A longirostrine tyrannosauroid from the Early Cretaceous of China

The fossil record of tyrannosauroid theropods is marked by a substantial temporal and morphological gap between small-bodied, Barremian taxa, and extremely large-bodied taxa from the latest Cretaceous. Here we describe a new tyrannosauroid, Xiongguanlong baimoensis n. gen. et sp., from the Aptian–Albian Xinminpu Group of western China that represents a phylogenetic, morphological, and temporal link between these disjunct portions of tyrannosauroid evolutionary history. Xiongguanlong is recovered in our phylogenetic analysis as the sister taxon to Tyrannosauridae plus Appalachiosaurus, and marks the appearance of several tyrannosaurid hallmark features, including a sharp parietal sagittal crest, a boxy basicranium, a quadratojugal with a flaring dorsal process and a flexed caudal edge, premaxillary teeth bearing a median lingual ridge, and an expanded axial neural spine surmounted by distinct processes at its corners. Xiongguanlong is characterized by a narrow and elongate muzzle resembling that of Alioramus. The slender, unornamented nasals of Xiongguanlong are inconsistent with recent hypotheses of correlated progression in tyrannosauroid feeding mechanics, and suggest more complex patterns of character evolution in the integration of feeding adaptations in tyrannosaurids. Body mass estimates for the full-grown holotype specimen of Xiongguanlong fall between those of Late Cretaceous tyrannosaurids and Barremian tyrannosauroids, suggesting that the trend of increasing body size observed in North American Late Cretaceous Tyrannosauridae may extend through the Cretaceous history of Tyrannosauroidea though further phylogenetic work is required to corroborate this.     

Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez,Central México

Huehuecuetzpalli mixtecus gen. et sp. nov. is characterized by a combination of characters unlike those of any of the previously described Late Jurassic or Early Cretaceous lizards. It has most of the synapomorphies common to modern squamates, but still retains primitive features rare in living taxa. Autapomorphic characters include an anteroposteriorly elongated premaxilla that results in the elongation of the snout and the apparent retraction of the external nares. A small rounded postfrontal and a parietal foramen on the frontoparietal suture suggest affinities with iguanians, but the retention of divided premaxillae, amphicoelous vertebrae, thoracolumbar intercentra, entepicondylar foramen, and a second distal tarsal supports the hypothesis that Huehuecuetzpalli has a more basal position relative to the extant squamates. Although its appearance is late in the fossil record of lizards, Huehuecuetzpalli is the first report of a basal squamate. It provides important information on early transformation of characters in lizard evolution. Many primitive characters present in some modern squamates are usually explained by paedomorphosis; however, these characters are common in early lizards suggesting that derived states may have been fixed later in lizard evolution. If Huehuecuetzpalli is an iguanian, then it would be the earliest known representative of this lineage and extends their fossil record into the Albian. This paper presents an extensive review of the characters and character states used in previously published cladistic analyses of the Squamata.     

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.     

The origin and early evolution of metatherian mammals: the Cretaceous record

Metatherians, which comprise marsupials and their closest fossil relatives, were one of the most dominant clades of mammals during the Cretaceous and are the most diverse clade of living mammals after Placentalia. Our understanding of this group has increased greatly over the past 20 years, with the discovery of new specimens and the application of new analytical tools. Here we provide a review of the phylogenetic relationships of metatherians with respect to other mammals, discuss the taxonomic definition and diagnosis of Metatheria, outline the Cretaceous history of major metatherian clades, describe the paleobiology, biogeography, and macroevolution of Cretaceous metatherians, and provide a physical and climatic background of Cretaceous metatherian faunas. Metatherians are a clade of boreosphendian mammals that must have originated by the Late Jurassic, but the first unequivocal metatherian fossil is from the Early Cretaceous of Asia. Metatherians have the distinctive tightly interlocking occlusal molar pattern of tribosphenic mammals, but differ from Eutheria in their dental formula and tooth replacement pattern, which may be related to the metatherian reproductive process which includes an extended period of lactation followed by birth of extremely altricial young. Metatherians were widespread over Laurasia during the Cretaceous, with members present in Asia, Europe, and North America by the early Late Cretaceous. In particular, they were taxonomically and morphologically diverse and relatively abundant in the Late Cretaceous of western North America, where they have been used to examine patterns of biogeography, macroevolution, diversification, and extinction through the Late Cretaceous and across the Cretaceous-Paleogene (K-Pg) boundary. Metatherian diversification patterns suggest that they were not strongly affected by a Cretaceous Terrestrial Revolution, but they clearly underwent a severe extinction across the K-Pg boundary.     

A report on biocompounds from palm fossil of India

The occurrence of a large number of fossil woods having resemblance in anatomical features with the modern palm genus, Phoenix L in Deccan Intertrappean fossil flora of Maastrichtian-Danian age (i. e. Late Cretaceous and Earliest Tertiary (65-67 my)) indicates the most primitive record of date palm. Present discovery of biocompounds from fossil wood of Phoenix collected from Deccan Intertrappean having affinity with the biocompounds known from modern plant further exemplify the earliest documentation of Phoenix in Indian peninsula.     

The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain

Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids– the Struthiosaurinae.     

The genus Macroteleia Westwood in Middle Miocene amber from Peru (Hymenoptera,Platygastridae s.l., Scelioninae)

A new species of the scelionine genus Macroteleia Westwood (Platygastridae s.l., Scelioninae) is described and figured from a female beautifully preserved in Middle Miocene amber from Peru. Macroteleia yaguarum Perrichot & Engel, sp. n., shows a unique combination of characters otherwise seen independently within its congeners. It is most similar to the modern M. surfacei Brues, but differs from it by the non-foveolate notauli, the contiguous punctures of the vertex, and the continuous propodeum. The new species is the first New World fossil of the genus, suggesting a Cretaceous origin for the group and a relatively old age of the South American, tropical African, and Australian faunas, and a younger age of the modern Holarctic faunas.     

Identification of a New Hesperornithiform from the Cretaceous Niobrara Chalk and Implications for Ecologic Diversity among Early Diving Birds

The Smoky Hill Member of the Niobrara Chalk in Kansas (USA) has yielded the remains of numerous members of the Hesperornithiformes, toothed diving birds from the late Early to Late Cretaceous. This study presents a new taxon of hesperornithiform from the Smoky Hill Member, Fumicollis hoffmani, the holotype of which is among the more complete hesperornithiform skeletons. Fumicollis has a unique combination of primitive (e.g. proximal and distal ends of femur not expanded, elongate pre-acetabular ilium, small and pyramidal patella) and derived (e.g. dorsal ridge on metatarsal IV, plantarly-projected curve in the distal shaft of phalanx III:1) hesperornithiform characters, suggesting it was more specialized than small hesperornithiforms like Baptornis advenus but not as highly derived as the larger Hesperornis regalis. The identification of Fumicollis highlights once again the significant diversity of hesperornithiforms that existed in the Late Cretaceous Western Interior Seaway. This diversity points to the existence of a complex ecosystem, perhaps with a high degree of niche partitioning, as indicated by the varying degrees of diving specializations among these birds.     

Breeding system diversification and evolution in American Poa supersect. Homalopoa (Poaceae: Poeae: Poinae)

Background and Aims Poa subgenus Poa supersect. Homalopoa has diversified extensively in the Americas. Over half of the species in the supersection are diclinous; most of these are from the New World, while a few are from South-East Asia. Diclinism in Homalopoa can be divided into three main types: gynomonoecism, gynodioecism and dioecism. Here the sampling of species of New World Homalopoa is expanded to date its origin and diversification in North and South America and examine the evolution and origin of the breeding system diversity.Methods A total of 124 specimens were included in the matrix, of which 89 are species of Poa supersect. Homalopoa sections Acutifoliae, Anthochloa, Brizoides, Dasypoa, Dioicopoa, Dissanthelium, Homalopoa sensu lato (s.l.), Madropoa and Tovarochloa, and the informal Punapoa group. Bayesian and parsimony analyses were conducted on the data sets based on four markers: the nuclear ribosomal internal tanscribed spacer (ITS) and external transcribed spacer (ETS), and plastid trnT-L and trnL-F. Dating analyses were performed on a reduced Poa matrix and enlarged Poaceae outgroup to utilize fossils as calibration points. A relaxed Bayesian molecular clock method was used.Key Results Hermaphroditism appears to be pleisiomorphic in the monophyletic Poa supersect. Homalopoa, which is suggested to have originated in Eurasia 8·4–4·2 million years ago (Mya). The ancestor of Poa supersect. Homalopoa radiated throughout the New World in the Late Miocene–Early Pliocene, with major lineages originating during the Pliocene to Pleistocene (5–2 Mya). Breeding systems are linked to geographic areas, showing an evolutionary pattern associated with different habitats. At least three major pathways from hermaphroditism to diclinism are inferred in New World Homalopoa: two leading to dioecism, one via gynodioecism in South America and another directly from hermaphroditism in North America, a result that needs to be checked with a broader sampling of diclinous species in North America. A third pathway leads from hermaphroditism to gynomonoecism in Andean species of South America, with strictly pistillate species evolving in the highest altitudes.Conclusions Divergence dating provides a temporal context to the evolution of breeding systems in New World Poa supersect. Homalopoa. The results are consistent with the infrageneric classification in part; monophyletic sections are confirmed, it is proposed to reclassify species of sect. Acutifoliae, Dasypoa and Homalopoa s.l. and it is acknowledged that revision of the infrageneric taxonomy of the gynomonoecious species is needed.     

A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs

Cretaceous ichthyosaurs have typically been considered a small, homogeneous assemblage sharing a common Late Jurassic ancestor. Their low diversity and disparity have been interpreted as indicative of a decline leading to their Cenomanian extinction. We describe the first post-Triassic ichthyosaur from the Middle East, Malawania anachronus gen. et sp. nov. from the Early Cretaceous of Iraq, and re-evaluate the evolutionary history of parvipelvian ichthyosaurs via phylogenetic and cladogenesis rate analyses. Malawania represents a basal grade in thunnosaurian evolution that arose during a major Late Triassic radiation event and was previously thought to have gone extinct during the Early Jurassic. Its pectoral morphology appears surprisingly archaic, retaining a forefin architecture similar to that of its Early Jurassic relatives. After the initial latest Triassic radiation of early thunnosaurians, two subsequent large radiations produced lineages with Cretaceous representatives, but the radiation events themselves are pre-Cretaceous. Cretaceous ichthyosaurs therefore include distantly related lineages, with contrasting evolutionary histories, and appear more diverse and disparate than previously supposed.     

Anatomy and relationships of Pachyrhachis problematicus,a primitive snake with hindlimbs

The anatomy of Pachyrhachis problematicus, an elongate, limb-reduced squamate from the Upper Cretaceous of Israel, is described and evaluated in detail. Previously considered a snake-like ''lizard'' of uncertain affinities, it is here shown to be the most primitive snake, and the sister-group to all other snakes. Pachyrhachis exhibits numerous derived characters uniting it with modern snakes (scolecophidians and alethinophidians): e.g. mobile premaxilla-maxilla articulation, braincase enclosed by frontals and parietals, sagittal parietal crest, absence of tympanic recess, single postdentary bone, over 140 presacral vertebrae, and complete loss of shoulder girdle and forelimb. However, it is more primitive than all modern snakes in retaining some strikingly primitive (lizard-like) features: presence of a jugal, squamosal, normal sacral attachment, and well-developed hindlimb composed of femur, tibia, fibula, and tarsals. Pachyrhachis provides additional support for the hypothesis that snakes are most closely related to Cretaceous marine lizards (mosasauroids). Almost all of the derived characters proposed to unite snakes and mosasauroids are highly developed in Pachyrhachis: the mobile mandibular symphysis, intramandibular joint, long and recurved pterygoid teeth, quadrate suspended by the supratemporal, loosely united pelvic elements (ilium, ischium, and pubis), and separate astragalus and calcaneum.     

Cambrian stem-group annelids and a metameric origin of the annelid head

The oldest fossil annelids come from the Early Cambrian Sirius Passet and Guanshan biotas and Middle Cambrian Burgess Shale. While these are among the best preserved polychaete fossils, their relationship to living taxa is contentious, having been interpreted either as members of extant clades or as a grade outside the crown group. New morphological observations from five Cambrian species include the oldest polychaete with head appendages, a new specimen of Pygocirrus from Sirius Passet, and an undescribed form from the Burgess Shale. We propose that the palps of Canadia are on an anterior segment bearing neuropodia and that the head of Phragmochaeta is formed of a segment bearing biramous parapodia and chaetae. The unusual anatomy of these taxa suggests that the head is not differentiated into a prostomium and peristomium, that palps are derived from a modified parapodium and that the annelid head was originally a parapodium-bearing segment. Canadia, Phragmochaeta and the Marble Canyon annelid share the presence of protective notochaetae, interpreted as a primitive character state subsequently lost in Pygocirrus and Burgessochaeta, in which the head is clearly differentiated from the trunk.     

Convergent Evolution in Aquatic Tetrapods: Insights from an Exceptional Fossil Mosasaur

Mosasaurs (family Mosasauridae) are a diverse group of secondarily aquatic lizards that radiated into marine environments during the Late Cretaceous (98–65 million years ago). For the most part, they have been considered to be simple anguilliform swimmers – i.e., their propulsive force was generated by means of lateral undulations incorporating the greater part of the body – with unremarkable, dorsoventrally narrow tails and long, lizard-like bodies. Convergence with the specialized fusiform body shape and inferred carangiform locomotory style (in which only a portion of the posterior body participates in the thrust-producing flexure) of ichthyosaurs and metriorhynchid crocodyliform reptiles, along with cetaceans, has so far only been recognized in Plotosaurus, the most highly derived member of the Mosasauridae. Here we report on an exceptionally complete specimen (LACM 128319) of the moderately derived genus Platecarpus that preserves soft tissues and anatomical details (e.g., large portions of integument, a partial body outline, putative skin color markings, a downturned tail, branching bronchial tubes, and probable visceral traces) to an extent that has never been seen previously in any mosasaur. Our study demonstrates that a streamlined body plan and crescent-shaped caudal fin were already well established in Platecarpus, a taxon that preceded Plotosaurus by 20 million years. These new data expand our understanding of convergent evolution among marine reptiles, and provide insights into their evolution''s tempo and mode.     

Extreme convergence in stick insect evolution: phylogenetic placement of the Lord Howe Island tree lobster

The ‘tree lobsters’ are an enigmatic group of robust, ground-dwelling stick insects (order Phasmatodea) from the subfamily Eurycanthinae, distributed in New Guinea, New Caledonia and associated islands. Its most famous member is the Lord Howe Island stick insect Dryococelus australis (Montrouzier), which was believed to have become extinct but was rediscovered in 2001 and is considered to be one of the rarest insects in the world. To resolve the evolutionary position of Dryococelus, we constructed a phylogeny from approximately 2.4 kb of mitochondrial and nuclear sequence data from representatives of all major phasmatodean lineages. Our data placed Dryococelus and the New Caledonian tree lobsters outside the New Guinean Eurycanthinae as members of an unrelated Australasian stick insect clade, the Lanceocercata. These results suggest a convergent origin of the ‘tree lobster’ body form. Our reanalysis of tree lobster characters provides additional support for our hypothesis of convergent evolution. We conclude that the phenotypic traits leading to the traditional classification are convergent adaptations to ground-living behaviour. Our molecular dating analyses indicate an ancient divergence (more than 22 Myr ago) between Dryococelus and its Australian relatives. Hence, Dryococelus represents a long-standing separate evolutionary lineage within the stick insects and must be regarded as a key taxon to protect with respect to phasmatodean diversity.     

Sclerotic Rings in Mosasaurs (Squamata: Mosasauridae): Structures and Taxonomic Diversity

Mosasaurs (Squamata: Mosasauridae) were a highly diverse, globally distributed group of aquatic lizards in the Late Cretaceous (98–66 million years ago) that exhibited a high degree of adaptation to life in water. To date, despite their rich fossil record, the anatomy of complete mosasaur sclerotic rings, embedded in the sclera of the eyeball, has not been thoroughly investigated. We here describe and compare sclerotic rings of four mosasaur genera, Tylosaurus, Platecarpus, Clidastes, and Mosasaurus, for the first time. Two specimens of Tylosaurus and Platecarpus share an exact scleral ossicle arrangement, excepting the missing portion in the specimen of Platecarpus. Furthermore, the exact arrangement and the total count of 14 ossicles per ring are shared between Tylosaurus and numerous living terrestrial lizard taxa, pertaining to both Iguania and Scleroglossa. In contrast, two species of Mosasaurus share the identical count of 12 ossicles and the arrangement with each other, while no living lizard taxa share exactly the same arrangement. Such a mosaic distribution of these traits both among squamates globally and among obligatorily aquatic mosasaurs specifically suggests that neither the ossicle count nor their arrangement played major roles in the aquatic adaptation in mosasaur eyes. All the mosasaur sclerotic rings examined consistently exhibit aperture eccentricity and the scleral ossicles with gently convex outer side. Hitherto unknown to any squamate taxa, one specimen of Platecarpus unexpectedly shows a raised, concentric band of roughened surface on the inner surface of the sclerotic ring. It is possible that one or both of these latter features may have related to adaptation towards aquatic vision in mosasaurs, but further quantitative study of extant reptilian clades containing both terrestrial and aquatic taxa is critical and necessary in order to understand possible adaptive significances of such osteological features.     

First skull of Antillothrix bernensis,an extinct relict monkey from the Dominican Republic

The nearly pristine remains of Antillothrix bernensis, a capuchin-sized (Cebus) extinct platyrrhine from the Dominican Republic, have been found submerged in an underwater cave. This represents the first specimen of an extinct Caribbean primate with diagnostic craniodental and skeletal parts in association, only the second example of a skull from the region, and one of the most complete specimens of a fossil platyrrhine cranium yet discovered. Cranially, it closely resembles living cebines but is more conservative. Dentally, it is less bunodont and more primitive than Cebus, with crowns resembling Saimiri (squirrel monkeys) and one of the oldest definitive cebines, the late Early Miocene Killikaike blakei from Argentina. The tricuspid second molar also resembles the enigmatic marmosets and tamarins, whose origins continue to present a major gap in knowledge of primate evolution. While the femur is oddly short and stout, the ulna, though more robust, compares well with Cebus. As a member of the cebid clade, Antillothrix demonstrates that insular Caribbean monkeys are not monophyletically related and may not be the product of a single colonizing event. Antillothrix bernensis is an intriguing mosaic whose primitive characters are consistent with an early origin, possibly antedating the assembly of the modern primate fauna in greater Amazonia during the La Venta horizon. While most Greater Antillean primate specimens are quite young geologically, this vanished radiation, known from Cuba (Paralouatta) and Jamaica (Xenothrix) as well as Hispaniola, appears to be composed of long-lived lineages like several other mainland clades.     

A New Notosuchian from the Late Cretaceous of Brazil and the Phylogeny of Advanced Notosuchians

A new notosuchian crocodyliform from the Late Cretaceous Bauru Group found in the southeastern State of São Paulo (Brazil) is described here. The new taxon, Caipirasuchus stenognathus, is referred as a new species of the recently erected genus Caipirasuchus within the clade Sphagesauridae based on a phylogenetic analysis of basal mesoeucrocodylians. Caipirasuchus stenognathus is represented by an almost complete skull and lower jaw that has autapomorphic characters that distinguish it from other species of Sphagesauridae. These autapomorphies include: maxilla forming part of the orbital margin (absence of lacrimal-jugal contact), nasal with smooth depressions on the posterior region close to the contact with the maxilla and lacrimal, postorbital with posterior palpebral facet that extends posteriorly underneath the ear-flap groove, and a distinct anterior process of the medial flange of the retroarticular process. Additionally, the new taxon lacks autapomorphic features described in other sphagesaurids. The phylogenetic analysis results in a monophyletic genus Caipirasuchus, that is the sister group of a clade fomed by Sphagesaurus huenei, Caryonosuchus pricei, and Armadillosuchus arrudai. Sphagesaurids also include a basal clade formed by Adamantinasuchus navae and Yacarerani boliviensis. Other notosuchian taxa, such as Mariliasuchus amarali, Labidiosuchus amicum, Notosuchus terrestris, and Morrinhosuchus luziae are successive sister taxa of Sphagesauridae, forming a clade of advanced notosuchians that are restricted to the Late Cretaceous of South America. These results contrast with most previous phylogenetic hypotheses of the group that depicted some members of Sphagesauridae as more closely related to baurusuchids, or found Asian (e.g., Chimaerasuchus) or African (Malawisuchus, Pakasuchus) forms nested within advanced notosuchians that are, according to our analysis, endemic of the Late Cretaceous of South America.     

Angiosperm pollen grains of San Just site (Escucha Formation) from the Albian of the Iberian Range (north-eastern Spain)

The most complete angiospermous palynological assemblage for the middle-earliest late Albian of north-eastern Spain is reported from the Middle Member of the Escucha Formation at the San Just site. Nine palynological samples yielded mainly monosulcates (Clavatipollenites, Dichastopollenites, Hammenia, Jusinghipollis, Liliacidites, Pennipollis including proposed new species P. escuchensis, Retimonocolpites, and Transitoripollis) and a few tricolpates (Phimopollenites, Rousea, Striatopollis, and Tricolpites). These assemblages, mainly composed of angiosperm pollen grains, suggest that significantly diversified flowering plants flourished both in brackish and freshwater swamp plain environments represented by the Escucha Formation in Albian times, although they were less diverse than angiosperms in the Late Cretaceous. The palynological assemblages share greatest similarities with those of other southern Laurasian localities, although there are some taxa more typical of palynofloras of northern Gondwana.     

The paleoenvironments of azhdarchid pterosaurs localities in the Late Cretaceous of Kazakhstan

Five pterosaur localities are currently known from the Late Cretaceous in the northeastern Aral Sea region of Kazakhstan. Of these, one is Turonian-Coniacian in age, the Zhirkindek Formation (Tyulkili), and four are Santonian in age, all from the early Campanian Bostobe Formation (Baibishe, Akkurgan, Buroinak, and Shakh Shakh). All so far collected and identifiable Late Cretaceous pterosaur bones from Kazakhstan likely belong to Azhdarchidae: Azhdarcho sp. (Tyulkili); Aralazhdarcho bostobensis (Shakh Shakh); and Samrukia nessovi (Akkurgan). These latter two taxa, both from the Bostobe Formation might be synonyms. Azhdarcho sp. from the Zhirkindek Formation lived in a tropical-to-subtropical relatively humid climate on the shore of an estuarine basin connected to the Turgai Sea. Known fossils were collected in association with brackish-water bivalves and so the overall paleoenvironment of this pterosaur was likely an estuarine marsh as indicated by the dominance of conifers and low relative counts of ferns and angiosperms. Aralazhdarcho bostobensis, from the Bostobe Formation, lived on a coastal fluvial plain along the Turgai Sea. This paleoenvironment was either floodplain (Akkurgan, Buroinak, and Shakh Shakh) or estuarine (Baibishe). In the Santonian – early Campanian, shallow waters near this coastal plain were sites for the intensive accumulation of phosphates under upwelling conditions caused by strong winds from the ancient Asian landmass. These winds also caused significant aridization of the climate during this time. We speculate that pterosaurs may have been attracted to this area by the abundant resources in the bio-productive estuaries and nearshore upwelling waters.