambocoeliid, and Truncalosia Communities. The Onondaga-age Eastern Americas Realm is divided into the Appohimchi Province in the Appalachian Basin and the Michigan Basin-Hudson Bay Lowland Province in the Midwest. The provincial assignment of the James Bay region of Ontario is uncertain; the Eastern Townships of Quebec are near the boundaries both of the two provinces of the Eastern Americas Realm, and of the Eastern Americas Realm and the Old World Realm, the latter realm being probably in the Canadian Maritime Provinces.     

<Emphasis Type="Italic">Protungulatum</Emphasis>, Confirmed Cretaceous Occurrence of an Otherwise Paleocene Eutherian (Placental?) Mammal

Neither pre-Cenozoic crown eutherian mammals (placentals) nor archaic ungulates (“condylarths”) are known with certainty based on the fossil record. Herein we report a new species of the Paleocene archaic ungulate (“condylarth”) Protungulatum from undisputed Late Cretaceous aged rocks in Montana USA based on an isolated last upper premolar, indicating rare representatives of these common early Tertiary mammals appeared in North America a minimum of 300 k  years before the extinction of non-avian dinosaurs. The other 1200 mammal specimens from the locality are characteristic Late Cretaceous taxa. This discovery overturns the current hypothesis that archaic ungulates did not appear in North America until after the Cretaceous/Tertiary (K/T) boundary and also suggests that other reports of North American Late Cretaceous archaic ungulates may be correct. Recent studies, including ours, cannot determine whether Protungulatum does or does not belong to the crown clade Placentalia.