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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., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

In the Iberian Peninsula, Miocene apes (Hominoidea) are generally rare and mostly restricted to the Vallès-Penedès Basin. Here we report a new hominoid maxillary fragment with M 2 from this basin. It was surface-collected in March 2017 from the site of Can Pallars i Llobateres (CPL, Sant Quirze del Vallès), where fossil apes had not been previously recorded. The locality of provenance (CPL-M), which has delivered no further fossil remains, is located very close (ca. 50 m) to previously known CPL outcrops, and not very far (ca. 500 m in NW direction) from the classical hominoid-bearing locality of Can Poncic 1. Here we describe the new fossil and, based on the size and proportions of the M 2 , justify its taxonomic attribution to Hispanopithecus cf. laietanus, a species previously recorded from several Vallesian sites of the Vallès-Penedès Basin. Based on the associated mammalian fauna from CPL, we also provide a biochronological dating and a paleoenvironmental reconstruction for the site. The associated fauna enables an unambiguous correlation to the Cricetulodon hartenbergeri - Progonomys hispanicus interval local subzone, with an estimated age of 9.98-9.73 Ma (late Vallesian, MN10). Therefore, CPL-M is roughly coeval with the Hispanopithecus laietanus-bearing localities of Can Llobateres 1 and Can Feu 1, and minimally older than those of La Tarumba 1 and Can Llobateres 2. In contrast, CPL-M is younger than the early Vallesian (MN9) localities of Can Poncic 1 (the type locality of Hispanopithecus crusafonti) as well as Polinyà 2 (Gabarró) and Estació Depuradora d'Aigües Residuals-Riu Ripoll 13, where Hispanopithecus sp. is recorded. The associated fauna from CPL indicates a densely forested and humid paleoenvironment with nearby freshwater. This supports the view that Hispanopithecus might have been restricted to dense wetland forests soon before its extinction during the late Vallesian, due to progressive climatic deterioration. Coupled with the existence of other fossiliferous outcrops in the area, this find is most promising for the prospect of discovering additional fossil hominoid remains in the future., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The extinct dryopithecine Hispanopithecus (Primates: Hominidae), from the Late Miocene of Europe, is the oldest fossil great ape displaying an orthograde body plan coupled with unambiguous suspensory adaptations. On the basis of hand morphology, Hispanopithecus laietanus has been considered to primitively retain adaptations to above-branch quadrupedalism-thus displaying a locomotor repertoire unknown among extant or fossil hominoids, which has been considered unlikely by some researchers. Here we describe a partial skeleton of H. laietanus from the Vallesian (MN9) locality of Can Feu 1 (Vallès-Penedès Basin, NE Iberian Peninsula), with an estimated age of 10.0-9.7 Ma. It includes dentognathic and postcranial remains of a single, female adult individual, with an estimated body mass of 22-25 kg. The postcranial remains of the rib cage, shoulder girdle and forelimb show a mixture of monkey-like and modern-hominoid-like features. In turn, the proximal morphology of the ulna-most completely preserved in the Can Feu skeleton than among previously-available remains-indicates the possession of an elbow complex suitable for preserving stability along the full range of flexion/extension and enabling a broad range of pronation/supination. Such features, suitable for suspensory behaviors, are however combined with an olecranon morphology that is functionally related to quadrupedalism. Overall, when all the available postcranial evidence for H. laietanus is considered, it emerges that this taxon displayed a locomotor repertoire currently unknown among other apes (extant or extinct alike), uniquely combining suspensory-related features with primitively-retained adaptations to above-branch palmigrady. Despite phylogenetic uncertainties, Hispanopithecus is invariably considered an extinct member of the great-ape-and-human clade. Therefore, the combination of quadrupedal and suspensory adaptations in this Miocene crown hominoid clearly evidences the mosaic nature of locomotor evolution in the Hominoidea, as well as the impossibility to reconstruct the ancestral locomotor repertoires for crown hominoid subclades on the basis of extant taxa alone.

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.

The great ape and human clade (Primates: Hominidae) currently includes orangutans, gorillas, chimpanzees, bonobos, and humans. When, where, and from which taxon hominids evolved are among the most exciting questions yet to be resolved. Within the Afropithecidae, the Kenyapithecinae (Kenyapithecini + Equatorini) have been proposed as the sister taxon of hominids, but thus far the fragmentary and scarce Middle Miocene fossil record has hampered testing this hypothesis. Here we describe a male partial face with mandible of a previously undescribed fossil hominid, Anoiapithecus brevirostris gen. et sp. nov., from the Middle Miocene (11.9 Ma) of Spain, which enables testing this hypothesis. Morphological and geometric morphometrics analyses of this material show a unique facial pattern for hominoids. This taxon combines autapomorphic features--such as a strongly reduced facial prognathism--with kenyapithecine (more specifically, kenyapithecin) and hominid synapomorphies. This combination supports a sister-group relationship between kenyapithecins (Griphopithecus + Kenyapithecus) and hominids. The presence of both groups in Eurasia during the Middle Miocene and the retention in kenyapithecins of a primitive hominoid postcranial body plan support a Eurasian origin of the Hominidae. Alternatively, the two extant hominid clades (Homininae and Ponginae) might have independently evolved in Africa and Eurasia from an ancestral, Middle Miocene stock, so that the supposed crown-hominid synapomorphies might be homoplastic.

The Late Aragonian (late middle Miocene) stratigraphic sequence of Abocador de Can Mata (ACM) from Els Hostalets de Pierola (Vallès-Penedès Basin, Catalonia, Spain), rich in fossil vertebrate localities, provides a unique opportunity to study the evolution of western Eurasian hominoids. Among these sites, Barranc de Can Vila 1 (BCV1) recently delivered a well-preserved hominoid partial skeleton of a new genus and species, Pierolapithecus catalaunicus. On the basis of the small-mammal fauna recovered at BCV1, we infer an early MN 7+8 age, between 12.5 and 12Ma, for this site. The spatial distribution of macromammal fossils, the relative abundances of skeletal elements, and their state of preservation suggest that different agents were involved in the accumulation of the P. catalaunicus individual and the remaining taxa. Carnivore marks occur on some bones of the P. catalunicus skeleton, documenting the action of predators and/or scavengers in this case. In contrast, carnivore marks are extremely rare on other macromammal remains, which seem to be derived from adjacent alluvial-fan plain areas. The small-mammal fauna from BCV1 and the large-mammal fauna from the ACM series, indicate the presence of considerably humid and warm forest environments. The compositions of the small-mammal fauna from BCV1 and from other Late Aragonian sites from the Vallès-Penedès area are similar to those from France and central Europe. The former are clearly distinct from those of Iberian inner basins, where the environment appears to have been drier, thus precluding the dispersal of hominoids into that area.

We describe a partial skeleton with facial cranium of Pierolapithecus catalaunicus gen. et sp. nov., a new Middle Miocene (12.5 to 13 million years ago) ape from Barranc de Can Vila 1 (Barcelona, Spain). It is the first known individual of this age that combines well-preserved cranial, dental, and postcranial material. The thorax, lumbar region, and wrist provide evidence of modern ape-like orthograde body design, and the facial morphology includes the basic derived great ape features. The new skeleton reveals that early great apes retained primitive monkeylike characters associated with a derived body structure that permits upright postures of the trunk. Pierolapithecus, hence, does not fit the theoretical model that predicts that all characters shared by extant great apes were present in their last common ancestor, but instead points to a large amount of homoplasy in ape evolution. The overall pattern suggests that Pierolapithecus is probably close to the last common ancestor of great apes and humans.