Middle Miocene tragulid remains from Abocador de Can Mata: The earliest record of Dorcatherium naui from Western Europe

In the past, the tragulid species Dorcatherium crassum has been considered a synonym of Dorcatherium naui, although they are currently considered distinct species. The former has an exclusively Aragonian (MN4 to MN6) stratigraphic range, whereas D. naui is considered a typically Vallesian species (MN9 and MN10) that survives until the Turolian (MN11). There are only a few previous citations of Dorcatherium from the European latest Aragonian (MN7+8), and only that of Przeworno (Poland) can be securely attributed to D. naui. Here we describe dentognathic and postcranial remains of Dorcatherium from several localities from the stratigraphic series of the Abocador de Can Mata (ACM), in els Hostalets de Pierola (Vallès-Penedès Basin, Catalonia, Spain). These Dorcatherium remains, from both the lower (ca. 11.9-12.5 Ma, MN7) and upper (ca. 11.6 Ma, MN8) portions of the ACM series, constitute the earliest record of D. naui from the Iberian Peninsula and Western Europe. They most closely resemble D. naui from the Vallesian, but still share several features with D. crassum. On the basis of the unique combination of features displayed by the ACM material, a new subspecies, D. naui meini is erected. Its distinct status from the nominotypical subspecies, D. naui naui, is reinforced by the chronological differences between the two taxa. The morphological similarities between Dorcatherium and the extant water chevrotain (Hyemoschus aquaticus) tentatively suggest the presence of forested and humid paleoenvironments in this area of the Vallès-Penedès Basin during the latest Aragonian. Nevertheless, Dorcatherium is very rare in ACM and displays a very patchy chronological distribution. Together with the similarly spotty distribution of both primates and castorids along the ACM series, this suggests that very humid conditions might have been only intermittently present in this area throughout the latest Aragonian.     

First record of Latonia gigantea (Anura,Alytidae) from the Iberian Peninsula

The single extant species of the anuran genus Latonia lives in Israel, but in the fossil record the genus is known mainly from Europe, spanning from the Oligocene to the early Pleistocene. Here we describe new remains of Latonia from the early to late Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula), coming from the following localities: Sant Mamet (MN4), Sant Quirze and Trinxera del Ferrocarril (MN7+8), and Castell de Barberà, Can Poncic 1 and Can Llobateres 1 (MN9). Fossils from the late Aragonian and early Vallesian are attributed to Latonia gigantea mainly because of the morphology of the ornamentation that covers the maxillae. In turn, an ilium from Sant Mamet is not diagnostic at the specific level and is assigned only to the genus Latonia. The newly reported remains represent the first record of L. gigantea in the Iberian Peninsula, where Latonia was previously known by a single report of Latonia cf. ragei from Navarrete del Río (MN2) and remains from other localities unassigned to species. Moreover, the Vallès-Penedès remains represent one of the southernmost records of the species known thus far. The presence of Latonia in these localities confirms the humid and warm environment suggested by the recorded mammal fauna.     

The scimitar-toothed cat Machairodus aphanistus (Carnivora: Felidae) in the Vallès-Penedès Basin (NE Iberian Peninsula)

Here, we revise all the published and unpublished scimitar-toothed cat remains from the Vallès-Penedès Basin (NE Iberian Peninsula), in order to confirm their taxonomic attribution to Machairodus aphanistus as well as to provide more precise information about its chronological distribution in this basin. The studied material (including dentognathic as well as postcranial remains) comes from the following localities: Can Mata indeterminate (late MN7+8 or MN9), Creu Conill 22 (MN9), Can Poncic 1 (MN9), Can Llobateres 1 (MN9), Santiga (MN9), La Tarumba 1 (MN10), Viladecavalls (MN10), Ronda Oest Sabadell ROS-D3 (MN10), and Torrent de Febulines (MN10). Most of the studied material fits well with the morphologic and metrical features characteristic of the Vallesian species M. aphanistus, with the exception of the remains from Creu Conill 22 (an undescribed partial P4 formerly attributed to this taxon), which belongs in fact to a medium-sized hyaenid. From a biostratigraphic viewpoint, the removal of the Creu Conill material from the hypodigm of M. aphanistus has important implications, because this locality (11.1 Ma) was considered to record the first appearance datum of this taxon in the Vallès-Penedès Basin. However, the report of a previously unpublished talus from Can Mata (late MN7+8 or MN9) indicates that this taxon was present in this basin at least by the earliest Vallesian. Therefore, our results indicate that the first appearance datum of Machairodus in the Vallès-Penedès Basin might be somewhat younger than previously assumed, although dating uncertainties for the Can Mata remains preclude a more precise assessment. In contrast, the new mandibular remains from ROS-D3 (MN10) are likely coeval with those from La Tarumba 1, with the last appearance datum of M. aphanistus in the Vallès-Penedès Basin corresponding to Torrent de Febulines (ca. 9.1 Ma). The postcranial material described from various Vallès-Penedès localities further indicates that M. aphanistus displayed less cursorial adaptations than its purported descendant Homotherium.     

A new species of Pliopithecus Gervais, 1849 (Primates: Pliopithecidae) from the Middle Miocene (MN8) of Abocador de Can Mata (els Hostalets de Pierola,Catalonia, Spain)

Pliopithecus (Pliopithecus) canmatensis sp. nov. is described from several Late Aragonian localities from Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès‐Penedès Basin, Catalonia, Spain), spanning from ～11.7 to 11.6 Ma (C5r.3r subchron), and being correlated to the MN8 (reference locality La Grive L3). The ACM remains display a pliopithecine dental morphology with well‐developed pliopithecine triangles on M/2 and M/3. This, together with other occlusal details, negates an attribution to the subgenus Epipliopithecus. Although slightly smaller, the ACM remains are most similar in size to comparable elements of P. piveteaui and P. antiquus. Several occlusal details (such as the greater development of the buccal cingulid in lower molars) and dental proportions (M/3 much longer than M/2), however, indicate greater similarities with P. antiquus from Sansan and La Grive. The ACM remains, however, differ from P. antiquus in dental proportions as well as occlusal morphology of the lower molars (including the less peripheral position of the protoconid and more medial position of the hypoconulid, the more mesial position of the buccal cuspids as compared to the lingual ones, the narrower but distinct mesial fovea, the higher trigonid, and the more extensive buccal cingulid, among others). These differences justify a taxonomic distinction at the species level of the ACM pliopithecid remains with respect to P. antiquus. Previous pliopithecid findings from the Vallès‐Penedès Basin, previously attributed to P. antiquus, are neither attributable to the latter species nor to the newly erected one. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Explaining the end of the hominoid experiment in Europe

The Vallesian Crisis involved the extinction of most of the hominoids that settled successfully in Europe during the middle and early Late Miocene, including Dryopithecus, Ankarapithecus and Graecopithecus. This event has been dated at 9.6 Ma, predating by more than one million years the spread of the C4 grasses and the retreat of forests over large parts of the globe at 7-8 Ma. The finding of macrofloral remains in the Terrassa section (Vallès-Penedès Basin) sheds new light on the nature of vegetational change associated with the hominoid extinction. This section presents an abundant late Vallesian vertebrate fauna and has been accurately dated at 9.2 - 9 Ma by paleomagnetism. Therefore, it provides the best indication of the kind of vegetation that occupied the area after the Vallesian Crisis. It is suggested that the extinction of the late Miocene Western European hominoids was not related to the spread of grasses, but to a significant increase of a floral association dominated by deciduous trees.     

The Miocene mammal record of the Vallès-Penedès Basin (Catalonia)

The land mammal record of the Vallès-Penedès Basin (Catalonia, NE Spain) ranges from the early Miocene (Ramblian) to the late Miocene (Turolian), that is from about 20 to 7 Ma. Here we present an updated review of the mammal succession focusing on biochronology as well as on environmental and faunal changes. Based on faunal similarities with central Europe, we interpret this basin as a transitional zone between the forested environments of northern regions and the more arid landscapes of the inner Iberian Peninsula. The quality of the Vallès-Penedès record and its chronostratigraphic control is clearly better for the late Aragonian and the Vallesian (between 12.6–9.0 Ma), especially for small mammals. Therefore, we analyze small mammal diversity dynamics during this interval. Contrary to previous analyses, which found an abrupt extinction event coinciding with the early/late Vallesian boundary (the Vallesian Crisis), our results show that this pattern is due to uneven sampling. Instead, taxonomic richness slowly decreased since the late Vallesian as a result of a series of extinctions that mostly affected forest-dwelling taxa.     

New remains of Dorcatherium crassum (Artiodactyla: Tragulidae) from the Early Miocene (MN4) of Els Casots (Subirats,Vallès-Penedès Basin)

New tragulid dental remains from the late Early Miocene (16.5–16.3 Ma, MN4) locality of Els Casots (Vallès-Penedès Basin, Catalonia, Spain) are described. This sample fits well, both in size and occlusal morphology, with the material of Dorcatherium crassum from the type locality (Sansan, France; MN6). We therefore attribute the tragulid from Els Casots to this species, also in agreement with its known chronostratigraphic range throughout Europe (MN4–MN6) and the previous records of this species from other MN4 localities of the Vallès-Penedès Basin. The described remains – which include the postcanine lower deciduous and permanent dentition, as well as several upper cheek teeth – constitute the most complete sample of D. crassum from the Iberian Peninsula. The presence of this species at Els Casots is consistent with the lacustrine depositional environment inferred from sedimentological evidence and associated fauna, and further confirms the nearby presence of densely forested environments with a humid climate with low seasonality.     

Taxonomy of subgenus Temnoclemmys Bergounioux, 1958 (Testudines: Geoemydidae: Ptychogasterinae) based on new material from the Vallès-Penedès Basin (NE Iberian Peninsula)

The taxonomy of the freshwater turtle subgenus Ptychogaster (Temnoclemmys) (Testudines, Geoemydidae, Ptychogasterinae) is revised based on new and previously-published material from the Vallès-Penedès Basin (NE Iberian Peninsula). Additional remains of subgenus Ptychogaster, first reported from this basin, are also described. Although the oldest remains of Temnoclemmys are left unassigned, most of the available remains are attributed to its type species, Ptychogaster (Temnoclemmys) batalleri, originally described from the Late Miocene (MN9 and MN10) of this basin. The new material expands the chronostratigraphic range of P. batalleri in the Vallès-Penedès Basin back to the MN7 + 8 and shows that “Testudo” celonica is its junior synonym. Emended diagnoses for Ptychogaster and Temnoclemmys are also provided.     

New Pseudaelurus and Styriofelis remains (Carnivora: Felidae) from the Middle Miocene of Abocador de Can Mata (Vallès-Penedès Basin)

New remains of felid jaws and teeth are described from several localities of the local stratigraphic series of Abocador de Can Mata (ca. 11.9 to 11.6 Ma, Middle Miocene; Vallès-Penedès Basin, Catalonia, Spain). Three different taxa are identified: Styriofelis turnauensis, Pseudaelurus romieviensis and Pseudaelurus quadridentatus. The described remains of P. romieviensis enable extending considerably the chronological range of this species in the Iberian Peninsula, in agreement with its record in the rest of Europe. Moreover, it is shown for the first time that P. romieviensis may possess a p2. The presence of this tooth therefore does not constitue a valid diagnostic feature to distinguish P. romieviensis from P. quadridentatus.     

First partial face and upper dentition of the Middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallès‐Penedès Basin,Catalonia, NE Spain): Taxonomic and phylogenetic implications

A well‐preserved 11.8‐million‐years‐old lower face attributed to the seminal taxon Dryopithecus fontani (Primates, Hominidae) from the Catalan site ACM/C3‐Ae of the Hostalets de Pierola area (Vallès‐Penedès Basin, Catalonia, NE Spain) is described. The new data indicate that D. fontani is distinct at the genus level from Late Miocene European taxa previously attributed to Dryopithecus, which are here reassigned to Hispanopithecus. The new facial specimen also suggests that D. fontani and the Middle Miocene Pierolapithecus catalaunicus are not synonymous. Anatomical and morphometric analyses further indicate that the new specimen shows a combination of lower facial features—hitherto unknown in Miocene hominoids—that resembles the facial pattern of Gorilla, thus providing the first nondental evidence of gorilla‐like lower facial morphology in the fossil record. Considering the current evidence, the gorilla‐like facial pattern of D. fontani is inferred to be derived relative to previously known stem hominids, and might indicate that this taxon is either an early member of the Homininae or, alternatively, a stem hominid convergent with the lower facial pattern of Gorilla. The biogeographic implications of both alternatives are discussed. This new finding in the Hostalets de Pierola section reinforces the importance of this area for understanding the elusive question of the Middle Miocene origin and early radiation of great apes. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Intraspecific variation and evolutionary trends of Alicornops simorrense (Rhinocerotidae) in Spain

The Spanish material of Alicornops simorrense constitutes the best representation of the species in Western Europe. It provides interesting data on the metrical and morphological intraspecific variation of A. simorrense and its evolutionary trends from late Middle Miocene to the early Late Miocene. From late Aragonian to early Vallesian, a slight increase in size is observed, but without clear limits among series. During the late Vallesian, A. simorrense evolved in central Spanish basins into a second species, A. alfambrense, greater in size and with more robust proportions. In the Vallés-Penedés basin, A. simorrense shows a noticeable increase in size, while maintaining its proportions, during the Vallesian. A. simorrense was a very abundant species, an open woodland dweller, with gregarious behaviour, whose extinction was probably linked to the climatic change that took place at the end of the Vallesian and the beginning of the Turolian.     

A New Skull of <Emphasis Type="Italic">Hyaenictis</Emphasis> Gaudry, 1861 (Carnivora,Hyaenidae) Shows Incipient Adaptations to Durophagy

The European Miocene records a wide diversity of hyaenid ecomorphotypes represented by multiple genera. Among these, Hyaenictis Gaudry, 1861, is one of the least known. This genus includes four species from the late Miocene and Pliocene of the Old World, but in Europe Hyaenictis is only represented by two species, recorded by scarce and fragmentary remains: Hyaenictis graeca Gaudry, 1861, from Pikermi (MN12; Greece) and Hyaenictis almerai Villalta Comella and Crusafont Pairó, 1948, from Sant Miquel de Toudell (MN10; Vallès-Penedès Basin, NE Iberia). Here, we describe a new skull of Hyaenictis aff. almerai from the Vallès-Penedès site of Ronda Oest Sabadell Sector D (MN10), representing the most complete European specimen of the genus. In the presence of m2 and virtual lack of m1 metaconid, the described cranium more closely resembles Hyaenictis rather than any other medium- to large-sized European hyaenid. However, the new skull does not fit well with previously known Hyaenictis species, more closely resembling the bone-cracking Adcrocuta Kretzoi, 1938, in the development of premolar accessory cuspids and the possession of relatively broad cheek teeth. These and other features (strong mandibular muscular insertions and enamel microstructure) denote more durophagous adaptations than previously documented in Hyaenictis (considered a cursorial/dog-like hyaena), and favor the inclusion of H. aff. almerai in the transitional bone-cracking hyaenid ecomorphotype.     

Knee function through finite element analysis and the role of Miocene hominoids in our understanding of the origin of antipronograde behaviours: the Pierolapithecus catalaunicus patella as a case study

Although extensive research has been carried out in recent years on the origin and evolution of human bipedalism, a full understanding of this question is far from settled. Miocene hominoids are key to a better understanding of the locomotor types observed in living apes and humans. Pierolapithecus catalaunicus, an extinct stem great ape from the middle Miocene (c. 12.0 Ma) of the Vallès-Penedès Basin (north-eastern Iberian Peninsula), is the first undoubted hominoid with an orthograde (erect) body plan. Its locomotor repertoire included above-branch quadrupedalism and other antipronograde behaviours. Elucidating the adaptive features present in the Pierolapithecus skeleton and its associated biomechanics helps us to better understand the origin of hominoid orthogrady. This work represents a new biomechanical perspective on Pierolapithecus locomotion, by studying its patella and comparing it with those drawn from a large sample of extant anthropoids. This is the first time that the biomechanical patellar performance in living non-human anthropoids and a stem hominid has been studied using finite element analysis (FEA). Differences in stress distribution are found depending on body plan and the presence/absence of a distal apex, probably due to dissimilar biomechanical performances. Pierolapithecus’ biomechanical response mainly resembles that of great apes, suggesting a similar knee joint use in mechanical terms. These results underpin previous studies on Pierolapithecus, favouring the idea that a relevant degree of some antipronograde behaviour may have made up part of its locomotor repertoire. Moreover, our results corroborate the presence of modern great ape-like knee biomechanical performances back in the Miocene.     

Fossil apes from the vallès‐penedès basin

Currently restricted to Southeast Asia and Africa, extant hominoids are the remnants of a group that was much more diverse during the Miocene. Apes initially diversified in Africa during the early Miocene, but by the middle Miocene they extended their geographical range into Eurasia, where they experienced an impressive evolutionary radiation. Understanding the role of Eurasian hominoids in the origin and evolution of the great‐ape‐and‐human clade (Hominidae) is partly hampered by phylogenetic uncertainties, the scarcity and incompleteness of fossil remains, the current restricted diversity of the group, and pervasive homoplasy. Nevertheless, scientific knowledge of the Eurasian hominoid radiation has significantly improved during the last decade. In the case of Western Europe, this has been due to the discovery of new remains from the Vallès‐Penedès Basin (Catalonia, Spain). Here, I review the fossil record of Vallès‐Penedès apes and consider its implications. Although significant disagreements persist among scholars, some important lessons can be learned regarding the evolutionary history of the closest living relatives of humans. © 2012 Wiley Periodicals, Inc.     

New dental remains of Anoiapithecus and the first appearance datum of hominoids in the Iberian Peninsula

New dental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula). These specimens correspond to maxillary fragments with upper teeth from two female individuals from two different localities: left P³–M¹ (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M¹–M² and left P⁴–M² (IPS35027) from ACM/C1-E* (12.3–12.2 Ma). Relative enamel thickness is also computed in the latter individual and re-evaluated in other Middle Miocene hominoids from ACM, in order to better assess their taxonomic affinities. With regard to maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lower-crowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M² occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.     

Taxonomic attribution of the La Grive hominoid teeth

The two hominoid teeth—a central upper incisor (NMB G.a.9.) and an upper molar (FSL 213981)—from the Middle Miocene site of La Grive‐Saint‐Alban (France) have been traditionally attributed to Dryopithecus fontani (Hominidae: Dryopithecinae). However, during the last decade discoveries in the Vallès‐Penedès Basin (Spain) have shown that several hominoid genera were present in Western Europe during the late Middle Miocene. As a result, the attribution of the dryopithecine teeth from La Grive is not as straightforward as previously thought. In fact, similarities with the upper incisor of Pierolapithecus have led to suggestions that either the latter taxon is present at La Grive, or that it is a junior synonym of Dryopithecus. Here, we re‐describe the La Grive teeth and critically revise their taxonomic assignment based on metrical and morphological comparisons with other Middle to Late Miocene hominoids from Europe and Turkey, with particular emphasis on those from the Vallès‐Penedès Basin. Our results suggest that the I¹ differs in several respects from those of Pierolapithecus and Hispanopithecus, so that an attribution to either Dryopithecus or Anoiapithecus (for which this tooth is unknown) seems more likely. The molar, in turn, most likely corresponds to the M¹ of a female individual. Compared to other Middle Miocene taxa, its occlusal morphology enables its distinction from Pierolapithecus, whereas relative crown height agrees well with Dryopithecus. Therefore, based on available evidence, we support the traditional attribution of the La Grive hominoid to D. fontani. Am J Phys Anthropol 151:558–565, 2013. © 2013 Wiley Periodicals, Inc.     

Enamel thickness in the Middle Miocene great apes Anoiapithecus,Pierolapithecus and Dryopithecus

On the basis of industrial computed tomography, relative enamel thickness (RET) is computed in three Middle Miocene (ca 11.9–11.8 Ma) hominoids from Abocador de Can Mata (Vallès-Penedès Basin, Catalonia, Spain): Pierolapithecus catalaunicus from BCV1 and Anoiapithecus brevirostris from C3-Aj, interpreted as stem hominids; and Dryopithecus fontani from C3-Ae of uncertain phylogenetic affinities. Pierolapithecus displays an average RET value of 19.5, Anoiapithecus of 18.6 and Dryopithecus of 10.6. The thick-enamelled condition of Pierolapithecus and Anoiapithecus is also characteristic of afropithecids, including the more derived kenyapithecins from the early Middle Miocene of Eurasia (Griphopithecus and Kenyapithecus). Given the presence of other dentognathic and craniofacial similarities, thick enamel may be interpreted as a symplesiomorphy of the Hominidae (the great ape and human clade), which would have been later independently modified along several lineages. Given the correlation between thick enamel and hard-object feeding, our results suggest that thick enamel might have been the fundamental adaptation that enabled the out-of-Africa dispersal of great-ape ancestors and their subsequent initial radiation throughout Eurasia. The much thinner enamel of Dryopithecus is difficult to interpret given phylogenetic uncertainties, being either a hominine synapomorphy or a convergently developed feature.     

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

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