The Miocene primate Pliobates is a pliopithecoid.

The systematic status of the small-bodied catarrhine primate Pliobates cataloniae, from the Miocene (11.6 Ma) of Spain, is controversial because it displays a mosaic of primitive and derived features compared with extant hominoids (apes and humans). Cladistic analyses have recovered Pliobates as either a stem hominoid or as a pliopithecoid stem catarrhine (i.e., preceding the cercopithecoid-hominoid divergence). Here, we describe additional dental remains of P. cataloniae from another locality that display unambiguous synapomorphies of crouzeliid pliopithecoids. Our cladistic analyses support a close phylogenetic link with poorly-known small crouzeliids from Europe based on (cranio)dental characters but recover pliopithecoids as stem hominoids when postcranial characters are included. We conclude that Pliobates is a derived stem catarrhine that shows postcranial convergences with modern apes in the elbow and wrist joints-thus clarifying pliopithecoid evolution and illustrating the plausibility of independent acquisition of postcranial similarities between hylobatids and hominids.

Paleoenvironmental inferences on the Late Miocene hominoid-bearing site of Can Llobateres (NE Iberian Peninsula): An ecometric approach based on functional dental traits.

Hispanopithecus laietanus from the Late Miocene (9.8 Ma) of Can Llobateres 1 (CLL1; Vallès-Penedès Basin, NE Iberian Peninsula) represents one of the latest occurrences of fossil apes in Western mainland Europe, where they are last recorded at ∼9.5 Ma. The paleoenvironment of CLL1 is thus relevant for understanding the extinction of European hominoids. To refine paleoenvironmental inferences for CLL1, we apply ecometric models based on functional crown type (FCT) variables-a scoring scheme devised to capture macroscopic functional traits of occlusal shape and wear surfaces of herbivorous large mammal molars. Paleotemperature and paleoprecipitation estimates for CLL1 are provided based on published regional regression models linking average FCT of large herbivorous mammal communities to climatic conditions. A mapping to Whittaker's present-day biome classification is also attempted based on these estimates, as well as a case-based reasoning via canonical variate analysis of FCT variables from five relevant biomes. Estimates of mean annual temperature (25 °C) and mean annual precipitation (881 mm) classify CLL1 as a tropical seasonal forest/savanna, only in partial agreement with the canonical variate analysis results, which classify CLL1 as a tropical rainforest with a higher probability. The former biome agrees better with previous inferences derived from fossil plants and mammals, as well as preliminary isotopic data. The misclassification of CLL1 as a tropical forest is attributed to the mixture of forest-adapted taxa with others adapted to more open environments, given that faunal and plant composition indicates the presence of a dense wetland/riparian forest with more open woodlands nearby. The tested FCT ecometric approaches do not provide unambiguous biome classification for CLL1. Nevertheless, our results are consistent with those from other approaches, thus suggesting that FCT variables are potentially useful to investigate paleoenvironmental changes through time and space-including those that led to the extinction of European Miocene apes.

A revised (earliest Vallesian) age for the hominoid-bearing locality of Can Mata 1 based on new magnetostratigraphic and biostratigraphic data from Abocador de Can Mata (Vallès-Penedès Basin, NE Iberian Peninsula).

The Abocador de Can Mata (ACM) composite stratigraphic sequence (els Hostalets de Pierola, Vallès-Penedès Basin, NE Iberian Peninsula) has yielded a diverse primate assemblage from the late Aragonian (Middle to Late Miocene). Detailed litho-, bio-, and magnetostratigraphic control has enabled an accurate dating of these fossil remains. Comparable data, however, were lacking for the nearby locality of Can Mata 1 (CM1), which yielded a dryopithecine canine of a female individual. Given the lack of hipparionin equids and giraffids, CM1 has been correlated to the latest Aragonian (Mammal Neogene [MN] zone MN7+8). Here we revise the age of CM1 based on fieldwork and associated paleomagnetic samplings undertaken in 2018-2021. Our results extend the ACM composite sequence upward and indicate that CM1 correlates to the earliest Vallesian (MN9). The updated ACM sequence has a thickness of ∼300 m and comprises 12 magnetozones correlated to subchrons C5Ar.1r to C5n.2n (∼12.6-11.1 Ma; latest MN6 to earliest MN9, late Aragonian to earliest Vallesian). CM1 is correlated to C5r.1r (11.146-11.056 Ma), with an interpolated age of 11.11 Ma, thus postdating the dispersal of hipparionin horses into the Vallès-Penedès Basin-which is correlated to the previous subchron C5r.1n, with an interpolated age of 11.18 Ma, and by definition marks the beginning of the Vallesian. CM1 also minimally postdates the earliest record of giraffids at ACM-representing their earliest well-dated occurrence in the basin-being correlated to C5r.1n with an interpolated age of 11.11 Ma. We conclude that CM1 has an earliest Vallesian (MN9) age of ∼11.1 Ma, intermediate between the Aragonian dryopithecins and the Vallesian hispanopithecins. Ongoing paleontological surveillance at ACM thus offers the prospect to yield additional earliest Vallesian ape remains, which are essential to clarify their taxonomic allocation as well as to confirm whether hispanopithecins evolved locally from dryopithecins rather than immigrating from elsewhere during MN9.

Palaeoecological differences underlie rare co-occurrence of Miocene European primates.

BACKGROUND: The two main primate groups recorded throughout the European Miocene, hominoids and pliopithecoids, seldom co-occur. Due to both their rarity and insufficiently understood palaeoecology, it is currently unclear whether the infrequent co-occurrence of these groups is due to sampling bias or reflects different ecological preferences. Here we rely on the densely sampled primate-bearing sequence of Abocador de Can Mata (ACM) in Spain to test whether turnovers in primate assemblages are correlated with palaeoenvironmental changes. We reconstruct dietary evolution through time (ca. 12.6-11.4 Ma), and hence climate and habitat, using tooth-wear patterns and carbon and oxygen isotope compositions of enamel of the ubiquitous musk-deer Micromeryx. RESULTS: Our results reveal that primate species composition is strongly correlated with distinct environmental phases. Large-bodied hominoids (dryopithecines) are recorded in humid, densely-forested environments on the lowermost portion of the ACM sequence. In contrast, pliopithecoids inhabited less humid, patchy ecosystems, being replaced by dryopithecines and the small-bodied Pliobates toward the top of the series in gallery forests embedded in mosaic environments. CONCLUSIONS: These results support the view that pliopithecoid primates preferred less humid habitats than hominoids, and reveal that differences in behavioural ecology were the main factor underpinning their rare co-occurrence during the European Miocene. Our findings further support that ACM hominoids, like Miocene apes as a whole, inhabited more seasonal environments than extant apes. Finally, this study highlights the importance of high-resolution, local investigations to complement larger-scale analyses and illustrates that continuous and densely sampled fossiliferous sequences are essential for deciphering the complex interplay between biotic and abiotic factors that shaped past diversity.

Bio- and magnetostratigraphic correlation of the Miocene primate-bearing site of Castell de Barberà to the earliest Vallesian.

Castell de Barberà, located in the Vallès-Penedès Basin (NE Iberian Peninsula), is one of the few European sites where pliopithecoids (Barberapithecus) and hominoids (cf. Dryopithecus) co-occur. The dating of this Miocene site has proven controversial. A latest Aragonian (MN7+8, ca. 11.88-11.18 Ma) age was long accepted by most authors, despite subsequent reports of hipparionin remains that signaled a Vallesian age. On the latter basis, Castell de Barberà was recently correlated to the early Vallesian (MN9, ca. 11.18-10.3 Ma) on tentative grounds. Uncertainties about the provenance of the Hippotherium material and the lack of magnetostratigraphic data precluded more accurate dating. After decades of inactivity, fieldwork was resumed in 2014-2015 at Castell de Barberà, including the original layer (CB-D) that previously delivered most of the fossils. Here we report magnetostratigraphic results for the original outcrop and another nearby section. Our results indicate that CB-D is located in a normal polarity magnetozone in the middle of a short (∼20 m-thick) stratigraphic section. The composite magnetostratigraphic section (∼50 m) has as many as four to six magnetozones. These multiple reversals, coupled with the in situ recovery of a Hippotherium humerus from CB-D in 2015, make it unlikely that any of the sampled normal polarity magnetozones correlate with the long normal polarity subchron C5n.2n (11.056-9.984 Ma), which is characteristic of the early Vallesian. Our results support instead a correlation of CB-D with C5r.1n (11.188-11.146 Ma), where the Aragonian/Vallesian boundary is situated, and therefore indicate an earliest Vallesian age of ∼11.2 Ma for Castell de Barberà. Our results settle the longstanding debate about the Aragonian vs. Vallesian age of this site, which appears roughly coeval with the Creu de Conill 20 locality (11.18 Ma), where hipparionins are first recorded in the Vallès-Penedès Basin.

Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards.

Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively.

Oldest skeleton of a fossil flying squirrel casts new light on the phylogeny of the group.

Flying squirrels are the only group of gliding mammals with a remarkable diversity and wide geographical range. However, their evolutionary story is not well known. Thus far, identification of extinct flying squirrels has been exclusively based on dental features, which, contrary to certain postcranial characters, are not unique to them. Therefore, fossils attributed to this clade may indeed belong to other squirrel groups. Here we report the oldest fossil skeleton of a flying squirrel (11.6 Ma) that displays the gliding-related diagnostic features shared by extant forms and allows for a recalibration of the divergence time between tree and flying squirrels. Our phylogenetic analyses combining morphological and molecular data generally support older dates than previous molecular estimates (~23 Ma), being congruent with the inclusion of some of the earliest fossils (~36 Ma) into this clade. They also show that flying squirrels experienced little morphological change for almost 12 million years.

Ten years in the dump: An updated review of the Miocene primate-bearing localities from Abocador de Can Mata (NE Iberian Peninsula).

More than ten years of paleontological fieldwork during the enlargement of the Can Mata Landfill (Abocador de Can Mata [ACM]), in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula) led to the recovery of >60,000 Miocene vertebrate remains. The huge sampling effort (due to continuous surveillance of heavy machinery digging activity, coupled with manual excavation and screen-washing of sediments) enabled generally rare faunal elements such as pliopithecoid and hominoid primates to be found. Thanks to detailed litho-, bio- and magnetostratigraphic controls, accurate dating is possible for all the recovered primate remains from 19 of the 235 localities defined along the 234 m-thick composite stratigraphic sequence of the ACM. Here we report updated estimated (interpolated) ages for these paleontological localities and review the timing of the primate succession in this area. Our results indicate that the whole ACM sequence is late Aragonian in age (MN6 and MN7+8) and includes seven magnetozones that are correlated to subchrons C5Ar.1r to C5r.2r (ca. 12.6 to 11.4 Ma). Great apes (dryopithecines) are first recorded at 12.4-12.3 Ma, but most of the finds (Anoiapithecus, Pierolapithecus and Dryopithecus) cluster between 12.0 and 11.9 Ma, followed by some indeterminate dryopithecine remains between 11.7 and 11.6 Ma. Pliopithecoids first appear at 12.1 Ma, being subsequently represented by Pliopithecus between 11.9 and 11.7 Ma. The small-bodied hominoid Pliobates is the youngest ACM primate, with an estimated age of 11.6 Ma. Although these primates probably overlapped in time, their co-occurrence is recorded only twice, at 11.9 Ma (a dryopithecine with Pliopithecus) and at 11.6 Ma (a dryopithecine with Pliobates). The rare co-occurrence between great apes and small-bodied catarrhines might be attributable to sampling biases and/or to presumed diverging ecological preferences of these groups. In the future, more detailed analyses of the fauna recovered from the long and densely-sampled ACM sequence will hopefully throw new light on this long-standing, unresolved question.

Miocene small-bodied ape from Eurasia sheds light on hominoid evolution.

Miocene small-bodied anthropoid primates from Africa and Eurasia are generally considered to precede the divergence between the two groups of extant catarrhines­hominoids (apes and humans) and Old World monkeys­and are thus viewed as more primitive than the stem ape Proconsul. Here we describe Pliobates cataloniae gen. et sp. nov., a small-bodied (4 to 5 kilograms) primate from the Iberian Miocene (11.6 million years ago) that displays a mosaic of primitive characteristics coupled with multiple cranial and postcranial shared derived features of extant hominoids. Our cladistic analyses show that Pliobates is a stem hominoid that is more derived than previously described small catarrhines and Proconsul. This forces us to reevaluate the role played by small-bodied catarrhines in ape evolution and provides key insight into the last common ancestor of hylobatids (gibbons) and hominids (great apes and humans).

An amphisbaenian skull from the European miocene and the evolution of Mediterranean worm lizards.

The evolution of blanid amphisbaenians (Mediterranean worm lizards) is mainly inferred based on molecular studies, despite their fossils are common in Cenozoic European localities. This is because the fossil record exclusively consists in isolated elements of limited taxonomic value. We describe the only known fossil amphisbaenian skull from Europe - attributed to Blanus mendezi sp. nov. (Amphisbaenia, Blanidae) - which represents the most informative fossil blanid material ever described. This specimen, from the Middle Miocene of Abocador de Can Mata (11.6 Ma, MN7+8) in the Vallès-Penedès Basin (Catalonia, NE Iberian Peninsula), unambiguously asserts the presence of Blanus in the Miocene of Europe. This reinforces the referral to this genus of the previously-known, much more incomplete and poorly-diagnostic material from other localities of the European Neogene. Our analysis - integrating the available molecular, paleontological and biogeographic data - suggests that the new species postdates the divergence between the two main (Eastern and Western Mediterranean) extant clades of blanids, and probably precedes the split between the Iberian and North-Western African subclades. This supports previous paleobiogeographic scenarios for blanid evolution and provides a significant minimum divergence time for calibrating molecular analyses of blanid phylogeny.

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 P(3)-M(1) (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M(1)-M(2) and left P(4)-M(2) (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 M(2) 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.

Kretzoiarctos gen. nov., the oldest member of the giant panda clade.

The phylogenetic position of the giant panda, Ailuropoda melanoleuca (Carnivora: Ursidae: Ailuropodinae), has been one of the most hotly debated topics by mammalian biologists and paleontologists during the last century. Based on molecular data, it is currently recognized as a true ursid, sister-taxon of the remaining extant bears, from which it would have diverged by the Early Miocene. However, from a paleobiogeographic and chronological perspective, the origin of the giant panda lineage has remained elusive due to the scarcity of the available Miocene fossil record. Until recently, the genus Ailurarctos from the Late Miocene of China (ca. 8-7 mya) was recognized as the oldest undoubted member of the Ailuropodinae, suggesting that the panda lineage might have originated from an Ursavus ancestor. The role of the purported ailuropodine Agriarctos, from the Miocene of Europe, in the origins of this clade has been generally dismissed due to the paucity of the available material. Here, we describe a new ailuropodine genus, Kretzoiarctos gen. nov., based on remains from two Middle Miocene (ca. 12-11 Ma) Spanish localities. A cladistic analysis of fossil and extant members of the Ursoidea confirms the inclusion of the new genus into the Ailuropodinae. Moreover, Kretzoiarctos precedes in time the previously-known, Late Miocene members of the giant panda clade from Eurasia (Agriarctos and Ailurarctos). The former can be therefore considered the oldest recorded member of the giant panda lineage, which has significant implications for understanding the origins of this clade from a paleobiogeographic viewpoint.

New dental remains of Hispanopithecus laietanus (Primates: Hominidae) from Can Llobateres 1 and the taxonomy of Late Miocene hominoids from the Vallès-Penedès Basin (NE Iberian Peninsula).

Here we report 12 teeth of the fossil great ape Hispanopithecus (Hominidae: Dryopithecinae: Hispanopithecini), recovered in 2011 from the locality of Can Llobateres 1 (MN9, early Vallesian, Late Miocene, ca. 9.7 Ma [millions of years ago]) in the Vallès-Penedès Basin (Catalonia, Spain). Besides an isolated dP(3) from layer CLL1.1b in the eastern (classical) sector of the site, all of the remaining teeth come from facies CLL1.0 (roughly equivalent to CLL1.2 and CLL1.1b), located in the newly excavated western sector, and representing at least two different individuals. Based on facet congruence and degree of wear, all of the upper cheek teeth, a central incisor and a lateral incisor most likely correspond to a single young adult individual of unknown sex, whereas a very worn I(2) and a female C(1) represent one or two additional individual(s). Morphological and metrical comparisons allow us to attribute these remains to Hispanopithecus laietanus, which is the single hominoid species recognized at CLL1. The newly described teeth represent a significant addition to the hypodigm of this taxon, enabling us to more completely assess the degree of variation displayed by several teeth. In light of the new specimens, the previous tooth position assignment of several upper molars from Can Llobateres and Can Poncic is revised, and the criteria employed to distinguish Hispanopithecus crusafonti from H. laietanus are critically evaluated. On the basis of the available upper cheek teeth from these localities, a distinction at the species level between both samples is tentatively favored, mainly on the basis of P(3), M(1) and M(2) proportions as well as I(1) lingual morphology and proportions. The results of the 2011 field season unambiguously confirm that hominoid-bearing fossiliferous layers from CLL1 are not exhausted. Additional excavations at this site are promising for the discovery of additional remains of H. laietanus in the near future.

The paleoenvironment of Hispanopithecus laietanus as revealed by paleobotanical evidence from the Late Miocene of Can Llobateres 1 (Catalonia, Spain).

The early Vallesian site of Can Llobateres 1 (Vallès-Penedès Basin, Catalonia, Spain) is one of the richest localities of the European Late Miocene, having yielded the most complete remains of the fossil great ape Hispanopithecus laietanus (Primates: Hominidae). Fossil plant remains had been previously reported from this site but mostly remained unpublished. Here we describe an assemblage of plant megaremains recovered in 2010, which provides valuable paleoenvironmental data. This assemblage consists of a mixture of parautochthonous and allochthonous detached organs (leaves, stems, reproductive structures) deposited in marshy areas. The source vegetation mainly consisted of abundant reeds, palms, evergreen laurels and figs that probably grew in or near the marsh boundaries or nearby riparian forests. This environmental picture is consistent with the mammalian fauna, which shows the prevalence of humid forested environments, although somewhat more open woodlands might have been present away from the wet areas. The occurrence of mega-mesothermal taxa, together with the absence of deciduous elements, is consistent with a subtropical to warm-temperate climate. Within this mosaic environment, H. laietanus would have preferred the more humid and forested habitats, which probably were still quite common in the Vallès-Penedès during the early Vallesian. Such habitats would have provided a continuous ripe fruit supply throughout the year to these frugivorous great apes. Paleobotanical data from older sites of the same area and nearby basins show that the zonal vegetation was a warm-temperate mixed forest defined by evergreen laurels, together with leguminous trees and shrubs as well as a significant proportion of deciduous elements. Tropical and subtropical taxa would have been restricted to humid areas in the lowlands. From the late Vallesian onwards, many of these taxa disappeared from the Vallès-Penedès, whereas deciduous trees became dominant in the forested areas and wetlands, thus likely having driven Hispanopithecus to extinction in the study area.

Brief communication: the oldest pliopithecid record in the Iberian Peninsula based on new material from the Vallès-Penedès Basin.

Here, we report on an isolated pliopithecid M3/ (IPS35028) from locality ACM/C3-B2 (12.0 Ma, MN7) of the late Middle Miocene stratigraphic series of Abocador de Can Mata (ACM, Vallès-Penedès Basin, NE Iberian Peninsula). This tooth is about 0.2 million years older than the remains of Pliopithecus canmatensis (11.8-11.7 ma), recorded from several localities from the ACM series. The unusual occlusal features of IPS35028, together with the lack of homologous material for several pliopithecid species, preclude a precise taxonomic attribution of the C3-B2 specimen, which does not fit the morphology of any known pliopithecid M3/. In particular, although an attribution to P. canmatensis would seem reasonable on the basis of size, identical geographic provenance, and similar age, the morphology of IPS35028 appears too primitive compared to the M1/ and M2/ of the former taxon. Instead, the C3-B2 pliopithecid displays several primitive features shared with the dionysopithecine Dionysopithecus and the pliopithecine Pliopithecus piveteaui. It therefore seems more likely that IPS35028 represents a previously unknown pliopithecid taxon, although a formal taxonomic recognition of its probable distinct status is not advisable, given the scarcity of the currently available material. Alternatively, this taxon might be more closely related to small-bodied African catarrhines (such as dendropithecids). However, the morphology of the ACM specimen is not particularly similar to that of the M3/ of these African taxa. Hence, based on age and geographic provenance, an attribution of IPS35028 to the Pliopithecidae is favored here.

Updated chronology for the Miocene hominoid radiation in Western Eurasia.

Extant apes (Primates: Hominoidea) are the relics of a group that was much more diverse in the past. They originated in Africa around the Oligocene/Miocene boundary, but by the beginning of the Middle Miocene they expanded their range into Eurasia, where they experienced a far-reaching evolutionary radiation. A Eurasian origin of the great ape and human clade (Hominidae) has been favored by several authors, but the assessment of this hypothesis has been hampered by the lack of accurate datings for many Western Eurasian hominoids. Here we provide an updated chronology that incorporates recently discovered Iberian taxa and further reevaluates the age of many previously known sites on the basis of local biostratigraphic scales and magnetostratigraphic data. Our results show that identifiable Eurasian kenyapithecins (Griphopithecus and Kenyapithecus) are much younger than previously thought (ca. 14 Ma instead of 16 Ma), which casts serious doubts on the attribution of the hominoid tooth from Engelswies (16.3-16.5 Ma) to cf. Griphopithecus. This evidence is further consistent with an alternative scenario, according to which the Eurasian pongines and African hominines might have independently evolved in their respective continents from similar kenyapithecin ancestors, resulting from an early Middle Miocene intercontinental range extension followed by vicariance. This hypothesis, which would imply an independent origin of orthogrady in pongines and hominines, deserves further testing by accurately inferring the phylogenetic position of European dryopithecins, which might be stem pongines rather than stem hominines.

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 approximately 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.