First record of Beremendia fissidens (Mammalia,Soricidae) in the Pleistocene of the Iberian Peninsula,with a review of the biostratigraphy,biogeography and palaeoecology of the species

This paper reports the first paleontological record of Beremendia fissidens (Mammalia, Soricidae) in the Iberian Peninsula during the second third of the Early Pleistocene. The species is exclusively present at the lowermost levels (Lower Red Unit: TE8–14) of the Sima del Elefante site, one of the largest stratigraphic sections of the Atapuerca cave complex (Burgos, Spain). The age of Sima del Elefante shows that this large-sized type of red-toothed venomous shrew inhabited the Sierra de Atapuerca more than 1.1 Myr ago, coexisting with an extremely rich and diverse faunal association of nearly 40 small and large mammalian species, including hominines. The presence of this species in the Atapuerca locality has important palaeoecological, palaeobiogeographical and biostratigraphic implications, which are extensively discussed here, throwing light on aspects largely left aside for this important group of red-toothed shrews, previously relegated to nothing but mere faunal lists, at least in the Iberian Peninsular context.     

New foot remains from the Gran Dolina-TD6 Early Pleistocene site (Sierra de Atapuerca, Burgos, Spain)

This paper presents and describes new foot fossils from the species Homo antecessor, found in level TD6 of the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These new fossils consist of an almost complete left talus (ATD6-95) and the proximal three-quarters of a right fourth metatarsal (ATD6-124). The talus ATD6-95 is tentatively assigned to Hominin 10 of the TD6 sample, an adult male specimen with which the second metatarsal ATD6-70+107 (already published) is also tentatively associated. Analysis of these fossils and other postcranial remains has made possible to estimate a stature similar to those of the specimens from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain). The morphology of the TD6 metatarsals does not differ significantly from that of modern humans, Neanderthals and the specimens from Sima de los Huesos. Talus ATD6-95, however, differs from the rest of the comparative samples in being long and high, having a long and wide trochlea, and displaying a proportionally short neck.     

A very diverse amphibian and reptile assemblage from the late Middle Pleistocene of the Sierra de Atapuerca (Sima del Elefante,Burgos, Northwestern Spain)

Among the Pleistocene localities of the Sierra de Atapuerca, the site called Sima del Elefante is famous for having delivered in its lower level (TE-LRU) some hominid remains currently considered the oldest in Western Europe (about 1.2 Ma). In its upper part (TE-URU), this site has fossiliferous levels pertaining to the late Middle Pleistocene (350-250 ka). Here we describe for the first time the amphibians and reptiles of these upper levels (TE18 and TE19), which have proved to be one of the richest assemblages of all the localities of the Sierra de Atapuerca. The faunal list is composed of 18 taxa made up of urodeles (Salamandra salamandra and Lissotriton helveticus), anurans (Discoglossus sp., Alytes sp. Pelobates cultripes, Pelodytes punctatus, Bufo bufo, Bufo calamita, Hyla arborea and cf. Rana sp.), a terrestrial tortoise (Testudo s.l.), lizards (Lacerta s.l., Podarcis sp. and Anguis fragilis) and snakes (Natrix natrix, Natrix maura, Coronella cf. girondica and Vipera latasti). For the first time, the genera Lissotriton, Discoglossus and Podarcis are mentioned in the Pleistocene localities of the Sierra de Atapuerca. TE19 also corresponds to the earliest mention for L. helveticus in the Iberian Peninsula. With the exception of tortoises, all these species are currently present in the province of Burgos, and thus suggests that the herpetofauna has a modern aspect since the Middle Pleistocene. This association of amphibians and reptiles suggests a slightly warmer climate than the current one, but may correspond to a “cold and dry” period if compared with other Pleistocene localities of the Sierra de Atapuerca. The landscape was probably composed of a gallery forest along a quiet water river within a Mediterranean environment alternating laterally between dry meadows, rocky or stony areas and open scrubland.     

Patterns of morphological variation amongst semifossorial shrews in the highlands of Guatemala,with the description of a new species (Mammalia,Soricomorpha, Soricidae)

Members of the Cryptotis goldmani group of small‐eared shrews (Mammalia, Soricomorpha, Soricidae) represent a clade within the genus that is characterized by modifications of the forelimb that include broadened forefeet, elongated and broadened foreclaws, and massive humeri with enlarged processes. These modifications are consistent with greater adaptation to their semifossorial habits than other members of the genus. The species in this group occur discontinuously in temperate highlands from southern Tamaulipas, Mexico, to Honduras. In Guatemala, there are three species: the relatively widespread Cryptotis goodwini and two species (Cryptotis lacertosus, Cryptotis mam) endemic to highland forests in the Sierra de los Cuchumatanes of western Guatemala. Ongoing studies focusing on the relationships of variation in cranial and postcranial skeletal morphology have revealed a fourth species from remnant cloud forest in the Sierra de Yalijux, central Guatemala. In this paper, I describe this new species and characterize its morphology relative to other species in the C. goldmani group and to other species of Cryptotis in Guatemala. In addition, I summarize available details of its habitat and ecology. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1267–1288.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

Twentieth anniversary of Homo antecessor (1997‐2017): a review

It has been twenty years since diagnosis and publication of the species Homo antecessor.¹ Since then, new human fossils recovered from the TD6 level of the Gran Dolina site (Sierra de Atapuerca, northern Spain) have helped to refine its taxonomic and phylogenetic position. In this paper, we present a synthesis of the most characteristic features of this species, as well as our interpretation derived from the latest investigations. We focus on the phylogenetic interpretation of Homo antecessor, taking into account the most recent paleogenetic analyses and a reassessment of the European Middle Pleistocene hominin record. We try to show that, twenty years after its publication, H. antecessor provides a good opportunity to address the morphology of the last common ancestor of Neandertals and modern humans.     

Enamel and dentine dimensions of the Pleistocene hominins from Atapuerca (Burgos,Spain): A comparative study of canine teeth

Enamel and dentin patterns have awakened a considerable interest in phylogenetic studies. However, almost nothing is known about the dental tissue proportions of European Pleistocene hominins, apart from Neanderthal populations. This study aims to assess the three-dimensional dental tissue proportions of permanent canines belonging to the extensive sample of hominin teeth at Sierra de Atapuerca (Spain) through the use of microtomographic techniques. Our results show that early and middle Pleistocene populations from Atapuerca exhibit large coronal and root dentine dimensions, as well as a thinly enamelled pattern, which has been traditionally considered an autapomorphic Neanderthal trait. Therefore, these results might support an early enamel thickness decrease which is already observed 800 kyr ago in Homo antecessor and maintained in later groups such as Sima de los Huesos and Neanderthal populations during the middle Pleistocene.     

L’approvisionnement en matières premières lithiques au Pléistocène inférieur et moyen dans la Sierra de Atapuerca, Burgos (Espagne)

This paper summarizes the latest studies on lithic raw material raising that are currently undertaken within the Sierra de Atapuerca research project. During the past two years, several works have provided new petrographical and technological data, contributing to the knowledge of the kinds of materials exploited by the Atapuerca hominids, as well as the areas where those materials were raised. The sites yielding this information are Gran Dolina (Lower and Middle Pleistocene) and Galeria (Middle Pleistocene), both located in the Trinchera del Ferrocarril archeopaleontological complex.     

Eight new species of Marasmodes (Asteraceae,Anthemideae) from South Africa

During a stay at the Kirstenbosch Research Centre in Cape Town (South Africa), several groups of Asteraceae were studied. One of these was the genus Marasmodes (tribe Anthemideae). After a careful taxonomic study of additional material, including the first species described by A. P. de Candolle, the author has concluded that eight collections should be considered as new species. These new species are described and their relationships with the most similar species of the genus are discussed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 330–342.     

Updated Atapuerca biostratigraphy: Small-mammal distribution and its implications for the biochronology of the Quaternary in Spain

The Sierra de Atapuerca (Burgos, Spain) contains the most comprehensive archaeological and paleontological evidence of human evolution in Europe during the Quaternary. The time scale based on the microfossil content of the sedimentary infillings of the caves of Atapuerca has been used to attribute relative ages to the various paleontological and archaeological levels and correlate amongst the different sites. Microfossils are particularly significant as they are the dominant constituents of the caves from the Pleistocene to the Holocene of the Sierra de Atapuerca. The microfossils of the Atapuerca sites are mainly composed of isolated bones and teeth from small vertebrates, and the hard tissue of the vertebrate skeleton can be taxonomically classified. In revisiting the distribution and the classification of the small-mammal taxa along five of the main stratigraphic sequences from Atapuerca (Sima del Elefante, Gran Dolina, Galería-Zarpazos, Sima de los Huesos, and Portalón), we apply a new method, the “Unitary Associations” method, in order to refine the existing biozones, adding five new Faunal Units, and interpret their relative age. Finally, we correlate Atapuerca with other Pleistocene sites of Europe.     

Étude analytique d’une clavicule complète de subadulte d’Homo antecessor (site de Gran Dolina, Sierra d’Atapuerca, Burgos, Espagne)

This study reports on the skeletal remains of an infant clavicle - specimen ATD6-37 - belonging to the Homo antecessor species, unearthed at Lower Pleistocene level TD6 of the Gran Dolina site (Sierra de Atapuerca). Studied alongside a further adult specimen - ATD6-50 -, they provide us with significant information on two key paleobiological aspects of these early humans: body shape and development. Based on the analytical results, the paper proposes a more accurate proportional method for determining age at death is applied to the fossilized infant clavicle under study. It goes on to hypothesize on postcranial growth and body shape and discusses morphological patterns and age at death of these early humans through comparisons with a wide range of infant dental samples and clavicular specimens in early and modern humans.     

Endocranial traits of the Sima de los Huesos (Atapuerca,Spain) and Petralona (Chalkidiki,Greece) Middle Pleistocene ursids. Phylogenetic and biochronological implications

A complete cranium of Ursus deningeri from Petralona (Chalkidiki, Greece) covered by a thick carbonate crust has been virtually liberated through CT scan and 3D image reconstruction, and compared with a specimen from the Sima de los Huesos, Sierra de Atapuerca (Spain) Middle Pleistocene site. X-ray computed tomography allowed for the identification of previously unrecognized and phylogenetically relevant traits. Analysis of these traits and an additional series of classically defined traits demonstrate the near identity of these specimens, thereby, confirming an early stage of the speloid “(cave bear)” condition. The replicate luminescence ages obtained for the SH deposit provide a combined minimum age estimate of 427 ± 12 ka for the underlying fossils. Based on a series of primitive and derived morphological features of these two crania, a similar age is here proposed for the Petralona and SH U. deningeri.     

Early and Middle Pleistocene hominins from Atapuerca (Spain) show differences in dental developmental patterns

The Bayesian statistical approach considers teeth as forming a developmental module, as opposed to a tooth-by-tooth analysis. This approach has been employed to analyze Upper Pleistocene hominins, including Neandertals and some anatomically modern humans, but never earlier populations. Here, we show its application on five hominins from the TD6.2 level of the Gran Dolina site (Homo antecessor, Early Pleistocene) and the Sima de los Huesos site (Middle Pleistocene) of the Sierra de Atapuerca (Burgos, northern Spain). Our results show an advanced development of the third molars in both populations with respect to modern Homo sapiens. In addition, the Sima de los Huesos hominins differ from H. sapiens and H. antecessor in the relatively advanced development of their second molar. The relative mineralization of I1/M1 in H. antecessor appears to be similar to that of modern humans, as opposed to that of Neandertals, which appear to be unique. These observations, combined with reduced enamel formation times and the advanced development of the third molars, appear to indicate a shorter ontogenetic period in the hominins from Gran Dolina and Sima de los Huesos in comparison to modern human average.     

Carnivores from the Early Pleistocene hominid-bearing Trinchera Dolina 6 (Sierra de Atapuerca, Spain).

The carnivores from Trinchera Dolina level 6 (TD6) in Sierra de Atapuerca include Ursus sp., Crocuta crocuta, Mustela palerminea, Lynx sp., Canis mosbachensis and Vulpes praeglacialis. Approximately 80 human remains belonging to Homo antecessor were found in the Aurora Stratum (AS): located in the upper part of the TD6 level (TD6, T36-43). The carnivores from AS are taxonomically similar to the rest of the TD6 level, differing only in the occurrence of Lynx sp. The spotted hyaena (Crocuta crocuta) inhabited the Atapuerca Sierra during the Early Pleistocene and up to the Middle Pleistocene, after which it is absent. According to palaeomagnetic, U/Th and ESR results, the fossils from TD6 are dated to the Early Pleistocene, which is the earliest certain occurrence in Europe of Crocuta crocuta. It is associated with Mimomys savini, the arrival of which in Europe can be correlated with the beginning of the Early Biharian. Crocuta crocuta may have arrived during the late Early Pleistocene ("Bavelian complex"), and subsequently invaded the rest of the continent during the Middle and Late Pleistocene. Atapuerca TD6-AS represents the most ancient deposits in Europe where Homo and spotted hyaenas coexisted and, they probably competed ecologically. The carnivores from TD6 are biochronologically consistent with the end of the Early Pleistocene or early Cromerian (corresponding with the Biharian biochron). The timespan could correspond with oxygen isotope stages 19, 20 or 21.     

Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca,Burgos, Spain)

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this study has identified unique morphological characteristics of the SH tali. They are vertically shorter than those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.     

Exceptional biting capacities of the Early Pleistocene fossil shrew Beremendia fissidens (Soricidae,Eulipotyphla, Mammalia): new taphonomic evidence

The discovery of small, very well-defined and perfectly preserved toothmarks on a humerus of a mole Talpa cf. europaea from level TE9 of Sima del Elefante (Sierra de Atapuerca, Burgos) with a chronology of the Early Pleistocene is extraordinary. In a previous paper, this bite was compared with current small carnivores such as Mustela nivalis molars and with fossil remains of Mustela palerminea and the soricid (Eulipotyphla and Mammalia) Beremendia fissidens with the purpose of identifying the predator. It was hypothesised that Beremendia fissidens could be the bite maker. However, it was not possible to rule out other predators due to the shortage of Beremendia fissidens remains. Recently, new fossil remains of this insectivore have been found in Sima del Elefante levels, allowing the upper dentition to be measured. These new data suggest that effectively Beremendia fissidens may have had the capacity to bite prey larger than itself, even though it did not possess morphological characteristics specialised for the consumption of small mammals. The addition of small vertebrates (like talpids) to complement a diet based on insects could have been a way of responding to the needs of the high metabolic rate characteristic of Beremendia fissidens.     

A new multiflorous species of Leymus (Poaceae: Triticeae) from western China

A new species of Leymus section Racemosus, L. pluriflorus L.B.Cai & T.L.Zhang, is described and illustrated. It grows in the eastern part of Qinghai Province and the southern part of Gansu Province, China. It most closely resembles L. crassiusculus L.B.Cai, from which it differs in having longer rachis internodes, some pedicellate spikelets, more florets per spikelet, glabrous lemmas, shorter paleas and shorter anthers. It differs from all other Chinese species taxa in Leymus with regard to the large number (8–12) of florets in its spikelets, and from all species of Leymus in adjacent countries in having three to four spikelets per node. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 343–348.