A new, nearly complete stem turtle from the Jurassic of South America with implications for turtle evolution

Turtles have been known since the Upper Triassic (210Myr old); however, fossils recording the first steps of turtle evolution are scarce and often fragmentary. As a consequence, one of the main questions is whether living turtles (Testudines) originated during the Late Triassic (210Myr old) or during the Middle to Late Jurassic (ca 160Myr old). The discovery of the new fossil turtle, Condorchelys antiqua gen. et sp. nov. from the Middle to Upper Jurassic (ca 160-146Myr old) of South America (Patagonia, Argentina), presented here sheds new light on early turtle evolution. An updated cladistic analysis of turtles shows that C. antiqua and other fossil turtles are not crown turtles, but stem turtles. This cladistic analysis also shows that stem turtles were more diverse than previously thought, and that until the Middle to Upper Jurassic there were turtles without the modern jaw closure mechanism.     

A new nanhsiungchelyid turtle from the Late Cretaceous of Neixiang,Henan Province,China

A new genus and species of Nanhsiungchelyidae (Testudines: Cryptodira), Yuchelys nanyangensis n. g. n. sp., are described on the basis of a partial skeleton from the Late Cretaceous Gaogou Formation of Neixiang, Nanyang Basin, Henan Province. It is the first nanhsiungchelyid record in the east-central part of China and fills the geographical gap between the western (Uzbekistan, Mongolia, Northern and Southern China) and eastern (Japan) distribution areas of the family Nanhsiungchelyidae.     

Paleofloristic and paleoenvironmental implications of a Late Jurassic palynoflora from the Tochikubo Formation,North-East Japan

The first Jurassic palynofloral assemblage from Japan is reported from the Oxfordian Tochikubo Formation of the Soma-Nakamura Group, Fukushima Prefecture. Palynomorphs are quite diversified, with 32 genera and 41 species of lycopod and fern spores, gymnosperm pollen and freshwater algae. The composition of the assemblage is consistent with the Oxfordian age of this nonmarine formation and confirms a fluvio-lacustrine paleoenvironment, further detailing the paleovegetation reconstructed from macrofloral remains. Previous palynofloral reports of eastern Asia are reviewed and indicate that northeast Japan shares most similarities with coeval paleofloras of southeastern Russia. Consequences for paleofloristic provincialism and climates of the region during the Late Jurassic are discussed.     

A new cryptoclidid Plesiosaur from the Oxfordian (Late Jurassic) of Cuba

Cryptocleidus ? cuervoi caroliDE LA TORRE & ROJAS, 1949, a partial skull with associated mandible and atlas-axis from the Oxfordian of Cuba, has been completely prepared for the first time. It is here redescribed and assigned to a new cryptoclidid genus, Vinialesaurus, for which the species caroli is retained. Vinialesaurus is mainly diagnosed by palatal characters such as double internal nares, an anteriorly rounded vomer and a lack of anterior interpterygoid vacuities. It shares with other cryptoclidids large orbits and external nares, a small vertical jugal and a reduced tooth ornamentation. The occurrence of Vinialesaurus caroli in association with pliosauroids, ophthalmosaurian ichthyosaurs, metriorhynchid crocodilians and pleurodiran marine turtles, strongly suggests that a marine seaway was present in the Caribbean during the Oxfordian, connecting the western Tethys with the Oriental Pacific.     

A new Xinjiangchelyid turtle (Testudines, Eucryptodira) from the Jurassic Qigu Formation of the southern Junggar Basin, Xinjiang, north-west China

A new eucryptodiran turtle, Xinjiangchelys qiguensis sp. nov. from the Upper Jurassic (Oxfordian — ?Kimmeridgian) Qigu Formation of the southern Junggar Basin (north‐west China) is described. The type material consists of a partial skeleton, including the complete carapace, plastron, nearly all cervical vertebrae, both scapulae, the pelvis and one ulna. It is clearly identifiable as a basal eucryptodire since it lacks the mesoplastron. It is distinguished from other species of Xinjiangchelys by several autapomorphies of the carapace and plastron, such as the first and fifth vertebrals extending on the peripherals, the plastron with three pairs of gulars, and an intergular which does not contact the hyoplastron. In the postcranium, the scapula with a long acromial and a small scapular process, the pelvis with a short ilial shaft and the elongated cervical vertebrae are characteristic. A new phylogenetic analysis of the in‐group phylogeny of the Xinjiangchelyidae is proposed and discussed, resulting in a new classification of the family. Xinjiangchelys (Toxocheloides) narynensis is regarded as a nomen dubium. Shartegemys is referred to Xinjiangchelys, whereas the holotypes of ‘Plesiochelys’chungkingensis and ‘P’. latimarginalis are excluded from the genus Xinjiangchelys but included in the Xinjiangchelyidae.     

A revision of xinjiangchelyid turtles from the Late Jurassic of Sichuan Basin,China

The type specimens of the Jurassic turtles from the Sichuan Basin (China) previously referred to plesiochelyid Plesiochelys are revised, which confirm that they belong to the family Xinjiangchelyidae. The study of a large number of additional shell material shows, as the dominant group, the Xinjiangchelyidae were greatly diversified in the Late Jurassic of the Sichuan Basin. By the absence of mesoplastron and other shell characters, Chengyuchelys baenoides is moved to Xinjiangchelyidae and considered as a valid taxon. Of the xinjiangchelyids from the Late Jurassic of Sichuan Basin, four genera are recognized: Chengyuchelys, Tienfuchelys, Yanduchelys and cf. Protoxinjiangchelys. The phylogenetic analysis results in that Chengyuchelys includes C. baenoides, C. latimarginalis and C. radiplicatus, and Tienfuchelys consists of T. tzuyangensis, T. chungkingensis and T. zigongensis. All xinjiangchelyids from the Sichuan Basin are more primitive than Xinjiangchelys in the plastron sutured to the carapace.     

Coral biostromes of the Middle Jurassic from the Subbetic (Betic Cordillera,southern Spain): facies,coral taxonomy,taphonomy, and palaeoecology

Coral biostromes from the Camarena Formation (External Subbetic, Betic Cordillera) are reviewed under palaeoecologic, taphonomic, and palaeontologic aspects. The biostromes are dominated by phaceloid forms and are characterized by a typical shallow-marine microencruster assemblage with photophilic microencrusters and scarce microbial crusts. The abundance of stylinid corals and light-dependant microencrusters suggests oligotrophic conditions. Coral colonies were located among oolitic shoals that were unfavorable for coral growth. The corals were developed in phases without oolitic production alternating with phases of oolitic production, forming metric-scale sequences. A relative sea-level fall would have reduced the ooidal production and led to the deposition of thin layers of micritic facies in intertidal areas. The cementation and hardening of the bottom resulted in a hardground that was colonized by corals after a subsequent relative sea-level rise. The progressive increase of the energetic conditions induced an increasing production of ooids and the migration of oolitic shoals, which covered and finished the coral biostromes. Repetition of this process gave rise to sequences reflecting small pulses of oscillations in the relative sea level.     

Conservation implications of complex population structure: lessons from the loggerhead turtle (Caretta caretta)

Complex population structure can result from either sex-biased gene flow or population overlap during migrations. Loggerhead turtles (Caretta caretta) have both traits, providing an instructive case history for wildlife management. Based on surveys of maternally inherited mtDNA, pelagic post-hatchlings show no population structure across the northern Atlantic (phi(ST) < 0.001, P = 0.919), subadults in coastal habitat show low structure among locations (phi(ST) = 0.01, P < 0.005), and nesting colonies along the southeastern coast of the United States have strong structure (phi(ST) = 0.42, P < 0.001). Thus the level of population structure increases through progressive life history stages. In contrast, a survey of biparentally inherited microsatellite DNA shows no significant population structure: R(ST) < 0.001; F(ST) = 0.002 (P > 0.05) across the same nesting colonies. These results indicate that loggerhead females home faithfully to their natal nesting colony, but males provide an avenue of gene flow between regional nesting colonies, probably via opportunistic mating in migratory corridors. As a result, all breeding populations in the southeastern United States have similar levels of microsatellite diversity (H(E) = 0.70-0.89), whereas mtDNA haplotype diversity varies dramatically (h = 0.00-0.66). Under a conventional interpretation of the nuclear DNA data, the entire southeastern United States would be regarded as a single management unit, yet the mtDNA data indicate multiple isolated populations. This complex population structure mandates a different management strategy at each life stage. Perturbations to pelagic juveniles will have a diffuse impact on Atlantic nesting colonies, mortality of subadults will have a more focused impact on nearby breeding populations, and disturbances to adults will have pinpoint impact on corresponding breeding populations. These findings demonstrate that surveys of multiple life stages are desirable to resolve management units in migratory marine species.     

Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy

Distinct faunal aggregates are described from the Middle Jurassic (uppermost Bajocian/lowermost Bathonian and Middle Bathonian) clay deposits of Częstochowa area, south-central Poland. These aggregates are composed of molluscs (scaphopods, gastropods, bivalves, ammonites and belemnites), articulate brachiopods and echinoderms (asteroids, crinoids and echinoids). A large percentage of the fossils, especially bivalves, are fragmented, but some fossils are complete. Although most of the fossils are crushed and fragmented, they are still identifiable to at least the genus level. Thorough statistical analysis of taphonomic features indicates that the preservation of asteroid marginal plates is distinct from the ossicles derived from the host clays. The high frequency of bite marks and the good state of preservation suggest that the accumulations are the products of predation activities and most probably are the effect of regurgitation. Taking into account the rich and diverse fauna, the predator was a bottom-feeding generalist. Possible predators include palaeospinacid sharks, a tooth of which was collected from the same bedding surface, but not associated with regurgitated remains. Although the bite marks on the asteroid ossicles point to sharks as potential producers of regurgitates, other vertebrates, like durophagous pycnodontiform fish, cannot be excluded.     

New internally structured decapod microcoprolites from Germany (Late Triassic/Early Miocene), Southern Spain (Early/Middle Jurassic) and Portugal (Late Jurassic): Taxonomy, palaeoecology and evolutionary implications

Fossil faecal pellets with internal structures are described from non-marine environments for the first time. In the Late Triassic Lehrberg Beds (Middle Keuper, Carnian) of Germany, the oldest stratigraphic record of the form genusHelicerina Brönnimann &Masse,Helicerina keuperina Seegis n. sp., occurs.Favreina kalankyra Ebli from the Norian Hauptdolomit Formation is also attributed to this genus. Another new speciesof Helicerina, H. ruttei Schweigert n. sp., is described from the Early Miocene calcareous freshwater tufas of Southern Germany. It is attributed to the brachyuran freshwater crabProballaya quenstedti (Zittel) which occurs abundantly at the same locality. Hence, the coprolite genusHelicerina is the product of brachyuran decapods or their Triassic ancestors, and not of anomuran decapods as thought before. From younger Early to Middle Jurassic pelagic ferrugineous limestones of the Subbetic area (Southern Spain), the new speciesFavreina belandoi Fels n. sp. is described besidesHelicerina siciliana Senowbari-Daryan, Schäfer &Catalano, and the better known speciesPalaxius salataensis Brönnimann, Cros &Zaninetti. The number of canals withinFavreina belandoi varies but is clearly correlated with different ontogenetic stages of the same producer. From the Late Jurassic of Portugal, the new speciesPetalina hexalunulata Leinfelder n. gen. n. sp. is reported. It co-occurs withFavreina prusensis (Paréjas). The new genusPetalina is distinct fromFavreina by its hemispherical cross sections of the canals. In Portugal, both species are restricted to intrajurassic marine karstic fissures of the reefal Ota Limestone.     

A new phylogenetic hypothesis of turtles with implications for the timing and number of evolutionary transitions to marine lifestyles in the group

Evolutionary transitions to marine habitats occurred frequently among Mesozoic reptiles. Only one such clade survives to the present: sea turtles (Chelonioidea). Other marine turtles originated during the Mesozoic, but uncertain affinities of key fossils have obscured the number of transitions to marine life, and the timing of the origin of marine adaptation in chelonioids. Phylogenetic studies support either a highly‐inclusive chelonioid total‐group including fossil marine clades from the Jurassic and Cretaceous (e.g. protostegids, thalassochelydians, sandownids) or a less inclusive chelonioid total‐group excluding those clades. Under this paradigm, these clades belong outside Cryptodira, and represent at least one additional evolutionary transition to marine life in turtles. We present a new phylogenetic hypothesis informed by high resolution computed tomographic data of living and fossil taxa. Besides a well‐supported Chelonioidea, which includes protostegids, we recover a previously unknown clade of stem‐group turtles, Angolachelonia, which includes the Late Jurassic thalassochelydians, and the Cretaceous–Palaeogene sandownids. Accounting for the Triassic Odontochelys, our results indicate three independent evolutionary transitions to marine life in non‐pleurodiran turtles (plus an additional two‐three in pleurodires). Among all independent origins of marine habits, a pelagic ecology only evolved once, among chelonioids. All turtle groups that independently invaded marine habitats in the Jurassic–Cretaceous (chelonioids, angolachelonians, bothremydid pleurodires) survived the Cretaceous–Palaeogene mass extinction event. This highlights extensive survival of marine turtles compared to other marine reptiles. Furthermore, deeply‐nested clades such as chelonioids are found by the middle Early Cretaceous, suggesting a rapid diversification of crown‐group turtles during the Early Cretaceous.     

Late Mississippian vertebrate palaeoecology and taphonomy,Buffalo Wallow Formation,western Kentucky,USA

Aquatic vertebrates are reported from several facies of the Late Mississippian (Chesterian/Elviran/Serpukhovian) Buffalo Wallow Formation in western Kentucky, USA. Rhizodont bones and the partially articulated skeleton of a large anthracosaur (proterogyrinid) were found in rocks interpreted as a fluvial‐estuarine palaeochannel. Smaller, disarticulated tetrapod remains (anthracosaurs, whatcheeriids) were found in a weathered siltstone in an apparent channel margin‐sand flat facies. A putative oxbow‐abandoned channel facies contains skeletal elements of rhizodonts (cf. Rhizodus), colosteiids, Gyracanthus, Ageleodus, Cynopodius, xenacanthoids and palaeonisiciforms. Near the top of the channel fill, lungfish (cf. Tranodis) are found in carbonate‐rich nodules, which appear to be aestivation burrows. A presumed lacustrine facies contained a near‐complete colosteid. Thinning section, palaeosols, pedogenically‐altered carbonates and missing strata suggest tectonic and climatic overprints upon these depositional sequences. Multiple, incised channels in a low‐accommodation setting are interpreted to have provided local faunal traps for aquatic vertebrates. Late Mississippian palaeoclimate changes may have caused water table fluctuations, which might have aided in trapping and preserving aquatic vertebrates.     

Morphology and phylogenetic interpretation of a new Cambrian edrioasteroid (Echinodermata) from Spain

Abstract: The edrioasteroid, Aragocystites belli gen. nov. sp. nov. from the middle Cambrian Murero Formation of Spain, is described based on a small number of very well‐preserved specimens. Important anatomical characteristics include star‐shaped and pseudoclavate theca, rare or absent epispires, well‐developed interradially positioned oral plates and several unorganized cover plates associated with each widely exposed flooring plate. A phylogenetic analysis including several Cambro–Ordovician species shows it is more derived than Stromatocystites and Totiglobus but is a sister group to a clade comprising Cambraster and Edriodiscus. Ontogenetic observations based on juveniles of 5 mm in diameter suggest that this species changed thecal shape markedly during growth. A. belli gen. nov. sp. nov. probably lived in quiet environments where it attached directly to the sea floor on stabilized substrates.     

Sciuridae (Rodentia,Mammalia) from the Late Miocene of Venta del Moro (Cabriel Basin,Spain): biostratigraphical,phylogenetic and palaeoecological inferences

Abstract

Fossil squirrels are relatively abundant in Aragonian and Ramblian assemblages, but very scarce in the Late Turolian and Ruscinian of the Iberian Peninsula. Until now, the locality of Venta del Moro has yielded over 4500 micromammal dental remains, but just nine of them are sciurids. In this assemblage, we have identified the taxa Pliopetaurista pliocaenica, Heteroxerus cf. mariatheresae and Atlantoxerus cf. margaritae. This is the first co-occurrence of the genera Pliopetaurista, Heteroxerus and Atlantoxerus in the same locality, being also one of the oldest records of P. pliocaenica, the oldest record of a form that could be related to A. margaritae, and the youngest record of the genus Heteroxerus. In addition, we hypothesize about the phylogeny of Pliopetaurista in the light of recent discoveries. Finally, the presence of the genera Heteroxerus and Atlantoxerus suggests an open environment and relatively dry environmental conditions, while Pliopetaurista is purportedly a flying squirrel, dweller of forested habitats and wet climates. Based on this, we infer an open environment of grassland type, bordered by forested habitats, agreeing with the conditions supposed for Venta del Moro according to its macromammal and floral assemblages.     

A compsemydid turtle from the Upper Cretaceous of Var,southern France

Calissounemys matheroni gen. et sp. nov. (Testudines) is described on the basis of a skull and shell elements from the Upper Cretaceous of Var, southern France. This new taxon is assigned to the family Compsemydidae and characterized by a thick-boned, robust skull, a shallow temporal emargination, a crista supraoccipitalis not extending beyond the posterior edge of the skull roof, large nasals meeting along the midline for their full length; frontals retracted from the orbital margin, absence of a cheek emargination, a large jugal forming a substantial part of the orbital margin, absence of a secondary palate and an uneven upper triturating surface; and the shell with vertebral 1 clearly wider than vertebrals 2-3, with the lateral margins strongly divergent towards the anterior border and wider than long vertebrals 2-3. This find increases the diversity of the Late Cretaceous turtle fauna from southern France, and fills a stratigraphical gap in the fossil record of Compsemydidae between the Early Cretaceous and the Paleocene in Europe.     

A contribution to tropical rain forest taphonomy: retrieval and documentation of chimpanzee remains from Kibale Forest, Uganda

The absence of Pan and Gorilla fossils from Africa has led some to suggest that African rain forests are not conducive to bone preservation. The absence of fossils is unfortunate as it hampers phylogenetic and socioecological interpretations on the divergence of the earliest hominids. For the most part, taphonomic studies have been restricted to cave and open country contexts. With this in mind, we have initiated a taphonomic project in a tropical rain forest, the Kibale Forest of western Uganda. In the course of bone gathering activities over the past 4 years, we have documented the retrieval of skeletal remains representing nine chimpanzees (Pan troglodytes schweinfurthii). Crania and mandibles are most commonly recovered, followed by elements of the axial skeleton, and finally, appendages. Vertical segregation of small compact bones can occur in areas with a soft substrate. Scavenging activity suggests the role of suids, but this has not been proven. Geochemical tests suggest that Kibale soils are neutral and may be conducive to bone preservation. Our independent preliminary data from bone weathering/survival experiments indicate that bones appear undamaged after several years and are able to accumulate on the forest floor. These results contrast with popular assumptions on the potential of African rain forests as potential fossil reservoirs.     

Taphonomy of fossil macro-invertebrate assemblages as a tool for ecostratigraphic interpretation in Upper Jurassic shelf deposits (Prebetic Zone, southern Spain)

Composition and taphonomy of macro-invertebrate fossil assemblages, together with facies analysis, have been approached in order to interpret shifting paleoenvironmental conditions in the External Prebetic (S-SE Spain) during the early Late Jurassic (Middle Oxfordian). In oolitic and spongiolitic limestones, the size of fossil remains, mode of preservation, within-bed position, corrasion, fragmentation, epibiont and biogenic encrustation, disarticulation and uncoupling, allow recognition of two taphofacies, respectively. Identified ecostratigraphic events and trends accord with rapid flooding under high-energy conditions related to ecospace enlargement for cephalopods and then the persistence of lower energy, long-lasting exposure of skeletals and higher sedimentary rates. The paleoenvironmental interpretation is consistent with neritic environments shifting from shallow carbonate to hemipelagic sedimentation and enlarging of shelf ecospace for marine invertebrates.     

Late Maastrichtian small-sized herpetofauna from Valencia province,eastern Spain

A Late Maastrichtian microvertebrate assemblage which includes amphibian remains was recovered from continental deposits of the palaeontological site of La Solana, Valencia Province, Spain. This site is composed of variegated mudstones, pedogenically modified, interbedded with fluvial sand bodies and freshwater limestones lenses, and has also yielded plant debris, freshwater and oligohaline invertebrates, abundant fish remains (isolated bones and scales), turtle plates and archosaur bones. This fossil assemblage, dominated by aquatic forms, also includes semiaquatic and terrestrial elements, and may be interpreted as the palaeofauna of a wetland environment with terrestrial environs. The new material described here consists of fragmentary remains of an indeterminate albanerpetontid, a salamandrid and two anuran taxa (an alytid and a palaeobatrachid). The amphibians from La Solana are typical Laurasiatic taxa. This faunal association shows broad similarities to other coeval faunas of the Iberian Peninsula and contrasts with the Upper Campanian–Lower Maastrichtian sites where Gondwanan elements are frequent.     

Benthic foraminiferal morphogroups of mid to outer shelf environments of the Late Jurassic (Prebetic Zone, southern Spain): Characterization of biofacies and environmental significance

Benthic foraminiferal assemblages belonging to a mixed carbonate–siliciclastic shelf succession of the Oxfordian (Upper Jurassic) in the Prebetic, southern Spain, were analyzed. The faunal data, obtained on thin sections of strongly lithified sediments, allowed the detailed differentiation of a foraminiferal morphogroups system to be applied to the interpretation of paleoenvironmental conditions. Eleven morphogroups were differentiated according to shell composition, general morphology, number of chambers and mode of coiling. Paleoenvironmental analysis combines these features with inferred life-style (epifaunal, shallow infaunal and deep infaunal) and feeding strategy (suspension-feeder, deposit-feeder, herbivores, etc.) of the foraminifera. Analogies and differences regarding previously established morphogroup systems could be outlined accordingly.

In the Prebetic Oxfordian, distribution trends of foraminiferal morphogroups allow for a comparison of mid to outer shelf environments occupying the South-Iberian paleomargin, indicating that nutrient supply was the most important factor controlling morphogroup development. The mid shelf settings are dominated by epifaunal active herbivores–phytodetritivores, in shallow to deep substrate position of the redox boundary. The outer shelf is dominated by agglutinated and infaunal detritivore-bacterial scavenger morphogroups revealing a deep substrate position of the redox boundary. Particularly favorable trophic conditions in mid shelf environments are related with higher phytodetritus input from more proximal and shallower shelf areas.