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 Serravallian (Middle Miocene) shark fauna from Southeastern Spain and its palaeoenvironment significance

Abstract

The study of a new Serravallian (Middle Miocene) locality from the Southeastern Spain has yielded a shark assemblage characterized by microremains of at least seven taxa (Deania calcea, ?Isistius triangulus, ?Squaliolus cf. S. schaubi, ?Paraetmopterus sp., Pristiophorus sp., Scyliorhinus sp. and a cf. Squaliformes indet) of three different orders (Squaliformes, Pristiophoriformes and Carcharhiniformes). In addition, associated macroremains have also been found, including teeth of ?Cosmopolitodus hastalis, Isurus sp., Hemipristis serra, Odontaspis sp., Carcharhinus spp. and ?Otodus (Megaselachus) megalodon. The assemblage contains taxa with disparate environmental preferences including not only neritic and epipelagic sharks but also an important number of meso and bathypelagic representatives. The migration of deep water taxa to shallower waters through submarine canyons/coastal upwelling is proposed as the most plausible cause for explaining the origin of such assemblage. Interestingly, the composition of the deep-water taxa here reported contrast with the chondrichthyans assemblages from the Pliocene and extant Mediterranean communities. This entails a complex biogeographic history, where the Messinian salinity crisis strongly affected the posterior evolution of the Mediterranean ecosystems but some other factors, such us the existence of anoxic events during the Quaternary, could have also played an important role.     

Middle Miocene Cephalotaxus (Taxaceae) from Yunnan,Southwest China,and its implications to taxonomy and evolution of the genus

The infrageneric taxonomy of the genus Cephalotaxus Siebold et Zuccarini ex Endlicher (Taxaceae) is controversial as morphological characters have been doubted to change coherently among species. Although the epidermal features are considered as important characters in the taxonomy of conifers, they have not been systematically studied in this genus. Neogene foliage shoots with well-preserved epidermal structure not only can provide systematic links to the extant species, but also have implications for the taxonomical definition of the extant species by their epidermal features. In this paper, the morphologies of leafy shoots combining epidermal structure of a Neogene Cephalotaxus, which was collected from the middle Miocene of Southwest China, and ten extant species (variants) were studied. Morphology as well as epidermal structures was compared among species (variants) in Cephalotaxus. Based on the comparisons, the taxonomical definition of extant species in genus Cephalotaxus are discussed. Our results indicate that leaf morphological characters in combination with the features of epidermal structure can distinguish species in genus Cephalotaxus. The fossil foliage shoots from the middle Miocene of Southwest China were assigned to a new species, C. maguanensis n. sp., which shows close affinities to the extant C. hainanensis. On the basis of leaf and epidermal information, we suggest there are seven extant species in Cephalotaxus. With global cooling and the uplift of the Qinghai-Tibet Plateau since the middle Miocene, the cooling climate induced the differentiation and southward expansions of C. maguanensis, and its extant offspring, C. hainanensis, is distributed to Hainan Island and the Indo-Chinese peninsula.     

Facies and palaeoecology of Upper Jurassic (Middle Oxfordian) coral reefs in England

Summary This study documents the facies and fauna of Late Jurassic (Middle Oxfordian) coral reefs in England. Sedimentological and palaeoecological analysis of these reefs distinguishes three generic reef types: (1) small reef patches and thickets associated with siliciclastic deposits; (2) small reef patches and thickets associated with siliciclastic-free bioclastic grainstones and packstones; and (3) biostromal units associated with deep water facies. The depositional environments of these reef types are discussed. Two coral assemblages are identified: (1) the microsolenid assemblage; and (2) theThamnasteria, Isastraea, Fungiastraea andThecosmilia assemblage (Thamnasteria assemblage). TheThamnasteria assemblage developed in all shallow water environments in the study area, regardless of local environmental conditions. The fauna is very eurytopic,r-selected and can tolerate significant environmental fluctuations on short temporal scales (sub-seasonal). The main control on the development of the microsolenid assemblage was low light intensity, low background sedimentation rates and low hydrodynamic energy levels.     

Benthic foraminiferal assemblages of the Middle and Upper Jurassic sediments from the northeastern Alborz and western Koppeh Dagh,Iran: Systematic palaeontology and palaeoecology

This research is focused on a poorly studied Jurassic sequence in the eastern part of Alborz and Western Koppeh Dagh basins. The foraminifera are reported from Callovian (Middle Jurassic) to Tithonian (Upper Jurassic) sediments of the Farsian and Chaman Bid formations. The benthic foraminiferal associations belong to the families Ammodiscidae, Epistominidae, Hauerinidae, Hormosinidae, Ichthyolariidae, Lagenidae, Nodosariidae, Nubeculariidae, Ophthalmidiidae, Polymorphinidae, Rhabdamminidae, Saccamminidae, Spirillinidae, Spirocyclinidae, Textulariidae, Trocholinidae, and Vaginulinidae. In the studied area, the foraminifera are dominated by the families Nodosariidae, Vaginulinidae and Epistominidae. Palaeoecological analysis of the foraminiferal assemblages from the Farsian Formation indicates that these sediments were deposited on the inner to mid shelf environment, with normal salinity (euhaline), and relatively well oxygenated waters, with eutrophic conditions in the Goznawwi area and oligotrophic conditions in the Chaman Bid area. The foraminiferal assemblages in this study are the first record from Jurassic sediments of the eastern part of the Alborz and Koppeh Dagh basins in northern Iran.     

New species of Hispanomys (Rodentia,Cricetodontinae) from the Upper Miocene of Batallones (Madrid,Spain)

Material of Hispanomys (Rodentia, Cricetodontinae) is described from various localities at Batallones (MN10) (Madrid, Spain). All of it belongs to a single species, which differs from the other known species of the genus and a new taxon, Hispanomys moralesi sp. nov. , is created for it. The samples from the various localities show differences interpreted as being the result of slight age disparities amongst the different sites. Although they were previously thought to be coeval, Batallones 10 is probably older than Batallones 1, which is possibly older than Batallones 3. Hispanomys moralesi sp. nov. is characterized by several morphological features such as the lack of cingula and mesolophs, the presence of well‐developed ectolophs, four‐ or five‐rooted M1, short or absent mesolophids, and reduced and simplified M3. Hispanomys moralesi sp. nov. is a relatively derived species, the evolutionary stage of which is comparable to those of other members of the genus from the Upper Vallesian. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 725–747.     

Sierra del Colmenar 1A,a new late Messinian (Late Miocene) locality in the Bajo Segura Basin (SE Spain): biostratigraphic and palaeoenvironmental implications

Abstract

In this paper, we have studied a new micromammal site from the Sierra del Colmenar section (Elche, SE Spain), named Sierra del Colmenar 1A (SCO-1A), representing the uppermost levels of Messinian age of the Bajo Segura Basin. The sedimentary context of this locality corresponds to a costal lagoon with marine influence. The fossil site has yielded remains of Apodemus aff. gorafensis, Paraethomys meini, Apocricetus alberti, Occitanomys alcalai, Ruscinomys sp., Eliomys cf. truci, Muscardinus sp., Parasorex ibericus, Prolagus michauxi and Soricidae indet. Based on this assemblage, we propose a Late Miocene age (MN13) for the locality SCO-1A, matching the previously inferred age for the stratigraphic unit in which the fossil site is situated. According to its faunal composition, we infer warm and humid climate conditions in the moment of formation of this locality, agreeing with the environmental conditions of the Messinian age in a parallic sedimentary context.     

Slope breccias colonized by bivalves and serpulids during the Middle Jurassic (Subbetic,SE Spain)

The Middle Jurassic of the Intermediate Domain, in the northern Subbetic (External zones of the Betic Cordillera, SE, Spain), reveals the transition from hemipelagic limestones (Baños Formation) to overlying shallow-water oolitic limestones (Jabalcuz Formation). Near La Guardia (Jaén), the basal part of the succession that records this transition comprises clast-supported calcareous breccias 24 m thick. Transverse sections reveal a fan shape close to a normal fault. The upper parts of some breccia beds were colonized by bivalve patches that locally evolve to bivalve-serpulid bioherms. The bioherms are massive, with low relief and a metre-scale lenticular shape. Two main parts have been differentiated: a bivalve bioherm and a serpulid bioherm. The bivalve bioherm consists of densely packed mytiloids in life position with secondary serpulids, where the matrix is a grainstone of bioclasts, peloids, and ooids. Laterally, there are accumulations of reworked bivalves. The serpulid bioherm is made up of serpulid aggregates in a bioclast wackestone. On an unstable slope with abundant breccias, bioherm development took place in favourable periods of low sedimentation rate. The pioneer colonisers of the sea-bottom were mytiloid bivalves in relatively high-energy waters. Shell comissures were aligned parallel to the dip of the slope as an adaptation to feeding from the plankton and seston inputs associated with palaeocurrents. The intercalation of reworked bivalve beds likely indicates brief periods of higher-energy storm events. The serpulid bloom in the upper part of the buildups was probably related to a stage of low-energy conditions (as evidenced by the delicacy of these structures), which is congruent with the wackestone texture. The relief provided by the bivalves favoured the development of the serpulid bioherm phase in an elevated and advantageous position for suspension-feeding. The dense packing in these bioherms offered clear advantages in this unstable palaeoenvironment: (1) dense populations are less susceptible than isolated individuals to overgrowth by other sessile organisms or predator attacks; and (2) the interconnection among individuals enhances stability and resistance against high-energy events, and increases the growth potential of future generations. The interruption of bioherm growth was probably related to the deposition of new breccia beds and reworked oolitic limestones.     

Palaeofloristic assemblages and palaeoecology of the Lower Oligocene Bembridge Marls, Hamstead Ledge, Isle of Wight

A sequence of nine, plant-macrofossil-bearing horizons has been recognized in the Lower Oligocene from the Isle of Wight, England. The plant fossils are contained in finely laminated, olive-brown to black silty-clays and clays. Some horizons exhibit rhythmic alternation of sediment couplets; others show evidence of bioturbation and rootlet penetration. These horizons are included within a sequence of finely laminated greenish grey silty-clays and clays, largely devoid of plant macrofossil material. The plant-bearing horizons contain abundant fruits, seeds, fern sporangia, Azolla megaspores, and rare leaves. The nearest living relatives of the fossils all inhabit an aquatic or marginal aquatic environment. The associated fossil fauna includes lake bottom and lake margin species. The palynoflora of the plant-bearing and immediately underlying horizons includes large numbers of planktonic algae and limited representatives of a forest vegetation. It shows a strong relationship between the occurrences of Sparganiaceaepollenites pollen and of Typha macrofossils. Palaeoecological interpretations of this plant-fossil-bearing sequence are discussed with particular reference to modern environments.
One new genus and three new species are described. Emendations are presented for four other species. Stratiotes leaf margin teeth are described.     

Middle and Upper Jurassic bivalve associations from the Iberian Range (Spain)

The four sections richest in bivalves from the Middle and Upper Jurassic of the Iberian Range (Spain) were selected for a quantitative palaeoecological analysis of the bivalve fraction of the macrobenthos. Five bivalve associations and two assemblages were recognized with the help of a Q-mode hierarchical cluster analysis (Ward method). The main environmental factors controlling bivalve associations are thought to be substrate, water energy and distribution of organic matter. The bivalves exhibit a distinct spatial and temporal distribution pattern within the Aragonian Branch of the Iberian Range. Four of the bivalve associations occur in the Upper Oxfordian (Sot de Chera Fm) and one association in the Lower Callovian (Chelva Fm). In the Sot de Chera and Loriguilla formations, the abundance of bivalves decreases from NW to SE i.e., from relatively close to the shore line towards the distal-most part of the carbonate platform. In the Chelva Fm, bivalves are abundant in the Ariño region, interpreted as a palaeogeographic high. The spatial distribution of bivalves might have been largely controlled by the availability of nutrients.     

Fossil frogs (Anura) from Shanwang (Middle Miocene; Shandong Province, China)

The early middle Miocene biota from the Shanwang locality in eastern China is one of the most diverse and well preserved in the world. This overall richness contrasts with its low diversity of anuran amphibians. The most abundant anurans are representatives of the family Ranidae, which remains the dominant group in the extant anuran fauna of China. Besides Rana basaltica Young, some other ranids can be recognised at the locality, and are clearly distinguishable from R. basaltica based on their larger size and different proportions. Ranids are also represented by a developmental series of tadpoles, ranging from approximately Nieuwkoop-Faber (NF) stage 43 to a metamorphosing tadpole of NF stage 57. This is the first known developmental series of fossil non-pipoid Anura outside the Pelobatidae. In addition to the ranids, Bufo shandongensis nov. sp. (replacement name for B. linquensis), represented by one of the earliest known articulated bufonid skeletons, is described and illustrated. Another large toad, Macropelobates linquensis (Yang) nov. comb., is re-described; it can be assigned to the Pelobatoidea, with a possible relationship to the Scaphiopodidae, and can be considered, based on the presence of a large spade, a tibiofibula shorter than the femur, and a short astragalus and calcaneus, to be the earliest known anuran burrower.     

Biochronology and paleoclimatic implications of Middle Eocene to Oligocene planktic foraminiferal faunas

Planktic foraminiferal assemblages have been analyzed quantitatively in six DSDP sites in the Atlantic (Site 363), Pacific (Sites 292, 77B, 277), and Indian Ocean (Sites 219, 253) in order to determine the nature of the faunal turnover during Middle Eocene to Oligocene time. Biostratigraphic ranges of taxa and abundance distributions of dominant species are presented and illustrate striking similarities in faunal assemblages of low latitude regions in the Atlantic, Pacific and Indian oceans. A high resolution biochronology, based on dominant faunal characteristics and 55 datum events, permits correlation between all three oceans with a high degree of precision. Population studies provide a view of the global impact of the paleoclimatic and paleoceanographic changes occurring during Middle Eocene to Oligocene time.Planktic foraminiferal assemblage changes indicate a general cooling trend between Middle Eocene to Oligocene time, consistent with previously published oxygen isotope data. Major faunal changes, indicating cooling episodes, occur, however, at discrete intervals: in the Middle Eocene 44-43 Ma (P13), the Middle/Late Eocene boundary 41-40 Ma ( ), the Late Eocene 39-38 Ma ( ), the Eocene/Oligocene boundary 37-36 Ma (P18), and the Late Oligocene 31-29 Ma ( ). With the exception of the boundary, faunal changes occur abruptly during short stratigraphic intervals, and are characterized by major species extinctions and first appearances. The Eocene/Oligocene boundary cooling is marked primarily by increasing abundances of cool water species. This suggests that the boundary cooling, which marks a major event in the oxygen isotope record affected planktic faunas less than during other cooling episodes. Planktic foraminiferal faunas indicate that the boundary event is part of a continued cooling trend which began during the Middle Eocene.Two hiatus intervals are recognized in low and high latitude sections at the Middle/Late Eocene boundary and in the Late Eocene ( ). These hiatuses suggest that vigorous bottom water circulation began developing in the Middle Eocene, consistent with the onset of the faunal cooling trend, and well before the development of the psychrosphere at the boundary.     

Cenomanian to Coniacian ostracodes from the Nkalagu area (SE Nigeria): biostratigraphy and palaeoecology

Ostracode faunas from three sections in the Nkalagu area, including the type section of the Nkalagu Formation, were analysed qualitatively and quantitatively. Based on the results, an ostracode zonation is proposed as follows: aCythereis sp. 2 Interval Zone (late Turanian to Coniacian), aCythereis vitiliginosa reticulata Interval Zone (Turonian) and aCytherella spp. Abundance Zone (Cenomanian). This zonation may be only of local (basin wide) character but should allow a micropalaeontological correlation of strata in the Lower Benue Trough. By comparison with sedimentological and palaeontological parameters, aCytherella dominated shelf/depleted oxygen-assemblage can be distinguished from aCythereis/Ovocytheridea dominated upper bathyal/suboxic to oxic-assemblage. The stratigraphic and palaeoecologic characteristics of the ostracode assemblages will be specified and presented in this contribution. Some of the species (Leguminocythereis sp.,Cytherella cf.comanchensis) are assumed to be replaced and to reflect the turbiditic activity in the area, resulting in carbonate deposits of economic significance.     

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.     

Micromammal biostratigraphy of the Upper Miocene to lowest Pleistocene continental deposits of the Guadix basin,southern Spain

Minwer‐Barakat, R., García‐Alix, A., Suárez, E.M., Freudenthal, M. & Viseras, C. 2012 xx xx: Micromammal biostratigraphy of the Upper Miocene to lowest Pleistocene continental deposits of the Guadix basin, southern Spain. Lethaia, Vol. 45, pp. 594–614. Recent study of the small mammals (rodents and insectivores) from several fossil‐bearing sites situated in the central sector of the Guadix Basin (Southern Spain) has notably increased the knowledge of the mammal assemblages that existed in Southern Iberia from the latest Miocene to the earliest Pleistocene. On the basis of this new information, we propose a biozonation for the continental deposits of the Guadix Basin, which consists of six biozones ranging in age from the late Turolian (MN13) to the early Villanyian (MN17). These biozones, defined according to the rules of the International Stratigraphical Guide, include not only the mentioned recently discovered fossil sites, but also other, previously known, localities of the basin. Finally, we integrate the described biozones in the Neogene Mammal units and the European Land Mammal Ages, correlate them with several classical mammal sites from other Iberian basins and the rest of Europe, and establish an approximate numerical age for the lower and upper limits of each biozone. □Biostratigraphy, Guadix Basin, rodents, insectivores, Upper Miocene, Pliocene, Pleistocene.     

中国北方上中新统的早期玛姆象属(Mammut)及其在玛姆象科(Mammutidae)分化和演化中的意义

玛姆象属是长鼻类玛姆象科这一重要类群的最终成员.虽然这一属在上新世的欧亚大陆和更新世的北美大陆广泛分布,它早期的进化历史却鲜为人知.报道了中国北方上中新统发现的斜脊玛姆象(相似种)(Mammut cf.M.obliquelophus)的新材料,包括一个几乎完整的幼年头骨,这些材料显示了玛姆象科的许多原始特征,因此很好地解释了玛姆象属形态特征的形成过程.斜脊玛姆象(相似种)具有强烈向两侧扩展的枕部,在门齿窝的基部具有收缩,这些特征与莫罗托始轭齿象(Eozygodon morotoensis)和广河豕脊齿象(Choerolophodon guangheensis)均具有相似性,后两者分别为玛姆象科与豕脊齿象科的早期代表.因此,玛姆象科与豕脊齿象科(Choerolophodontidae)具有近的亲缘关系,二者同位于象形类(Elephantimorpha)系统发育中的基部.支序分析支持了这一结论.     

中国北方上中新统的早期玛姆象属(Mammut)及其在玛姆象科(Mammutidae)分化和演化中的意义（英文）

玛姆象属是长鼻类玛姆象科这一重要类群的最终成员。虽然这一属在上新世的欧亚大陆和更新世的北美大陆广泛分布,它早期的进化历史却鲜为人知。报道了中国北方上中新统发现的斜脊玛姆象(相似种)(Mammut cf.M.obliquelophus)的新材料,包括一个几乎完整的幼年头骨,这些材料显示了玛姆象科的许多原始特征,因此很好地解释了玛姆象属形态特征的形成过程。斜脊玛姆象(相似种)具有强烈向两侧扩展的枕部,在门齿窝的基部具有收缩,这些特征与莫罗托始轭齿象(Eozygodon morotoensis)和广河豕脊齿象(Choerolophodon guangheensis)均具有相似性,后两者分别为玛姆象科与豕脊齿象科的早期代表。因此,玛姆象科与豕脊齿象科(Choerolophodontidae)具有近的亲缘关系,二者同位于象形类(Elephantimorpha)系统发育中的基部。支序分析支持了这一结论。