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.     

Rodents and insectivores from the late Miocene of Romerales (Fortuna Basin,Southern Spain)

Abstract

The Neogene Fortuna Basin (Murcia Region, SE Spain) is rich in microvertebrate sites. Its continental sections include localities extended from the middle Turolian (MN12) to the early Ruscinian (MN14). However, there are few works dealing with the taxonomy of these fossil assemblages. In this paper we provide a complete taxonomic study of the rodents and insectivores from three levels (ROM-2B, ROM-2C and ROM-3A) within the Romerales section. We infer a late Turolian age (late Messinian) for these levels, among which the richest and most diverse level is ROM-C, including at least 11 different taxa. In addition, the paleoecological analysis of these fossil assemblages suggests the dominance of open herbaceous meadows under temperate climate during the formation of these sites, with a slight decrease in temperature and humidity from ROM-2B to ROM-2C.     

Miocene barnacle assemblages from southern Spain and their palaeoenvironmental significance

The Tabernas Basin in southeast Spain is a Neogene intermontane basin which filled with sub-aerial and submarine fan conglomerates grading into turbidite sandstones towards the basin centre. The basin history involved the development of Seravallian-Tortonian subaerial fans which coalesced southwards from the basin margin. Increasing water depth in the Tortonian led to a submarine development of these fans. The coarse fan conglomerates are colonized almost solely by barnacles of the species Megabalanus tintinnabulum , known from Miocene-Recent deposits. The barnacles occur in seven preservational-state assemblages, which provide a record of the colonization of stabilized mass-flow conglomerates and of subsequent reworking episodes. M. tintinnabulum is known from intertidal environments today. In situ clusters developed on the fan conglomerates record the transition of the fans from subaerial to submarine environments. Mass-flow deposits rich in barnacle shell debris are representative of basin-wards reworking of intertidal clusters and illustrate relative increase in water depth. A further shallowing episode is identifiable by the recolonization of mass flows by in situ barnacle clusters. This case history is demonstrative of the utility of some barnacles in determining basin history with respect to minor sea level fluctuations.     

Eocene siliceous sponges from the Ebro Basin (Catalonia, Spain)

The Eocene (Bartonian) marls of the La Guixa Member and Gurb Member, Vic Marls Formation (Ebro Basin, Catalonia, Spain), contain a very rich and diversified siliceous sponge fauna. The fauna is dominated by hexactinellids; lithistids and other demosponges are rare. It consists of 16 species representing 16 genera. Eleven new species and two new genera are proposed for these sponges: Reguantella cavernosa nov. gen. nov. sp., Regadrella concinna nov. sp. (both Hexactinellida, Lyssacinosa), Eurete clava nov. sp., Pleuroguettardia iberica nov. sp., Aphrocallistes almeriae nov. sp., Hexactinella informis nov. sp. (all Hexactinellida, Hexactinosa), Brachiolites munterensis nov. sp., Centrosia viquensis nov. sp., Callicylix eocenicus nov. sp., Rhizocheton robustus nov. sp. (all Hexactinellida, Lychniscosa), Propetrosia pristina nov. gen. nov. sp. (Demospongia, Haplosclerida). Some genera of sponges in this fauna are still extant, but, in general, the predominant ones are very close in morphology, and, without doubt, closely related to the Late Cretaceous sponges. This fauna also differs considerably, in terms of composition, from most other described faunas of Tertiary sponges from the Mediterranean region, which are dominated by lithistid sponges. Lithistid sponges are rare in this investigated assemblage, which seems most similar to an as yet undescribed Eocene fauna from Italy. There is ecological differentiation in the proportions of particular sponges in various outcrops and/or stratigraphical levels that is clearly associated with water-depth-related controlling factor(s): Munter, Tona and Sta. Cecilia represent the deepest facies, Gurb is intermediate, and St. Roc and Vespella are the most shallow. The exact bathymetric position of the sponge fauna is difficult to estimate, but it seems that 100 m (but probably 200 m and more in the case of the deepest parts) of water depth may be inferred for this facies.     

New charophyte assemblage from middle Miocene lacustrine deposits of Moneva (Ebro Basin,Spain)

Two new fossil charophyte species from middle Miocene lacustrine deposits of Moneva (Ebro Basin, NE Spain) are described and illustrated. Sphaerochara miocenica nov. sp. is represented by medium sized gyrogonites with a characteristic spheroidal shape and ornamented with numerous small and regularly spaced tubercles arranged along the spiral cells. Several specimens show an unusual carbonate encrusted layer that covers completely or partially the gyrogonite. Psilochara monevaensis nov. sp. is characterized by large gyrogonites, with typical prolate shape and protruding apical poles. A second population of S. miocenica nov. sp. from a distant early upper Miocene lacustrine basin located in the Middle East (Bekaa Valley, Lebanon), identified in a previous study as Sphaerochara sp., is described and illustrated here, allowing the comparison of these distant populations. Ecologically, the associated fauna and facies from both localities suggest that these taxa grew in very shallow athalassic lakes. Interestingly, S. miocenica nov. sp. having a wide geographic distribution in peri-Mediterranean lakes with variable salinity, may become a fossil guide for the Miocene, providing at the same time important palaeoecological information.     

Late Miocene Erinaceinae from the Teruel Basin (Spain)

Classifying fossil teeth of Erinaceinae (spiny hedgehogs) is a challenging task, because of their scanty record and systematic treatment that heavily relies on skull characteristics. In this paper we describe the complete set of isolated dental elements of Erinaceinae from the upper Miocene sediments of the Teruel Basin (eastern Central Spain). Four different species were recognized: Postpalerinaceus cf. vireti, Atelerix aff. depereti, Atelerix steensmai nov. sp., and a form classified as Erinaceinae genus and species indet. All four are relatively derived in showing multi-purpose dentitions, not showing only adaptations to insectivory, but also to carnivory, herbivory and possibly durophagy/malacophagy. The temporal occurrence of spiny hedgehogs during the middle to late Miocene in the Teruel Basin and neighboring Calatayud-Montalbán Basin peaks within periods of relative aridity, a correlation consistent with modern geographic distribution. Messinian cooling is the best candidate for explaining a remarkable demise of Erinaceinae at 7 Ma.     

Rodents, insectivores and chiropterans (Mammalia) from the late Aragonian of Can Missert (Middle Miocene, Vallès-Penedès Basin, Spain)

In this paper, the insectivores, chiropterans and rodents from the middle Miocene site of Can Missert are described. The faunal list of this locality includes the following species: Miosorex grivensis, Desmanella sp., Talpidae indet., Vespertilionidae indet., Hispanomys daamsi, Megacricetodon minor, Megacricetodon ibericus, Fahlbuschia crusafonti, Democricetodon brevis nemoralis, Eumyarion medium, Muscardinus hispanicus, Eomuscardinus sp., Paraglirulus werenfelsi and Spermophilinus bredai. H. daamsi is a new Cricetodontine species which is characterized by complete ectolophs and relatively long third lower molars. The rodent association of Can Missert enable one to assign this locality to the late Aragonian, MN 8, being close in age to other localities in the Vallès-Penedès Basin such as Castell de Barberà. However, the proportion in which each species is represented is very different in the two cases. This evidence points to the existence climatic pulses at the end of the Aragonian Mammal-Stage.     

Correlation between the Paleocene/Eocene boundary and the Ilerdian at Campo, Spain

The Ilerdian is a well-established Tethyan marine stage, which corresponds to an important phase in the evolution of larger foraminifera not represented in the type-area of the classical Northwest-European stages. This biostratigraphic restudy of its parastratotype in the Campo Section (northeastern Spain) based on planktic foraminifera, calcareous nannofossils, dinoflagellate cysts and the distribution of the stable isotopes ∂¹³C and ∂¹⁸O is an attempt to correlate the Paleocene/Eocene boundary based on a characteristic carbon isotope excursion (CIE) marking the onset of the Initial Eocene Thermal Maximum (IETM) and the Ilerdian stage. The base of this ∂¹³C excursion has been chosen as the criterion for the recent proposal of the Global Stratotype Section and Point (GSSP) of the base of the Eocene (= base of the Ypresian) in the Dababiya Section (Egypt) to which an age of 54.9 Ma has been attributed. This level is also characterized by a marked extinction among the deep-water benthic foraminifera (Benthic Foraminifera Extinction Event, BFEE), a flood of representatives of the planktic foraminiferal genus Acarinina and the acme of dinoflagellate cysts of the genus Apectodinium. In the Campo Section, detailed biozonations (planktic foraminifera, calcareous nannofossils, dinoflagellate cysts) are recognized in the Lower and Middle Ilerdian. The correlation with the Ypresian stratotype is based on dinoflagellate cysts and calcareous nannofossils. The base of the Ilerdian is poor in planktic microfossils and its precise correlation with the redefined Paleocene/Eocene boundary remains uncertain.     

Reproducibility and reliability of the Pliensbachian calcareous nannofossil biohorizons from the Basque-Cantabrian Basin (Northern Spain)

Based on calcareous nannofossil assemblages identified in four expanded and well-dated sections from the Basque-Cantabrian Basin, the main objective of this paper is to improve the knowledge of the Pliensbachian calcareous nannofossil events, and to calibrate these events to the ammonite zones established for this area. The semiquantitative analysis of more than 140 smear slides from the Tudanca and Santotis sections, and the re-analysis of more than 200 smear slides from the Camino and San Andrés sections, have been carried out in order to describe the succession of calcareous nannofossil assemblages. Related to their composition changes, we have recognized and calibrated to the ammonite zones two main events: the first occurrences (FOs) of Similiscutum cruciulus and Lotharingius hauffii, and six secondary events: the FOs of Biscutum novum, Biscutum grande, Biscutum finchii, Lotharingius barozii and the FCOs (first common occurrences) of Calcivascularis jansae and L. hauffii. In the Camino and San Andrés sections, we also identify the FOs of Biscutum dubium, Bussonius prinsii and Lotharingius sigillatus. The obtained data allow us to assess the degree of reproducibility of the Pliensbachian calcareous nannofossil events in the studied area. The biohorizon succession recognized in the Basque-Cantabrian Basin are compared to those proposed for NW Europe, Lusitanian Basin, Italy and Southern France.     

Eocene fossil rodent assemblages from the Erlian Basin (Inner Mongolia,China): Biochronological implications

Rodents from the Nuhetingboerhe-Huheboerhe area in the Erlian Basin, Inner Mongolia, differ stratigraphically: primitive ctenodactyloids such as Chenomys and Yuanomys are dominant in the upper part of the Nomogen Formation; Tamquammys dominates in the Arshanto Formation; Asiomys, Pappocricetodon, and Yuomys appear in the lower part of the Irdin Manha Formation and Tamquammys is rarer in this formation than in the Arshanto Formation. The assemblage of the upper part of the Nomogen Formation is similar to that of the Lingcha Formation of Hunan, the Wutu Formation of Shandong, the Yuhuangding Formation of Hubei, and the Bumban Member of the Nara-Bulak Formation of Mongolia. The assemblage in the upper part of the Arshanto Formation is correlated with that from the locality Andarak 2 in Kyrgyzstan. The assemblage from the lower part of the Irdin Manha Formation resembles that of the lower part of the Hetaoyuan Formation of Henan.On the basis of the comparison of the rodent assemblages, I consider that the age of upper part of the Nomogen Formation corresponds to the Bumbanian land mammal age. The Bumbanian, Arshantan, and Irdinmanhan land mammal ages are correlated respectively to the early Ypresian, the middle–late Ypresian, and the early Lutetian of the Geological Time Scale. The Bumbanian and Irdinmanhan land mammal ages are also correlated to the early Wasatchian and the early Uintan (or the later Bridgerian) of the North American Land Mammal Ages.     

Muridae from the late Miocene site of Venta del Moro (Eastern Spain)

Murids are usually the dominant faunas of the late Miocene and early Pliocene micromammal assemblages. The present work deals with the murid faunas of the well-known late Miocene locality of Venta del Moro, comprising over 2700 molars ascribed to the taxa Apodemus gorafensis, Occitanomys alcalai, Paraethomys meini and Stephanomys dubari. This list differs from those of previous studies by the presence of A. gorafensis instead of A. gudrunae. The presence of A. gudrunae in younger localities than Venta del Moro implies that both species, considered ancestor and descendant, coexisted for some time. In addition, the analysis of such an extensive collection has allowed us to check the variability of the studied species in great detail, such as the development of the longitudinal connections of O. alcalai, to a greater extent than previously described. Finally, we discuss the biometrical changes of the lineages of Paraethomys in Southern Europe from MN13 to MN15, and hypothesize about the mechanisms behind these variations in size.     

Biostratigraphy and sedimentary evolution of Late Miocene and Pliocene continental deposits of the Granada Basin (southern Spain)

The Late Miocene and Pliocene continental sediments in the Granada Basin (southern Spain) have yielded large amounts of fossil small mammals in 37 localities from 11 sections. The aim of this paper is to integrate faunistic, stratigraphic, and sedimentary criteria to unravel the geological history of the continental infilling of the basin. The palaeontological study has led to a detailed biozonation on the basis of rodents, which helps to correlate in detail the different sedimentary units found in the basin, and to follow the changes of the different sedimentary systems and their palaeogeographical evolution through time. Combination of the proposed biostratigraphy and the reinterpretation of the magnetostratigraphic analyses of the Barranco del Purcal section allows us to assign an absolute age slightly older than 5.23 Ma to the Turolian–Ruscinian boundary (MN13‐MN14).     

Hoof-Like Unguals,Skin, and Foot Movements Deduced from Deltapodus Casts of the Galve Basin (Upper Jurassic-Lower Cretaceous,Teruel, Spain)

Galve (Teruel, Spain) is a town in the interior of a synclinal fold with Upper Jurassic marine limestones along its flanks, and, in its core, Upper Jurassic–Lower Cretaceous continental and shoreline sediments crop out. The core sediments cover an area about 8 km², and contain a concentration of sites with footprints, bones, and eggshells of dinosaurs. The footprints are both shafts and natural casts. Some casts are attributed to stegosaurs (Deltapodus). The Deltapodus casts are characterized by features that allow us to make direct observations on the skin formed by polygonal scales, and ellipsoidal “hooves,” as well as deductions on the movement of the limbs during walking. According to the opinion of some authors, dinosaur footprints are indicators of the motion of their limbs and sometimes of the whole body. So far, results have been deduced from theropod, ornithopod, and sauropod footprints. This article shows the results obtained from analis of the aforementioned Deltapodus casts, i.e., forelimb movement similar to that of the forelimbs of sauropods, and the rigid structure of the autopodial part of the hind limb.     

Biostratigraphical precisions on the Early Serpukhovian (Late Mississippian), by means of a carbonate algal microflora (cyanobacteria, algae and pseudo-algae) from La Serre (Montagne Noire, France)

The microbial-dominated part of a Late Mississippian (Early Carboniferous) carbonate olistolite at La Serre (Montagne Noire, southern France) yielded interesting algae and pseudo-algae. The constituents are well known cyanobacteria (not described) and different representatives of poorly known or new species: cyanobacteria: Aphralysia capriorae, Hedstroemia (?) serrana n. sp., Dasycladales: Paraepimastopora sp., Nanopora sp., Cabrieropora pokornyi and E. occitanica, Anatoliporaceae: Richella incrustata, Caulerpales: Poncetellina (?) sp., questionable Archaeolithophyllaceae: Hortonella sp., Archaeolithophyllaceae: Neoprincipia with Neopricipia fluegeli, Neoprincipia petschoriaeformis n. sp., Neoprincipia claviformis n. sp. and Archaeolithophyllum ex gr. lamellosum; pseudo-algae Aoujgaliida: Stacheoidella spissa, Ungdarella uralica, and Fasciellidae: Fasciella and Praedonezella, with both polymorph species: Fasciella kizilia, and Praedonezella ex gr. cespeformis. This carbonate microflora is dated as Early Serpukhovian (Pendleian) by its association with the foraminifera Janichewskina, “Biseriella” and “Warnantella”. This dating is confirmed by the little more advanced algal assemblage, compared to the Latest Visean (Brigantian) assemblages from Montagne Noire (e.g. Japhet, Castelsec), and more primitive than the Late Serpukhovian (Arnsbergian) of Ardengost (Pyrenees). The importance of the calcifoliid pseudo-algae—Fasciella, Praedonezella, and Calcifolium (although this latter is locally lacking)—is suggested for the biozonation of the Visean/Serpukhovian boundary.     

The marine paleogene of the tremp region (NE Spain)-depositional sequences,facies history,biostratigraphy and controlling factors

Summary The marine Paleogene of the Tremp Basin in the Central Southern Pyrenees corresponds to four depositional sequences which are related to global eustatic third order cycles (Tejas A 2.3–2.6). Associated transgressive and downlap surfaces coincide with boundaries of biozones. Lowstand systems tracts consist of estuarine and braid delta systems. Transgressive and highstand systems tracts are composed of carbonate banks and reefs. Slow thrust-induced changes of the basin topography conditioned the basic type and the areal distribution of carbonate highstand and clastic lowstand systems. Rapid relative sea level changes controlled the activity and internal dynamic of the depositional systems. E-W directed blind thrust anticlines are covered during highstand periods by carbonate fringing banks withNummulites bars. N-S orientation of thrust anticlines leads to the evolution of reef-dominated barrier banks and shelf lagoonal homoclinal ramps. On-bank transport of carbonate sands dominates during transgressions, off-bank transport during highstand periods. Continuous thrusting during the Ilerdian caused angular unconformities only in combination with relative sea level fall. Sequence-internal onlap configurations result from contemporaneous tectonic tilting. Fourth order carbonate bank margin cycles contain well developed lowstand tracts due to increased subsidence rates. Fourth order flooding surfaces are marked by paleosoil horizons at their landward continuation.     

New Ichnospecies of Cardioichnus from the Miocene of the Guadalquivir Basin,Huelva, Spain

A new ichnospecies of Cardioichnus, Cardioichnus reniformis isp. nov., is documented and described from marine strata of the Late Miocene of southwestern Spain. It is a bilobate resting trace with a heart‐shaped outline and is related to the work of a wedge‐shaped irregular echinoid of the spatangoid group.     

High-frequency cycles in Upper-Miocene ramp-temperate carbonates (Sorbas Basin,SE Spain)

Uppermost-Tortonian temperate carbonates occur at the southern margin of the Sorbas Basin (Almería, SE Spain). These carbonates, included in the Azagador Member, formed in a gentle, shallow-water ramp. Six facies cycles in ramp deposits comprise alternating bivalve-shell concentrations and coralline algal beds. The basic cycle reflects the landward advance, as relative sea level rose, of coralline algal deposits, which were the facies of the outer ramp, over bivalve biostromes, which grew in the shallower areas of the mid-ramp. Biostromes were mainly built by oysters and locally by Isognomon. In many cases, however, the removal of smaller shells by storms left only thin, discontinuous patches of large bivalves as residual remains of the oyster biostromes. Some original cycles might be missing due to complete removal of bivalve shells from the biostromes. The six cycles recognised, therefore, should be considered as the minimum number of original cycles in the Azagador carbonates. The available age constraints suggest these cycles were forced by orbital precession or some higher-frequency process. Lithological cycles forced by precession are characteristic of the basinal deposits laterally equivalent to the Azagador carbonates.     

Glacial refugia of temperate, Mediterranean and Ibero-North African flora in south-eastern Spain: new evidence from cave pollen at two Neanderthal man sites

Aim To locate glacial refugia of thermophilous plant species in Spain. Location Two south‐eastern Spanish Neanderthal man sites in Murcia; namely, the inland Cueva Negra del Estrecho del Río Quípar and the coastal Sima de las Palomas del Cabezo Gordo. Methods We use pollen found in cave sediments as a source of palaeobotanical and palaeoecological information. The findings are discussed with regard both to animal remains from both sites, and also to other refugia in south‐eastern Spain and elsewhere in the Iberian Peninsula. Results Both sequences show persistence of abundant mesothermophilous trees during the last glacial stage, suggesting both localities were reservoirs of phytodiversity and woodland species. At both sites, deciduous and evergreen oaks are the most abundant components, followed by a wide variety of deciduous trees and sclerophyllous shrubs, including Ibero‐North African xerothermic scrub near the coast. Conclusions Incomplete information underlies a common misapprehension that Iberian glacial refugia were confined to southernmost parts of the peninsula. A rather different picture of Quaternary refugia emerges from consideration of pollen sequences from caves (and other inputs such as macroscopic charcoal, spatial genetic structure of present‐day populations, faunal remains, and present‐day distribution of thermophilous species). This picture offers a view of numerous viable areas for woodland species in southern Spain, in addition to others in the mountain ranges, both in continental central Spain and those of northern Spain: these stretch from the Mediterranean coast of Catalonia to the westernmost extent of the Bay of Biscay.     

A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain

Coastal plant communities in arid southeastern Spain are characterized by insect-pollinated scrub species, which fail to occur in Quaternary pollen sequences from valleys, marshlands and marine cores. We investigate pollen–vegetation relationships in samples from soil surfaces, animal dung, and sediments in depressions or basins that, in theory, should have pollen spectra that are comparable to those from sedimentary basins elsewhere. Pollen spectra from basins or depressions are very susceptible to long-distance wind and water transport. This can mask representation of pollen from the surrounding insect-pollinated vegetation, as can over-representation of basin-margin halophilous and hydrophilous pollen. Pollen spectra from biogenic materials of animal origin are the best analogues of local and regional vegetation, and show the best analytical potential in terms of pollen concentration and taxon diversity. Pollination properties of the species studied indicate they will rarely be found in most conventional pollen records. It cannot be stressed too strongly that insight into Quaternary vegetation of arid regions demands complementary pollen analysis of coprolites, urine-cemented deposits, and cave sediments with preserved biotic remains, in addition to water-lain sediments.