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Victoriamys, a new generic name for Chaline's vole from the Pleistocene of Western Europe
Abstract
A combination of dental and mandibular characters demonstrates that Chaline's vole, known from the Pleistocene of Spain and Italy, originally named as Allophaiomys chalinei and later referred by some authors to Arvicola and Tibericola, is unrelated to either Allophaiomys or Arvicola and probably is not closely related to Tibericola. It is here considered as the only member of a new genus, Victoriamys, named for the cave from which V. chalinei nov. comb. was first described. (c) 2012 Published by Elsevier Masson SAS.
... Nevertheless, there appears to have been a significant difference in arvicolid diversity between northern and southern Spain during the Pleistocene that may be unrelated to sampling bias. This difference is particularly pronounced when we compare arvicolid species diversity from the Atapuerca sequence of north-central Spain to that of the Baza Basin and Cueva Victoria in the south (Agustí et al., 2010;Cuenca-Bescós et al., 2010a;Martin, 2012a). It is difficult to explain this dichotomy by taphonomic bias, and it may be that the difference in diversity is related to environmental zonation during the Pleistocene, with more habitat complexity to the north than the south, perhaps mediated by climatic patterns that mantained a more arid, Mediterranean climate to the south. ...
... It is difficult to explain this dichotomy by taphonomic bias, and it may be that the difference in diversity is related to environmental zonation during the Pleistocene, with more habitat complexity to the north than the south, perhaps mediated by climatic patterns that mantained a more arid, Mediterranean climate to the south. Despite these differences, the combination of arvicolid records from the rich Atapuerca sequence with those of the Baza Basin, where a number of critical fossil assemblages are stratigraphically superposed and supplemented by a detailed paleomagnetic (pmag) history (Gibert et al., 2007;Cuenca-Bescós et al., 2010a;Martin, 2012a), allows the construction of an arvicolid biochronological Pleistocene sequence in Spain that is unparalleled for western Europe (Fig. 1). ...
... The answer to this problem seems to be that we need to branch out to molars other than the m1 and to mandibular anatomy. For instance, as noted by Gibert et al. (2007) and Martin (2012a), all known species of Arvicola have a complex m3 (with distinct T4) often mimicking the morphology of m2. Although this morphology may also be seen in some Mimomys species and Victoriamys, it is not seen in Microtus, in which T4 is usually reduced or absent. ...
... The last three genera occur at the reverse deposits of Atapuerca TE site but not at TD site (Cuenca-Bescos et al., 2015). Among the rodents, arvicolids tend to have the greatest value for biostratigraphy on the Iberian Peninsula (Cuenca-Besc os et al., 2010;Martin, 2012). The only arvicolid present at Cueva Victoria is V. chalinei, which has been reported from the sites of Cueva Negra, Atapuerca (Trinchera Dolina, levels 3e8), Almenara-Casablanca 3, Cal Guardiola in the Vallparadís section, and El Chaparral (Agustí and Galobart, 1986;Minwer-Barakat et al., 2011;L opez-García et al., 2012;Martin, 2012). ...
... Among the rodents, arvicolids tend to have the greatest value for biostratigraphy on the Iberian Peninsula (Cuenca-Besc os et al., 2010;Martin, 2012). The only arvicolid present at Cueva Victoria is V. chalinei, which has been reported from the sites of Cueva Negra, Atapuerca (Trinchera Dolina, levels 3e8), Almenara-Casablanca 3, Cal Guardiola in the Vallparadís section, and El Chaparral (Agustí and Galobart, 1986;Minwer-Barakat et al., 2011;L opez-García et al., 2012;Martin, 2012). V. chalinei (¼Allophaiomys chalinei), was initially reported from the late Early Pleistocene deposits of Trinchera Elefante at Atapuerca (Laplana and Cuenca-Bescos, 2000;Van der Made et al., 2003) but later cited only from the younger Trinchera Dolina (Cuenca-Bescos et al., 2015). ...
... V. chalinei (¼Allophaiomys chalinei), was initially reported from the late Early Pleistocene deposits of Trinchera Elefante at Atapuerca (Laplana and Cuenca-Bescos, 2000;Van der Made et al., 2003) but later cited only from the younger Trinchera Dolina (Cuenca-Bescos et al., 2015). Martin (2012) noted that V. chalinei had been recovered only from localities between the Jaramillo subchron and the base of the Brunhes (0.98e0.78 Ma), and referred Cueva Victoria to that time period. In addition, some new taxa have been described from Cueva Victoria: a new species of canid, Cuon rosi (Pons-Moy a and Moy a-Sol a, 1978), the arvicolid, A. chalinei (Alcalde et al., 1981), later assigned to V. chalinei (Martin, 2012), a new species of a lagomorph, Oryctolagus giberti (De Marf a, 2008), and a new species of deer, Megaloceros novocarthaginiensis (Van der Made, 2015). ...
Cueva Victoria has provided remains of more than 90 species of fossil vertebrates, including a hominin phalanx, and the only specimens of the African cercopithecid Theropithecus oswaldi in Europe. To constrain the age of the vertebrate remains we used paleomagnetism, vertebrate biostratigraphy and (230)Th/U dating. Normal polarity was identified in the non-fossiliferous lowest and highest stratigraphic units (red clay and capping flowstones) while reverse polarity was found in the intermediate stratigraphic unit (fossiliferous breccia). A lower polarity change occurred during the deposition of the decalcification clay, when the cave was closed and karstification was active. A second polarity change occurred during the capping flowstone formation, when the upper galleries were filled with breccia. The mammal association indicates a post-Jaramillo age, which allows us to correlate this upper reversal with the Brunhes-Matuyama boundary (0.78 Ma). Consequently, the lower reversal (N-R) is interpreted as the end of the Jaramillo magnetochron (0.99 Ma). These ages bracket the age of the fossiliferous breccia between 0.99 and 0.78 Ma, suggesting that the capping flowstone was formed during the wet Marine Isotopic Stage 19, which includes the Brunhes-Matuyama boundary. Fossil remains of Theropithecus have been only found in situ ∼1 m below the B/M boundary, which allows us to place the arrival of Theropithecus to Cueva Victoria at ∼0.9-0.85 Ma. The fauna of Cueva Victoria lived during a period of important climatic change, known as the Early-Middle Pleistocene Climatic Transition. The occurrence of the oldest European Acheulean tools at the contemporaneous nearby site of Cueva Negra suggest an African dispersal into SE Iberia through the Strait of Gibraltar during MIS 22, when sea-level was ∼100 m below its present position, allowing the passage into Europe of, at least, Theropithecus and Homo bearing Acheulean technology.
... Up to 39 species of bird have been recognized (Sánchez-Marco, 2004). A number of new taxa have been defined from Cueva Victoria, including the canid Cuon rosi (Pons-Moyà and Moyà-Solà, 1978); the arvicolid Allophaiomys chalinei (Alcalde et al., 1981), later reassigned to a new genus, Victoriamys by Martin (2012); and the lagomorph Oryctolagus giberti (de Marfà, 2008). The faunal association also includes Homo sp., based on an intermediate human phalanx (Gibert and Pons-Moyà, 1985;Pons-Moyà, 1985;Gibert et al., , 2002Santamaría and Gibert, 1992;Palmqvist et al., 1996), which some authors (Martínez-Navarro et al., 2005) have suggested may belong to Theropithecus (see discussion in Gibert et al., 2008, andMartínez-Navarro et al., 2008). ...
... Previous biostratigraphic data placed the fossils from Cueva Victoria in the Early Pleistocene, but younger than those from Venta Micena (Alcalde et al., 1981) at around 1.3 Ma (millions of years ago) . New paleomagnetic and biostratigraphic data provide a more accurate age of around 1.0 Ma, based on a reversal located in the capping flowstone (Fig. 1b), interpreted either as Jaramillo (Gibert et al., 2006) or Brunhes (Martin, 2012). ...
... It is known from Pliocene and Pleistocene deposits in South, East and North Africa. It has also been identified from Pleistocene sites outside Africa, in Mirzapur, India (Gupta and Sahni, 1981;Delson, 1993), dated by paleomagnetism at between w1.0 and 0.1 Ma (Azzaroli and Napoleone, 1982) and in Cueva Victoria, SE Spain (Gibert et al., 1995;this study), dated also at w1.0 Ma by both paleomagnetism and micromammal biostratigraphy (Gibert et al., 2006;Martin, 2012). ...
... For their relevance on the systematics of Pleistocene voles, m1s and M3s are often the only elements described and metrically studied (Cuenca-Bescós and Morcillo-Amo, 2022). To achieve this, we follow the terminology by López-García et al. (2015) and Martin (2012). Finally, the arvicoline first lower molars are metrically compared (through scatter plots and principal component analysis) with an assortment of Southern European sites relevant for their specific determination (López-García, 2008;, 2020. ...
... The buccal and lingual triangles are comparable in size. These characteristics are diagnostic features of Victoriamys chalinei (Martin, 2012;Cuenca-Bescós and Morcillo-Amo, 2022). Meanwhile, NOVA-FCT-DCT-5383 metrically falls within the variability of the specimens of Victoriamys chalinei from Cueva Victoria and El Chaparral (Fig. 8). ...
The Epivillafranchian (1.2 to 0.8 Ma) fossil vertebrate assemblages of Portugal are extremely poorly known compared with to those from the eastern half of Iberia. We review material from one of the two localities of this age previously known in Portugal, Algoz, and present a new microvertebrate locality, Santa Margarida. Both localities are situated in Algarve (South Portugal). The fauna of Algoz includes Hippopotamus antiquus, Eucladoceros sp., Metacervocerus rhenanus and an indeterminate leporid. There are remains of at least two individuals of each deer species from Algoz, although most of the remains are from the same juvenile individual of Eucladoceros sp., about 6 months old. The age of Algoz is determined to be ∼1.2 Ma given the combination of species and their biometrical and morphological characters. Santa Margarida has at least ten species of microvertebrates: Lacertidae indet., Crocidura sp., Sorex sp., Oryctolagus cf. cuniculus, Eliomys cf. quercinus, Apodemus cf. sylvaticus, Allocricetus bursae, Victoriamys chalinei, Iberomys huescarensis, and Iberomys brecciensis. This is the first occurrence of Victoriamys chalinei and Iberomys huescarensis –two vole species typical from the Early Pleistocene – in Portugal. Santa Margarida is younger than Algoz, ranging between ∼0.9 and ∼0.6 Ma, which means that the Early-Middle Pleistocene transition could be represented by this locality.
... The importance of the Short Haul record is that it documents an ancient age for the origination of Microtus and also suggests that early Pleistocene European Microtus are the result of a later dispersal, probably originating in Asia. Other arhizodont arvicoline genera appearing during the early Pleistocene include the Old World Arvicola, Chionomys, Tibericola, Stenocranius and Victoriamys (Agustí et al., 2013;Cuenca-Bescós et al., 2016;Martin, 2012). Early Pleistocene arhizodont voles also originating as end members of endemic mimomyine stems in East Asia include Huanonomys and Heteromimomys (Zhang et al., 2010;Zheng, 1992). ...
... While there may be a temporal difference between the B. del Paso and B. de los Conejos localities, we suspect the difference is slight, and neither is likely to be as old as the sites of Galera 2 and Zújar 14 from the K. gusii biozone. The combination of O. giberti, (arhizodont) Tibericola vandermeuleni and Mimomys oswaldoreigi defines the Mimomys oswaldoreigi biozone of Agustí et al. (2015) and Arvicolid Zone 1 of Martin (2012Martin ( , 2016. In eastern Europe, this interval correlates with the early Tamanian (= Odessian) faunal horizon of the eastern European "Eopleistocene" (Rekovets and Nadachowski, 1995) and regional biochrons MQR 11-10, equivalent to the early Biharian of western Europe (Tesakov, 2004). ...
A new arhizodont arvicoline rodent is described from early Pleistocene deposits of Barranco de los Conejos and Barranco del Paso in the Guadix-Baza Basin of southern Spain. The molars of Orcemys appear to represent a paedomorphic origination from a large rhizodont Mimomys. A few dental characters of Orcemys are superficially similar to those of the lagurines, but the presence of sparse cementum in reentrant folds and a Mimomys-kante formed opposite T5 on the first lower molar clearly identify Orcemys as an arvicoline. The character mosaic of Orcemys is unique among large early Pleistocene voles and the dentition of a potential ancestor probably resembled that of Mimomys medasensis with a tendency towards simplification. With Tibericola vandermeuleni and Mimomys oswaldoreigi, Orcemys represents one of the earliest experiments with arhizodonty among European voles. Including Mimomys medasensis at Barranco del Paso, this set of arvicolids redefines a previously recognized early Pleistocene MmQ1 biozone in Spain.
... Remarks: Victoriamys chalinei is a new combination to which Allophaiomys chalinei was recently allocated by Martin (2012). In the Iberian Peninsula, in addition to the type-locality of Cueva Victoria, this species has been reported from several late Early Pleistocene localities, e.g., Atapuerca (TD3-TD6 levels; Cuenca-Bescó s et al., 2010), Torrent de Vallparadís (layer Cal Guardiola D5;Minwer-Barakat et al., 2011), and Almenara-Casablanca (ACB 3 deposit; Agustí et al., 2011). ...
... These deposits habe been referred to the Allophaiomys chalinei biozone (0.98-0.79 Ma; Cuenca-Bescó s et al., 2010), i.e., in a much more recent time interval than the Italian site of Pietrafitta . Due to the revised biochronology of the Spanish Early Pleistocene deposits and the lack of a suitable ancestor (Cuenca-Bescó s et al., 2010;Martin, 2012), V. chalinei is no longer considered an Iberian endemic species. It might have been an immigrant to the Iberian Peninsula, with an older, still unclear, evolutionary history elsewere, as suggested by its occurrence in the Italian sites of Pietrafitta and Monte Argentario, significantly older than the Spanish sites Cuenca-Bescó s et al., 2010). ...
... In this paper we add to the diagnosis and descriptions of I. cabrerae by Chaline (1967Chaline ( , 1972 and Ayarzagüena and López-Martínez (1976) the fact that the mandibular foramen in the medial face is further back in comparison to other arvicolines (Fig. 4e), a character also noted by Gromov and Polyakov (1992). Of diagnostic value according to Martin (2012), the mandibular foramen is located in a more posterior po-sition than in Arvicola, Terricola, Microtus and Chionomys, in this respect resembling Victoriamys. The lower incisors are shorter than in other arvicolines, only reaching the beginning of the angular apophysis of the mandible (Fig. 4e), as also underscored by Gromov and Polyakov (1992). ...
... In our opinion, given the triangular character of the ACC in m1, the species that is morphologically closest to Iberomys is A. nutiensis. It is significant that in the Iberian Peninsula the species A. nutiensis (Agustí & Galobart 1986;Melero Rubio 2008;Cuenca-Bescós et al. 2010, 2012 is found in levels prior to the first appearance of I. huescarensis, in what we have called pre-Jaramillo faunas, in the A. lavocati Biozone (Cuenca- . ...
The extant Cabrera’s vole, Microtus cabrerae, differs in morphology and evolutionary
history from the other species of Microtus. This arvicoline has unique, derived features in the
cranium, mandible and dentition. Probably its most conspicuous features are its large size, the
high skull in lateral view, the long and distally broad nasals, and the triangle-shape of the
anteroconid complex, with a marked labio-lingual asymmetry of the occlusal surface of the
first lower molars. In this study, we propose a phylogenetic lineage that includes Cabrera’s
vole in what until now has been the Microtus subgenus Iberomys. Paleontological
information and several life history traits support the elevation of Iberomys to the rank of
genus. Genus Iberomys comprises species, which have appeared in succession during the
Quaternary: in the Early Pleistocene, the extinct I. huescarensis; in the Middle Pleistocene,
the extinct I. mediterraneus; and in the Late Pleistocene, the extant I. cabrerae. Interestingly,
the extant species shows several biological singularities, such as multiple polymorphic copies
of the SRY male-specific gene in both males and females, and the lowest basal metabolic rate
in relation to weight among arvicoline species. Likewise, its habitat requirement is unique
among the Iberian arvicolines. Accordingly, the biological and paleontological data that we
present in this work support the elevation of its taxonomic rank to that of genus. This study
also suggest a modification of nomenclature: Microtus (Iberomys) brecciensis is replaced
with Iberomys mediterraneus and the common name of the extant M. (I.) cabrerae changed
from ‘topillo’ to ‘iberon’ in order to improve conservation and protection actions.
... In this paper we add to the diagnosis and descriptions of I. cabrerae by Chaline (1967Chaline ( , 1972 and Ayarzagüena and López-Martínez (1976) the fact that the mandibular foramen in the medial face is further back in comparison to other arvicolines (Fig. 4e), a character also noted by Gromov and Polyakov (1992). Of diagnostic value according to Martin (2012), the mandibular foramen is located in a more posterior po-sition than in Arvicola, Terricola, Microtus and Chionomys, in this respect resembling Victoriamys. The lower incisors are shorter than in other arvicolines, only reaching the beginning of the angular apophysis of the mandible (Fig. 4e), as also underscored by Gromov and Polyakov (1992). ...
... In our opinion, given the triangular character of the ACC in m1, the species that is morphologically closest to Iberomys is A. nutiensis. It is significant that in the Iberian Peninsula the species A. nutiensis (Agustí & Galobart 1986;Melero Rubio 2008;Cuenca-Bescós et al. 2010, 2012 is found in levels prior to the first appearance of I. huescarensis, in what we have called pre-Jaramillo faunas, in the A. lavocati Biozone (Cuenca- . ...
The extant Cabrera's vole, Microtus cabrerae, differs in morphology and evolutionary history from the other species of Microtus. This arvicoline has unique, derived features in the cranium, mandible and dentition. Probably its most conspicuous features are its large size, the high skull in lateral view, the long and distally broad nasals, and the triangle-shape of the anteroconid complex, with a marked labio-lingual asymmetry of the occlusal surface of the first lower molars. In this study, we propose a phylogenetic lineage that includes Cabrera's vole in what until now has been the Microtus subgenus Iberomys. Paleontological information and several life history traits support the elevation of Iberomys to the rank of genus. Genus Iberomys comprises species, which have appeared in succession during the Quaternary: in the Early Pleistocene, the extinct I. huescarensis; in the Middle Pleistocene, the extinct I. mediterraneus; and in the Late Pleistocene, the extant I. cabrerae. Interestingly, the extant species shows several biological singularities, such as multiple polymorphic copies of the SRY male-specific gene in both males and females, and the lowest basal metabolic rate in relation to weight among arvicoline species. Likewise, its habitat requirement is unique among the Iberian arvicolines. Accordingly, the biological and paleontological data that we present in this work support the elevation of its taxonomic rank to that of genus. This study also suggest a modification of nomenclature: Microtus (Iberomys) brecciensis is replaced with Iberomys mediterraneus and the common name of the extant M. (I.) cabrerae changed from ‘topillo’ to ‘iberon’ in order to improve conservation and protection actions. This article is protected by copyright. All rights reserved
... The LAD (last appearance datum) for the species Beremendia fissidens, Asoriculus gibberodon, Sorex (D.) margaritodon, Crocidura kornfeldi, Ungaromys, Castillomys, Pliomys simplicior, and Allophaiomys occurs for the last time in the levels of the Sima del Elefante Lower Red Unit (TELRU: TE7eTE14) (Rofes and Cuenca-Besc os, 2006, 2009, 2011Cuenca-Besc os et al., 2010a (Fig. 2). In contrast, in Gran Dolina levels TD3e4 the species Stenocranius gregaloides, Pliomys episcopalis and Victoriamys chalinei (formerly classified under the genus Allophaiomys as Allophaiomys chalinei; see Martin, 2012), as well as species of modern Crocidura, Microtus, Iberomys and Allocricetus, appear for the first time in Sierra de Atapuerca (Cuenca-Besc os et al., 1995, 2005. ...
... The current distribution of these species is restricted more or less to the Mediterranean bioclimatic region (mainly meso-and thermo-Mediterranean areas); they are found mainly at less than 600 m a.s.l. P. cultripes generally lives in Table 1 Relation of the microfauna species found at the Early Pleistocene sites of the Iberian Peninsula L opez-Antoñanzas and Cuenca-Besc os, 2002;Agustí and Madurell, 2003;Blain, 2003;Furi o, 2003Furi o, , 2010van der Made et al., 2003;Cuenca-Besc os and García, 2007;Blain et al., 2008Carbonell et al., 2008;Rofes and Cuenca-Besc os, 2009, 2011Agustí et al., 2010;Bail on, 2010;Madurell-Malapeira et al., 2010;Minwer-Barakat et al., 2011;Martin, 2012 ...
... The site presents one of the best records of fossil fauna from the European Early Pleistocene, with approximately one hundred identified vertebrate species (Gibert & Ferràndez-Cañadell, 2015), mainly due to its condition as a den for the hyena P. brevirostris (Gibert et al., 1992). It is also noteworthy that taxa new to science have been discovered in Cueva Victoria, such as the lagomorph Oryctolagus giberti de Marfà, 2008, the arvicolid Victoriamys chalinei Martin, 2012, and the giant deer Megaloceros novocarthaginiensis Van der Made, 2015. ...
Cueva Victoria is a paleontological site of late Early Pleistocene age (ca. 0.9 Ma) located in the southeast of the Iberian Peninsula (Cartagena, Region de Murcia), which yielded diverse and abundant fossil remains, especially of large mammals. This accumulation is regarded as mainly deriving from the action of the giant hyena Pachycrocuta brevirostris, which used the cave as a den. Here, we present some preliminary taphonomic information deriving from the study of a portion of the large mammal collection from Cueva Victoria, including formerly uncatalogued material collected since the 1980s, which has not been considered in previous research. The study of this material started in 2018, when, owing to a suspension of the excavation activities in the cave, permissions were obtained to examine the uncatalogued specimens recovered during previous campaigns. Alongside a large number of remains of terrestrial mammals such as equids and cervids, Pachycrocuta brevirostris carried into the cave remains of groups uncommon in continental settings, such as marine mammals (cetaceans, seals). Beginning with this case, with then discuss the relevance of hyena-seal interactions in the Pleistocene of Europe in terms of ecology, as well as of seal documentation in the fossil record. From a critical review of the available evidence, Cueva Victoria stands out as a clear example of the consumption of seal by hyenas, with another likely case being represented by the much younger site of Lunel-Viel (late Middle Pleistocene, France). In other known cases of co-occurrence of hyenas and seals, the presence of seals is either due to other factors (e.g., hominins, deposition of marine sediments) or its origin cannot be ascertained for sure.
... Los restos fósiles recuperados en Cueva Victoria son de gran interés ya que, entre ellos, se han hallado varias especies y un género desconocidos previamente. Así, los taxones definidos en el yacimiento son 4: el cánido Cuon rosi (Pons-Moyà y Moyà-Solà, 1978), el lagomorfo Oryctolagus giberti (De Marfà, 2008), el cérvido Megaloceros novocarthaginiensis (Van der Made, 2015), y el arvicólido Allophaiomys chalinei (Alcalde et al., 1981), que pasó a ser la especie tipo de un nuevo género, Victoriamys (Martin, 2012). ...
La Cueva-Mina Victoria forma parte de un sistema cárstico de más de 3 km de longitud. Alberga un relleno Cuaternario con numerosos vertebrados, algunos descritos por primera vez. La actividad minera afectó a este enclave y permitió su descubrimiento, enriqueciendo al patrimonio natural con una historia y cultura singulares en la zona. Todo ello ha llevado a la protección legal del entorno como Bien de Interés Cultural. A pesar de esta riqueza en valores naturales y culturales, no se ha publicado, hasta la fecha, una valoración patrimonial del sitio que, de forma objetiva, exponga los criterios que aportan valor al bien y que permita un mínimo análisis o diagnóstico de los aspectos científicos más relevantes para su posterior gestión. Gracias a las ayudas económicas para la investigación e intervención del patrimonio arqueológico y paleontológico de la Región de Murcia y a la cofinanciación del Ayuntamiento de Cartagena, la Fundación Cidaris ha realizado, por primera vez, trabajos encaminados al conocimiento y gestión de los valores patrimoniales de la cueva. Se presentan los resultados del valor científico (Medio alto a Alto), discutiendo su significado y comparando los mismos con los de otros Lugares de Interés Geológico.
... The arvicolines are the most interesting taxa for age determination. Victoriamys chalinei is a typical species from the late Early Pleistocene of the Iberian Peninsula, being it restricted to the Arvicoline Zone 5 according to Martin, (2012) that ranges approximately between 1 million and 800.000 YBP. ...
The Quaternary fossil record of Portugal is important for our understanding of the paleobiodiversity in Iberia. In the present master thesis a series of studies augment our knowledge about this topic. A census of Quaternary paleobiodiversity is carried out in order to test how reliable the fossil record is for detecting living species, resulting in that ~38% of living terrestrial tetrapods are recognized in the fossil record for Portugal, although the number of species recognized varies between groups. The body mass of a Portuguese proboscidean (Palaeoloxodon antiquus) is calculated via numerical methods for the first time (11metric tons) and morphometric comparisons of this species with Mammuthus primigenius are presented using an extensive Proboscidean sample. A new fossil brown bear (Ursus arctos) locality, Algar do Vale da Pena, with numerous claw mark in the walls of the cave (the first of this type of marks described in Portugal) is presented and the fossil bear remains identified and compared to a sample from NW Spain. The bears from Algar do Vale da Pena contrast with other previously known Portuguese brown bear specimens by relative small size. A new microvertebrate locality from Algarve, Santa Margarida, is presented. It is an extraordinary rich site with one fossil for every two grams of sediment selected and processed. The locality provided the first record of two arvicoline taxa in Portugal (Iberomys huescarensis and Victoriamys chalinei), which allows giving a minimal age of around 800.000 YBP for at least part of it. This makes Santa Margarida one of the oldest three localities in the Pleistocene of Portugal.
... Las medidas del M1 inferior de Allophaiomys lavovati de la capa 7 del testigo norte están dentro de los rangos de tamaño de los ejemplares de la Sima del Elefante de Atapuerca (Laplana y Cuenca-Bescós, 2000), Genero Victoriamys (Martin, 2012) Victoriamys chalinei (Alcalde et al., 1981) Material: un M1 inferior de la capa 9 del Testigo Norte. Medidas: ver figura 2C. ...
RESUMEN El Alto de la Picarazas (Andilla-Chelva) es un yacimiento arqueológico con un amplio registro bioestratigráfico, correspondiente a un tramo final del Pleistoceno inferior y a los inicios del Pleistoceno medio. La cronología absoluta inferida a partir del estudio de micromamífersos y macromamíferos cubre una amplia horquilla cronológica de entre 0,78 Ma y 1,6 Ma. A lo largo de toda esta amplia secuencia se han documentado episodios de presencia humana, aspecto que no vamos a desarrollar en este trabajo.
... The chronology of this association, based on magnetostratigraphy, Th/U dating of flowstones, and micromammal biostratigraphy, is constrained to 0.90e0.85 Ma (Martin, 2012;Budsky et al., 2015;Gibert and Scott, 2015). Similarly to that from Vallparadís, the Cueva Victoria faunal assemblage has been related to an open environment surrounded by forest and with warm temperatures (Blain, 2015). ...
Several species of the genus Canis (Carnivora: Canidae) have been recorded from the European Early
Pleistocene, but the phylogenetic relationships among them and in relation to extant members of this
genus are still unclear. This is particularly true for the medium-sized and wolf-like extinct species Canis
mosbachensis. It has been considered by many researchers as a descendant of the larger Canis etruscus
and as a likely putative ancestor of extant wolves (Canis lupus). Other scholars, in contrast, have advocated
instead for a closer relationship between C. mosbachensis and the extinct Canis arnensis, and even a
close relationship between C. mosbachensis and C. lupus has been questioned. Here we describe the
previously unpublished medium-sized Canis remains from the late Early Pleistocene site of Vallparadís
Estacio, along with additional new Canis material from the roughly coeval site of Cueva Victoria (both in
the Iberian Peninsula), and compare them qualitatively and morphometrically with both extant and
extinct species of this genus. The described material most closely resembles in craniodental size and
shape the remains from Central and Southern Europe that have been previously assigned to
C. mosbachensis, to which they are hence formally attributed. The excellent preservation of the newly
described specimens (which include the most complete skull of this taxon) enables the description of
features previously unknown for this species, which further support a close phylogenetic link with living
wolves. Based on the described material, we review the role played by C. mosbachensis in the evolutionary
history of European fossil canids, and conclude that this extinct species is most closely related to
C. lupus and other closely-allied species, such as Canis anthus and Canis latrans.
... abaigari, P. abaigari or P. jaegeri does not co-occur with P. meini. In addition, although we can apply a 'time for space' substitution for the tentative recognition of a pattern analogous of character displacement along time in palaeontological contexts (Martin 2012), they should be restricted to the same area, while Figure 5 represents localities from the Teruel, Alcoy, Cabriel and Granada Basins, as well as the Table 3. Measurements (in millimetres) of the teeth of Paraethomys meini from Venta del Moro, according to the notation of the fossiliferous levels sensu Montoya et al. (2006 ; cr-6, crevillente 6 (de Bruijn, personal comment); aF-1'06, alcoi Forn 1'06 ; PEr-E, Peralejos E; Var, Villalba alta río (adrover et al. 1988); aBs-3a, alcoi Barranc sud 3a; ac-0 and ac-0B, alcoi cristian 0 and 0B ; lB2B, la Bullana 2B ; lg-4, la gloria 4; cEl-9, celadas 9 (adrover et al. 1993); gor-1, gorafe 1 (this work); al2-c and al2-D, alcoi 2c and 2D (Mansino et al. 2013); lJ, la Juliana (Montenat & de Bruijn 1976); MH, Mont-Hélène (aguilar et al. 1986); sE, séte (Michaux 1969;adrover 1986). ...
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.
... The chronology of this association, based on magnetostratigraphy, 230 Th/U dating of flowstones, and micromammal biostratigraphy, is constrained to 0.90e0.85 Ma (Martin, 2012;Budsky et al., 2015;Gibert and Scott, 2015). The faunal assemblage from Cueva Victoria has been also related to an open environment surrounded by forest and with warm temperatures (Blain, 2015). ...
The Iberian lynx (Lynx pardinus) is a critically endangered felid that, during the last fifty years, has been subject to an intensive conservation program in an attempt to save it from extinction. This species is first recorded at ca. 1.7–1.6 Ma (late Villafranchian, late Early Pleistocene) in NE Iberian Peninsula, roughly coinciding with the large faunal turnover that occurred around the middle to late Villafranchian boundary. Here we describe the largest collection of L. pardinus remains available to date from the Iberian late Early Pleistocene (Epivillafranchian), including localities from the Vallparadís Section (Vallès-Penedès Basin, NE Iberian Peninsula) and Cueva Victoria (Cartagena, SE Iberian Peninsula). The morphology and biometry of the studied material attests to the widespread occurrence of L. pardinus in the Mediterranean coast of the Iberian Peninsula since the latest Early Pleistocene, i.e., about 0.5 million years earlier than it was generally accepted (i.e., at the beginning of the Middle Pleistocene). Based on the features observed in the large sample studied in this paper, we conclude that Lynx spelaeus is a junior synonym of L. pardinus and further propose to assign all the Epivillafranchian and younger fossil lynxes from SW Europe to the extant species L. pardinus. Due to the arrival of the Eurasian lynx (Lynx lynx) into Europe at the beginning of the Late Pleistocene, the attribution of specimens younger than MIS 5e to either this species or L. pardinus solely on morphological grounds has proven equivocal. Here we discuss the main diagnostic features of both species of European lynxes and further review their evolutionary history and paleobiogeography throughout the Pleistocene.
... In the Old World, a late Pliocene and early Pleistocene radiation of rhizodont Mimomys and Mimomys-like species gave rise independently to a number of endemic arvicolinin genera without molar roots, such as Arvicola, Microtus, Iberomys, Stenocranius, Chionomys, Tibericola and Victoriamys (Chaline 1972;Rabeder 1981;Heinrich 1990;Agustí 1991;Agustí et al. 1993;Nadachowski 1991;Rekovets and Nadachowski 1995; Cuenca-Bescós et al. Tesakov et al. 2007;Martin 2012a). The relative positions of the arhizodont genera in Fig. 1 are somewhat speculative. ...
North American arvicolid rodent diversity is compared with global arvicolid diversity in a new phylogenetic model incorporating the possibility that the Arvicolidae as presently constituted is polyphyletic. It is likely that many early Pleistocene arhizodont arvicolinins (e.g. Arvicola, Microtus, Victoriamys) evolved from different Mimomys ancestors. North American phylogenetic patterns indicate successive waves of dispersal from Asia, coupled with endemic radiation of the Plophenacomyinae from an ancestral Promimomys, such as P. mimus.
... mm; W ¼ 1.14e1.54 mm, according to Alcalde et al., 1981;Martin, 2012). ...
In this paper the arvicoline from the Pirro 13 fissure are described (Pirro Nord, Apulia, southwestern Italy). Due to their wide geographical distributions and rapid evolutionary rates, arvicolines are especially useful for biochronological purposes. The Pirro 13 fissure has yielded one arvicoline species: Allophaiomys ruffoi. The presence of this vole species, along with a comparison of our data with those of other Early Pleistocene sites such as Sierra de Atapuerca and Orce, allows us to determine the age of Pirro 13, which is shown to range from 1.3 to 1.6 Ma, making Pirro 13 the most ancient locality with human evidence currently known in Western Europe.
... sylvaticus and Hystrix refossa (Minwer-Barakat et al., 2011;Martínez et al., 2010Martínez et al., , 2014. This association may be correlated with the Allophaiomys chalinei (recently redefined as Victoriamys chalinei by Martin, 2012) biozone defined by Cuenca-Besc os et al. (2010) at Atapuerca Gran Dolina for levels TD3 to TD8a, and ranging in age from the end of the Early Pleistocene (>0.78 Ma) to the beginning of the Middle Pleistocene (around 600 ka) (Falgu eres et al., 1999;Par es and P erez-Gonz alez, 1999;. According to Minwer-Barakat et al. (2011), the absence of V. chalinei in EVT-7 should not be interpreted as a biochronologic indication, but is probably due to the scarcity of the rodent material that has been found at Vallparadís. ...
ESR dating was applied to fossil teeth and optically bleached quartz grain samples from two units of the sequence at Vallparadís (Barcelona, Spain): weighted mean ESR age estimates of 858 ± 87 ka and 849 ± 48 ka were obtained for EVT-7, which includes the archaeological level 10, and EVT-8, respectively. These results are in good agreement with the existing magneto-biostratigraphic framework that constrain these deposits between 780 and 990 ka, and indicate that Vallparadís EVT-7 has a chronology very close to that of Atapuerca Gran Dolina TD-6 (Spain).
... Remarks: The size of the specimens falls within the range of those from the type locality of the species, Cueva Victoria (Alcalde et al., 1981); however, the mean size is larger in Barranc de la Boella (level 2). The studied teeth are slightly smaller than those of V. chalinei from levels TD4 and TD4B of Atapuerca Victoriamys is a recently erected genus (Martin, 2012). Previously V. chalinei was assigned to the genus Allophaiomys. ...
In 2007, remains of Mammuthus meridionalis and 334 lithic artefacts were recovered in level 2 of Barranc de la Boella. The presence of M. meridionalis allows us to assign an age older than 700 ka to this level, which represents one of the few sites of this age with palaeontological remains associated with lithic industry. It also enables us to shed light on questions pertaining to the chronology and the biogeography of the earliest European human settlements. In this paper, the small mammals from the Barranc de la Boella site are described for the first time, with special emphasis on the arvicolines. The presence of Victoriamys chalinei and Mimomys savini allows us to estimate the age of level 2. Level 2 of the Barranc de la Boella site ranges between the top of the Jaramillo subchron (0.99 Ma) and shortly after the Brunhes–Matuyama transition at 0.78 Ma, thus covering a time span of great importance for our knowledge of the earliest human occupations in Western Europe.
... The age of Pirro Nord LFA (?1.5e1.4 Ma) is thus bracketed between Pietrafitta and the Spanish Barranco L eon and Fuente Nueva 3 LFAs, where a more derived vole, M. (Allophaiomys) chalinei (= Victoriamys chalinei in Martin, 2012), is present (Agustí et al., 2010. 4.1.3. ...
This research aims to investigate the relationships between climate change and faunal dynamics in south-west Europe, disentangling the asynchronous and diachronous dispersal bioevents of large mammals across geographical and ecological boundaries, analysing biodiversity and its changes through time. The analysis of local versus regional biological dynamics may shed new light on whether turnovers and ecological and evolutionary changes developed because of global climate changes and related phenomena, or because of intrinsic biological factors. The SW European Quaternary fossil record is particularly suitable for studying the role of climate change at local and regional levels because of the complex physiographic and climatic heterogeneity of the study area, the presence of important geographical/ecological barriers and the complex history of invasions of species of varying geographical origin and provenance. The data base consists of taxonomically revised lists of large mammal species from selected SW European local faunal assemblages ranging in age from the Early to the late Middle Pleistocene (middle Villafranchian to early Aurelian European Land Mammal Ages). The new biochronological scheme proposed here allows for the comparison of local turnovers and biodiversity trends, yielding a better understanding of the action of geographical/ecological barriers that either prevented the range of some taxa from reaching some regions or caused delays in the dispersal of a taxon in some territories. The results obtained provide evidence that major environmental perturbations, triggering dispersal events and removing keystone species, modified the structure of the pre-existing mammalian faunas, merging previously independently-evolved taxa into new palaeo-communities. The coupled action of climatic changes and internal biotic dynamics thus caused the Quaternary SW European faunal complexes to significantly restructure. Diachroneity in local turnover across the study area probably relates to differences in local dynamic patterns of competition/coevolution, although different manifestations of global climate changes in different geographic settings would have contributed to the scale of local bioevents.
... The Jaramillo has not been identified in the Orce area and Iberomys huescarensis has not been recovered from sediments in the Baza basin during or prior to the Jaramillo. On a more trivial taxonomic note, "Allophaiomys" chalinei was recently allocated to the new genus Victoriamys (Martin, 2012). ...
Recent attempts to estimate the age of deposition of European fossil localities using mathematical equations derived from size change of the first lower molar in arvicolid rodent lineages as a function of time prompted an assessment of the value of this approach. The accuracy of “vole clocks” depends on accurate dating of a fossil system and establishment of a directional size change pattern through time in a given species from the dated system. Results of this review suggest that vole clocks have limited value for biochronology. In addition to several methodological and statistical problems with published studies, vole clocks in general are untenable because paleontological systems cannot resolve dating to the level of accuracy necessary to construct an accurate equation, size and shape change is never monotonic (constant velocity) in lineages, and size commonly reverses direction in lineages on all time scales.
... In several previous articles some of these groups have been studied in detail in a regional and transregional context (e.g., Ayarzaguena and Lopez-Martínez, 1976;Agustí, 1992;López-García et al., 2011;Petruso et al., 2011). Victoriamys, erected as an independent genus by Martin (2012) is grouped here within Allophaiomys since it includes primitive characters (low A/L values, primitive ACC pattern) and might show parallel evolution to other Microtus lineages. The most notable change through time observed in most of the Microtus lineages is an increase in complexity of the anteroconid complex in the m1 (A/L ratio). ...
Resumen En Cueva Victoria se han encontrado dos tipos de primates, el papionino Theropithecus oswa/di y el hominino Horno sp. El primero se definió por un molar inferior deciduo; recientemente, el descubrimiento de 4 nuevos dientes ha per-mitido una clasificación más ajustada como Theropithecus T oswa/di /eakeyi. Los restos de homininos hallados hasta ahora son una falange intermedia del V dedo de la mano derecha (CV-O) y dos fragmentos humerales (CV-1 y CV-2), lo que hace muy difícil su atribución específica, clasificándose como Horno sp. Respecto a la hipótesis publicada en 2005 de que la falange pertenezca en realidad a Theropithecus, se publicaron en 2008 una réplica y una contra réplica y en este trabajo se responde a la contrarréplica, ratificando la asignación a Horno sp. La presencia de estos primates en Cueva Victoria hace que este yacimiento sea fundamental para la discusión de las dispersiones de primates africanos a Europa, y refuerza la hipótesis de que probablemente la vía de entrada al continente europeo se produjo a través del Estrecho de Gibraltar. Palabras clave Theropithecus oswaldi, Horno sp., Dispersión, Falange, Pleistoceno inferior. Summary At Cueva Victoria, two types of primates have been found, the papionin Theropithecus oswaldi and the hominin Horno sp. The former was initially identified by a deciduous molar; recently, the discovery of 4 new teeth has allowed a more accurate classification as T T oswaldi leakeyi. Of the second, only one intermediate phalanx of the V finger of the right hand and two humeral fragments have been found up to now, which makes the specific attribution difficult, and they are classified as Horno sp. A claim, published in 2005, stating that the phalanx was of Theropithecus, was followed by a reply and a response in 2008. Here we reply this latter answer and we reaffirm the assignment to Homo sp. The presence of these primates in Cueva Victoria makes this site essential in the discussion of the dispersal of African primates into Europe, and reinforces the hypothesis of the Strait of Gibraltar as a probable route out of Africa.
In this paper we review the collection of fossil carnivores from the late Early Pleistocene site of Cueva Victoria (south-eastern
Iberian Peninsula) including the families Ursidae, Hyaenidae and Felidae. The studied material includes the species
Ursus deningeri, Pachycrocuta brevirostris, Homotherium latidens, Megantereon sp., Panthera gombaszoegensis,
Puma pardoides and Lynx pardinus. The former species are characteristic of the late Early Pleistocene assemblages of
the Mediterranean Europe corresponding to the Epivillafranchian biochron with an approximate age of 1.1-0.78 Ma.
Taking into account the known first and last appearances of the identified species the chronology of the site can be
restricted to ca. 1.0 Ma.
At Cueva Victoria, two types of primates have been found, the papionin Theropithecus oswaldi and the hominin Homo sp. The former was initially identified by a deciduous molar; recently, the discovery of 4 new teeth has allowed a more accurate classification as T. T. oswaldi leakeyi. Of the second, only one intermediate phalanx of the V finger of the right hand and two humeral fragments have been found up to now, which makes the specific attribution difficult, and they are classified as Homo sp. A claim, published in 2005, stating that the phalanx was of Theropithecus, was followed by a reply and a response in 2008. Here we reply this latter answer and we reaffirm the assignment to Homo sp. The presence of these primates in Cueva Victoria makes this site essential in the discussion of the dispersal of African primates into Europe, and reinforces the hypothesis of the Strait of Gibraltar as a probable route out of Africa.
Cueva Victoria is a karst-infilling site with early Pleistocene vertebrate fossil remains. It was discovered in the first half of the XXth century by mining explotation. The fossil record of Cueva Victoria is exceptional because of its large biodiversity and because it is the only locality in Europe where the African primate Theropithecus oswaldi occurs. Here, we summarize the history of paleontological research and excavation campaigns. We provide a catalogue of the different locations in the site, and a reference list of the publications on this site.
In the Early Pleistocene locality of Cueva Victoria, the insectivores are represented by several fossil teeth, mandibles and skull fragments. In the present work, we describe and provide measurements of these fossils for the first time. Insectivores show a rather low taxonomic diversity
in Cueva Victoria, with only two different species. Considering the environmental requirements for their extant relatives, the presence of only one hedgehog (Erinaceus cf. praeglacialis Brunner, 1933), and one shrew (Crocidura kornfeldi Kormos, 1934) points to the existence
of dry and warm conditions in the surrounding area of Cueva Victoria, unfavourable to other contemporary species of insectivores.
In this paper are described the arvicolines from El Chaparral site (Villaluenga del Rosario, C´ adiz, Southwestern
Spain). Due to their wide geographical distributions and rapid evolutionary rates, arvicolines are especially useful
for biostratigraphical purposes. The stratigraphic layers of El Chaparral have yielded arvicolines remains of the genera Allophaiomys,
Iberomys, Pliomys, and Terricola. The presence of these rodents together the other vertebrates recovered allows
to determinate the age of the El Chaparral site and compare our data with other Iberian Pleistocene sites, such as of the
Sierra de Atapuerca. El Chaparral site ranges between the Jaramillo Subchron (1.07-0.99 Ma) and shortly after the Brunhes-
Matuyama transition at 0.78 Ma covering a important time span for the knowledge of the earliest humans occupations in
western Europe.
INTRODUCTION The Arvicolidae is a cosmopolitan, mostly Holarctic, family of rodents that first appear in the fossil record during the late Miocene. The earliest records of this group may be from North America, although it is not likely that the family originated here. The arvicolids dispersed and explosively radiated at the higher latitudes during the late Tertiary and are represented throughout Eurasia and North America from that time onwards. Depending on the classification, there are approximately 10 extant genera in North America (see Appendix to this chapter). They range in size from less than 20 g (e.g., Clethrionomys gapperi) to over 1 kg (Ondatra zibethicus). While most modern Microtus are pastoral, found in relatively open, prairie or tundra environments, some North American Microtus and other genera can be found in forested (e.g., M. pinetorum, Clethrionomys, Phenacomys), bog (e.g., Synaptomys), and aquatic (e.g., Ondatra, Neofiber) habitats. Microtus, in particular, has very high reproductive rates in comparison with rodents of comparable size in other families, and the oscillating population cycles of arvicolids and their predators are well known. In many parts of the world, arvicolids represent the dominant prey species for carnivorous animals. There is an extensive literature on the systematics of fossil arvicolids. Summaries have been published by Miller (1896), Hinton (1926), Ognev (1950), Kretzoi (1955a, 1969), Rabeder (1981), Carleton and Musser (1984), Chaline (1987), Repenning, Fejfar, and Heinrich (1990), and Gromov and Polyakov (1992), and additional taxonomic changes have been made that apply to North American arvicolids in a variety of publications focused on specific groups.
Resumen
En este trabajo se describe la historia geológica de la cavidad, la formación de la brecha fosilífera, la edad de la fauna y la naturaleza humana de la falange CV-0 comparándola con las de Theropithecus.
Summary
This work describes the geological history of Cueva Victoria including the formation of the fossiliferous breccia, and the age of the fauna. This work also compares a phalanx classified as Homo with Theropitecus.
In the Early Pleistocene Red Lower Unit of the Sima del Elefante site (Sierra de Atapuerca karst complex, Burgos, Spain), levels TE9–TE13, dental and mandibular remains of an arvicoline are referred to as the new species Arvicola jacobaeus sp. nov. The new species has medium−sized hypselodont molars, with abundant cementum in the re−entrant folds, and thick enamel band with differentiation of the Mimomys−type. The occlusal morphology of M3 is simple. The dental morphology of the new species resembles that of Arvicola sapidus, though smaller. It is more derived, in size and morphology than the Middle Pleistocene species Arvicola mosbachensis. The morphologic affinities among Arvicola jacobaeus, Arvicola terrestris, and A. sapidus suggest a common ancestry. A preliminary phylogenetic analysis corroborates that Mimomys savini is the sister group of the Arvicola clade.
Se estudian los Arvicolidae (Rodentia, Mammalia) de los niveles inferiores de Gran Dolina (TD3, TD4 y TD4B), relleno cárstico de cerca de 18 metros de potencia que abarca parte del Pleistoceno Inferior y parte del Pleistoceno Medio, y que se encuentra dentro del complejo de yacimientos de Atapuerca (Burgos, España). Estos yacimientos son conocidos principalmente por el hallazgo de la mayor concentración de restos humanos del Pleistoceno Medio, con un estado de conservación excepcional. Además, la mayor parte de los yacimientos contienen una fauna diversa, abundante en prácticamente todos los puntos muestreados. Los roedores son el grupo más abundante y los arvicólidos son los más representados. En TD3, TD4 y TD4B se han reconocido los siguientes taxones de arvicólidos: lberomys sp., Stenocranius gregaloides, Terrícola arvalidens, Allophaiomys chalinei, Mimomys savini y Pliomys episcopalis. Estos taxones caracterizan en el Pleistoceno lnferior europeo el Bihariense tardío (Cromeriense en sentido amplio). La asociación de las especies de arvicólidos caracteriza un clima relativamente cálido y húmedo, en un contexto general de un intervalo interglaciar.
The analysis of the distribution of small mammals in the Italian mainland during the Plio-Pleistocene and their immigration in the Peninsula indicates the presence of many species of orien-tal European origin, a few iberoccitanic elements and some emdemic species. The Italian penin-sula belongs to a western Mediterranean bioprovince. The north-eastern region of Italy is a transi-tional biogeographical zone between this province and the central European and Balcanic areas.
In this paper, the rodents from the composite section of Torrent de Vallparadís (Terrassa, northeastern Spain) are described, with particular emphasis on the arvicolines. Due to their wide geographical distributions and rapid evolutionary rates, arvicolines are especially useful for biostratigraphical purposes. Eight stratigraphic layers have yielded rodent remains, including representatives of the genera Mimomys, Allophaiomys, Stenocranius, Iberomys, Microtus, Arvicola, Apodemus, Eliomys, and Hystrix. The presence of different rodent species, together with the available magnetostratigraphic data, allows a precise determination of the age of each layer and a detailed correlation with other Spanish Pleistocene sites, particularly those of the Sierra de Atapuerca. The complete studied sequence ranges from the pre-Jaramillo Biharian (1.4–1.2 Ma) to the early Toringian (less than 0.6 Ma.), constituting one of the most complete sequences of the Spanish Pleistocene and covering a time span of especial relevance in relation to the earliest human dispersal into western Europe.
Variation in enamel microhistology (the schmelzmuster) and other features were examined in adult Ogmodontomys poaphagus molars from the Meade Basin of southwestern Kansas. Measurements of total enamel thickness and thickness of tangential enamel
were taken on individual triangles of first upper molar samples from the superposed Ripley B, Hornet and Rexroad 2A local
faunas. Although thickness of total enamel layers and the tangential zone were not statistically different between triangles
and sites, the quality of tangential enamel on the trailing edges increased, from incipient tangential and well-developed
tangential enamel (Ripley B and Hornet) to solely well-developed tangential enamel (Rexroad 2A). The fundamental schmelzmuster
of O. poaphagus is radial enamel on the leading edges and radial plus incipient or well-developed tangential enamel on the trailing edges.
These results support the assertion that the schmelzmuster is a consistent and genetically controlled character mosaic, and
confirm suggestions in the literature that Ogmodontomys is distinct from Mimomys. Paired sample Student’s t-tests of trailing/leading edge samples for each triangle were used to test for differentiation.
Although the molars of Ogmodontomys poaphagus appear undifferentiated under the light microscope, and are indeed separable from the distinctly negatively differentiated
molars of Mimomys by this technique, our results show that the M1s of O. poaphagus are slightly negatively differentiated, with various triangles being either undifferentiated or negatively differentiated.
O. sawrockensis has a more primitive schmelzmuster than O. poaphagus, and the presence of discrete (primitive) lamellar enamel on the apices of at least one triangle in 55% of the sample from
Fallen Angel B, combined with the absence of lamellar enamel in molars of O. poaphagus from Fox Canyon and all subsequent Ogmodontomys samples, conclusively demonstrates that the Fallen Angel B population of O. sawrockensis was not ancestral to O. poaphagus in the Meade Basin. A unique mosaic of dental and size features indicates that the early Blancan O. pipecreekensis from Indiana is a sister species to O. sawrockensis and not ancestral to O. poaphagus.
The Orce fossil quarries, in the Baza Basin of southeastern Spain, are a rich source of Early Pleistocene Palaeolithic tools and vertebrate remains. Geologic fieldwork during the last decade has placed these fossiliferous strata within the context of a thick Neogene continental sequence. Detailed lithostratigraphic and magnetostratigraphic results indicate that at least the upper 60 m of this sequence are of Early Pleistocene age. The quarried strata (Venta Micena, Barranco León and Fuentenueva-3) are from a narrow time span (<100 ka) starting before 1.3 Ma. A new, lower excavation at ∼1.5 Ma (Fuentenueva-1 quarry) has a distinctly older fauna. These Orce strata provide a high resolution, Early Pleistocene record of grassland fauna that shows the end of Mammal Neogene fauna (MN17) in the Fuentenueva-1 site (with Gazella borbonica, Equus stenonis) and the beginning of more characteristic Pleistocene fauna in the Venta Micena site (with Hippopotamus antiquus, Equus granatensis, Homo sp.). Thus far, no evidence for human occupation has been found within the earlier Fuentenueva-1 quarry, although many of the same terminal MN17 species have been found with hominids on the other side of Europe at the Dmanisi site (earliest Pleistocene) in the Republic of Georgia.
Thousands of fossils of small vertebrates are recorded in one of the most complete stratigraphic sequences of the continental European Pleistocene, in the Sierra de Atapuerca. This sequence, with a few interruptions, spans the last 1.5 million years. The stratigraphic series is recorded in six karstic cavities in the Sierra de Atapuerca. From the oldest record to the most modern one, these are the Sima del Elefante, Gran Dolina, Galería–Zarpazos, Sima de los Huesos, El Portalón and El Mirador. The faunal successions of continental microvertebrates have made it possible to divide the Atapuerca sequence into 7 faunal units (Atapuerca Faunal Units, ATA FUs), which are based on the first and last local appearances of taxa (LO, HO). The Spanish Quaternary small vertebrate faunas are grouped into 5 distinct biozones, from the oldest to the most recent: Allophaiomys pliocaenicus, Allophaiomys lavocati, Allophaiomys chalinei, Iberomys brecciensis and Iberomys cabrerae. These are calibrated with radiometric and palaeomagnetic dating, allowing the proposal of a continental biochronological scale for the Pleistocene of the Iberian Peninsula.
The continental vertebrate fauna of Cúllar Baza-1 (Granada, Spain) occur immediately above the Matuyama/Brunhes polarity boundary, and therefore represent an initial record for the Middle Pleistocene. All polarity zones for the Late Pliocene through Middle Pleistocene are found in an 80 m section, which includes this fossil quarry. The existing collection of abundant remains of micro-mammals, macro-mammals and lithic artifacts (indicating human activity) can now be assigned an earliest Middle Pleistocene age of 0.75 Ma. Another section, 25 km across the Neogene Baza Basin, also has the Matuyama/Brunhes polarity boundary. However, in this case the Huéscar-1 fossil quarry and the Puerto Lobo sites are both below the polarity boundary, thus representing a record of the late Early Pleistocene at ∼0.9 Ma. Differences in micro-mammal species across these Matuyama/Brunhes boundaries are significant and justify an approximately coincident biostratigraphic boundary between the Biharian and Toringian stages.
A new species of Allophaiomys. A. chalinei, is described from the Lower Pleistocene of the Victoria cave (Murcia. Spain). This species is characterized by his great size, with confluent T 4 and T 5 and it has been also found at the lower levels of Atapuerca (Burgos) and Venta Micena (Granada). Peer reviewed
"Publikation des Sonderforschungsbereichs 53, 'Paläontologie mit besonderer Berücksichtigung der Palökologie' an der Universität Tübingen: Konstruktionsmorphologie Nr. 120." Habilitationsschrift--Frankfurt am Main. Summary also in English. Includes bibliographical references (p. 91-94).
The karstic filling of Cueva Victoria in southeastern Spain, dated from the latest early Pleistocene (ca. 1.1 Ma), is famous for providing primate fossil remains (Theropithecus) of typical African origin, in the general controversy on the antiquity of the first hominid settlements in Western Europe and their possible entrance into Europe through the Strait of Gibraltar. Cueva Victoria has also furnished the following fauna of anurans and squamate reptiles: cf. Pelodytes sp. (Pelodytidae), Bufo cf. B. bufo (Bufonidae), Blanus cinereus (Blanidae), Tarentola sp. (Geckonidae), Chalcides cf. Ch. bedriagai (Scincidae), Timon cf. T. lepidus and indeterminate small lacertids (Lacertidae), Natrix maura, Coronella girondica, Rhinechis scalaris and Malpolon cf. M. monspessulanus (Colubridae). This faunal association seems to suggest a mean annual temperature slightly fresher than nowadays (approximately 1°C less than at present in the area), with cooler winters but warmer summers and above all higher mean annual precipitations (+ 400 mm). The landscape may correspond to an open forest environment of a Mediterranean type, with some still water points.
Stone tools are durable reminders of the activities, skills and customs of early humans, and have distinctive morphologies that reflect the development of technological skills during the Pleistocene epoch. In Africa, large cutting tools (hand-axes and bifacial chopping tools) became part of Palaeolithic technology during the Early Pleistocene ( approximately 1.5 Myr ago). However, in Europe this change had not been documented until the Middle Pleistocene (<0.5 Myr ago). Here we report dates for two western Mediterranean hand-axe sites that are nearly twice the age of the supposed earliest Acheulian in western Europe. Palaeomagnetic analysis of these two sites in southeastern Spain found reverse polarity magnetozones, showing that hand-axes were already in Europe as early as 0.9 Myr ago. This expanded antiquity for European hand-axe culture supports a wide geographic distribution of Palaeolithic bifacial technology outside of Africa during the Early Pleistocene.
Arvicolids are rodents which have molars with a morphology formed by a sequence of enamel folds similar to the curve y = sin f(x). The morphology of the crown of the first lower molar (m1) of living species of Arvicola (large voles) is identified with six criteria, irrespective of tooth size. When rootless arvicolid fossil communities are analysed, it can be seen that the m1 morphology of Arvicola is present in those communities represented by specimens of small size at the beginning of the Quaternary. Before this data was known, the presence of Arvicola communities could only be detected in the second half of the Quaternary, when the specimens were comparable to the large size characterizing living species. The existence of communities of small-sized Arvicola at the beginning of the Quaternary implies that the m1 of Arvicola undergoes a continuous and accelerated growth throughout the entire Quaternary, which allows representatives the genus to be used as a chronological tool. These data mean that it is necessary to change the concept of the genus Allophaiomys and to formulate a new classification to reflect evolutionary relationships of quaternary arvicolids.
Simpson (1944, 1953) distinguished two major features of evolution, phyletic evolution or anagenesis (Rensch, 1954) on the one hand, and splitting or cladogenesis on the other. According to this concept, the two modes lead to the formation of new species either by gradual changes in lineages or by splitting of an ancestral species into two or more “sister” species.
Students of fossil arvicoline rodents are challenged to conform to a higher level of modern statistical and phylogenetic analysis. The terminology of the occlusal surface of arvicoline teeth of van der Meulen (1973, 1978) is proposed as the standard for future research. The current taxonomy of North American fossil voles referred to Allophaiomys, Pedomys, and Pitymys is reviewed and critiqued. These taxa are combined in the genus Microtus. Phaiomys (= Allophaiomys; = Orthriomys) and Pitymys (= Pedomys; = Herpetomys) are recognized as subgenera. A Pleistocene replacement chronology for Microtus dental morphotypes is presented, and a cladogram based on dental character states is constructed for North American Microtus. Microtus meadensis is considered to be an Old World immigrant and not closely related to M. quasiater. The Late Pleistocene Microtus reported by Repenning (1983) from El Tajo de Texquixquiac, Mexico as M. meadensis is tentatively referred to M. quasiater. Microtus quasiater is further considered to be distinct from M. pinetorum. Microtus (Pitymys) mcnowni is not a valid species and is tentatively combined under M. ochrogaster. Microtus (Pitymys) aratai is a valid species. Pleistocene specimens reported by Repenning (1983) from Centerville Beach, California now lack proper species reference. Microtus oaxacensis and M. guatemalensis, here classified in Pitymys, demonstrate both confluent T4–5 and closed T4–5 on m1, thus providing a possible evolutionary scenario deriving some species of the subgenus Microtus from Pitymys. This scenario is contrasted with that of Guthrie (1965), in which Microtus is descended from a species such as M. paroperarius, which has predominantly four closed triangles on m1.
Microtus (Pedomys) australis, sp. nov., is described from the Haile XVIA local fauna (1. f.) of Florida (ca. 1.3 to 1.6 Ma [million years ago]) and the Vera 1. f. of Texas (ca. 0.6 Ma). Microtus australis shares a number of plesiomorphic features with M. pliocaenicus and is related to both the subgenus Phaiomys, from which it evolved, and the subgenus Pedomys, to which it is referred. Microtus (Pedomys) guildayi is subsumed under M. pliocaenicus. The sample on which this taxon was based, from Cumberland Cave, is referred to as M. pliocaenicus/guildayi, a slightly modified form of the informal taxonomic usage proposed by Krishtalka and Stucky (1985). Microtus llanensis is synonymized with M. ochrogaster and is recognized as M. o./llanensis. It is considered an intermediate in the phyletic sequence from M. p. /guildayi to modern M. ochrogaster. Specimens reported by Repenning and Grady (1988) from the middle Pleistocene Hamilton Cave 1. f. as Pitymys hibbardi do not belong to that taxon and may represent a new species. Pitymys hibbardi is treated as a large form in a phyletic sequence leading to extant M. pinetorum. The Pleistocene replacement chronology of “pitymyinine” voles in Florida is as follows: M. australis-M. aratai-M. pinetorum/hibbardi-M. pinetorum/parvulus. A cladistic analysis of dental characters for select subgenera of Microtus is presented, and a revised classification is offered. Orthriomys, Pitymys, and Pedomys are recognized as endemic North American subgenera of Microtus, while representatives of Phaiomys (M. pliocaenicus) and Terricola (M. meadensis) are considered to be Old World immigrants. The relationships of M. quasiater, M. oaxacensis, M. guatemalensis, and M. aratai remain obscure.
The representation of Allophaiomys in Southern Europe has increased considerably thanks to the intensive samplings carried out in recent years in the Guadix-Baza basin and other sites in Spain (Casablanca and Cueva Victoria fissure infillings). In addition to the type-species, Microtus (Allophaiomys) pliocaenicus, these finds have provided information on two additional species: M. (Allophaiomys) chalineiAlcalde, Agusti & Villalta and M. (Allophaiomys) vandermeuleni n. sp. (type-locality: Barranco de los Conejos in the Guadix-Baza basin, Southern Spain). The latter species differs from other Allophaiomys representatives by its large size and its very deep LRA-3 (lingual reentrant angle 3), which contrasts with the primitive shape of the anteroconid complex. The evolutionary relationships of the different Allophaiomys species are analyzed by two different approaches. First, the phenetic view of the early evolution of Microtus in Europe is based exclusively on the A, B and C parameters commonly used in the systematics of this genus. A second phylogenetic approach is attempted also taking into account other morphological characters. As a consequence of the results shown by the mostly parsimonious cladogram, the subdivision of the Allophaiomys stock into two assemblages, the first one dealing with the “European” Pitymys (or Terricola, according to Brunet-Lecomte, 1988) and a second one dealing with the Chionomys group, is proposed. Moreover, this divergence can be traced back to the very early Pleistocene, instead of the middle Pleistocene age usually accepted for this splitting process. Thus, M. (A.) vandermeuleni shares a common ancestor with the Chionomys group, while M. (A.) chalinei is situated close to the origin of M. (A.) nutiensis. On the other hand, M. (A.) burgondiae is proposed as the sisterspecies of M (Allophaiomys) jordanicus (Haas) (not Arvicola). Therefore, a middle Pleistocene age is proposed for the site of Ubeidiya.
To the east of the Sea of Marmara, the North Anatolian fault (NAF) branches into two strands, namely the northern and the southern strands. The Adapazarı pull-apart basin is located in the overlapping zone of the Dokurcun and the İzmit–Adapazarı segments of the northern strand. The combined temporal ranges of the arvicolids from the Karapürçek formation (the first unit of the basin fill), deposited in the primary morphology of the Adapazarı pull-apart basin, cover the latest Villanyian (latest Pliocene) and the Biharian (Early Pleistocene) time interval. The Değirmendere fauna collected from the lowermost sediments of this formation suggests that the Adapazarı pull-apart basin started to form in the latest Pliocene. This, in turn, suggests that the dextral movement along the northern strand of the NAF commenced during the latest Pliocene. A new species, Tibericola sakaryaensis is also described.
Pleistocene exposures located near Cape Deceit on the south shore of Kotzebue Sound (Alaska) contain a complex sequence of organic sediments which have yielded fossil vertebrates of taxonomic, evolutionary, and zoogeographic significance. Cape Deceit Local Fauna from the Cape Deceit Formation, lowermost of three major stratigraphic units recognized at the site, contains representatives of the genera Canis, Rangifer, Cervus, Ochotona, Lemmus, Microtus, Pliomys, and a newly described genus and species, Predicrostonyx hopkinsi-predecessor of the extant genus Dicrostonyx. In addition to P. hopkinsi, new species of Ochotona, Pliomys, and Microtus are described from the Cape Deceit Local Fauna. the stage of evolution of Microtus deceitensis sp. n. and P. hopkinsi sp. n. indicate that the Cape Deceit Formation is at least of pre-Cromerian age. This assumption is substantiated by the stratigraphy of the overlying Inmachuk and Deering formations, fossils from these units (including Dicrostonyx torquatus and D. henseli), and C-14 dates. The Cape Deceit Local Fauna, therefore, contains the earliest North American record of several of the included genera in addition to the only North American record of Pliomys.
The Barranco de Orce (BO) section in the Baza basin (SE Spain) exposes several fossiliferous layers (O-1 to O-7) with Plio–Pleistocene micro- and macromammals. Biostratigraphic and magnetostratigraphic data from this and other sections in the basin have been extensively used to calibrate the Plio–Pleistocene chronology based on mammal biozonations. Because of its stratigraphic and geographic proximity, the BO section has also been used to date the paleontological and archeological sites of Barranco León, Fuentenueva-3 and Venta Micena. This study shows that the BO section crosses a mega-landslide that produces partial repetitions of the sedimentary sequence. The seven fossiliferous layers are actually the repetition of only two (O-6 and O-7) which are found in situ in the upper part of the ravine. New paleomagnetic results demonstrate the presence of Reverse magnetization throughout this section, contradicting the Normal event previously reported and assigned to the ‘Olduvai’ subchron (C2n). Published and new magnetostratigraphic data show that all archeological and paleontological sites in the Orce area are within a Reverse magnetochron, presumably C1r.2r (late Matuyama). The use of BO in the magnetobiostratigraphical calibration of the Pliocene/Pleistocene boundary for western Europe is not advised.
Eleven sections have been sampled for magnetostratigraphy in the fossil mammal-productive late Neogene basins of eastern Spain. The characteristic direction of magnetization was determined through progressive thermal and alternating field demagnetization. It was possible to correlate the magnetozones from the various sections to the GPTS as well as to the fossil MN zones within the sediment. The MN zone boundaries based on this correlation are as follows: MN 11/12 ⋍ 8.1 my; MN 12/13 ⋍ 6 my; MN 13/14 ⋍ 4.9 my; MN 14/15 = 4.18 my; MN 15/16 = 3.58 my. New fossil lists are presented for the Fuente del Viso and Loma del Castillo faunas in the Cabriel Basin.
Thesis (M.S.)--Murray State University, 1998. Includes bibliographical references (leaves [74]-79) and abstract.
Continental faunal units on the Plio-Pleistocene of France. Memoria della Societa Geologica Italiana 31
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On the vertebrate fauna of the Lower Pleistocene site Ubeidiya
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Variation and enamel differentiation of Microtus molars from the Java local fauna
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Mesial aspect of left mandibles in Victoriamys chalinei nov. comb. from Cueva Victoria
- Fig
Fig. 1. Mesial aspect of left mandibles in Victoriamys chalinei nov. comb. from Cueva Victoria (A, CV-MC-848;
Arvicola sp. -Ruiz Bustos, table 1 Microtus (Allophaiomys) chalinei -Kotsakis Holotype: Vi 15, R m1, in collections of the Section of Biostratigraphy of the Institute
- Allophaiomys Alcalde
- Agustí
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Allophaiomys chalinei -Alcalde, Agustí and Villalta, p. 203, fig. 1. 1999. Arvicola sp. -Ruiz Bustos, table 1. 2003. Microtus (Allophaiomys) chalinei -Kotsakis, Abbazzi, Angelone, Argenti, Barisone, Fanfani, Marcolini and Masini, p. 319, fig. 3. Holotype: Vi 15, R m1, in collections of the Section of Biostratigraphy of the Institute ''Jaime Almerí'' (C.S.I.C.).
fig. 1, pl. 1-1) and Agustí (1992: pl. 1, fig. H). Locality and age: Cueva Victoria
- Alcalde
Paratypes: additional specimens from Cueva Victoria illustrat-ed by Alcalde et al. (1981: fig. 1, pl. 1-1) and Agustí (1992: pl. 1, fig. H). Locality and age: Cueva Victoria; Early Pleistocene (0.98-0.90 Ma).
Amphibians and squamate reptiles from the latest early Pleistocene of Cueva Victoria
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latest early Pleistocene of Cueva Victoria (Murcia, southeastern Spain, SW
Mediterranean): paleobiogeographic and paleoclimatic interpretations. Geologica Acta 6, 345-361.
Mammalia) de los niveles inferiores de Gran Dolina (Pleistoceno inferior
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Burgos, Españ a). Revista Españ ola de Paleontología 10, 202-218.