Josep M. Robles

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.

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.

A unique Middle Miocene European hominoid and the origins of the great ape and human clade

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.

Biochronological, taphonomical, and paleoenvironmental background of the fossil great ape Pierolapithecus catalaunicus (Primates, Hominidae)

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.

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.

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.

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

Meet your gibbon cousin Apes are divided into two groups: larger-bodied apes, or hominoids, such as humans, chimps, and gorillas; and smaller-bodied hylobatids, such as gibbons. These two lineages are thought to have diverged rather cleanly, sharing few similarities after the emergence of crown hominoids. Alba et al. describe a new ape from the Miocene era that contains characteristics from both hominoids and small-bodied apes (see the Perspective by Benefit and McCrossin). Thus, early small-bodied apes may have contributed more to the evolution of the hominoid lineage than previously assumed. Science, this issue p. 10.1126/science.aab2625; see also p. 515 Early small-bodied apes may have contributed more to the evolution of the hominoid lineage than previously thought. [Also see Perspective by Benefit and McCrossin] INTRODUCTION Reconstructing the ancestral morphotype from which extant hominoids (apes and humans) evolved is complicated by the mosaic nature of ape evolution, the confounding effects of independently evolved features (homoplasy), and the virtual lack of hylobatids (gibbons and siamangs) in the Miocene fossil record. For several decades, small-bodied anthropoid primates from Africa and Eurasia have not played an important role in this debate, because they generally lack the shared derived features of extant catarrhines (hominoids and Old World monkeys) and are thus considered to precede their divergence. Even some small-bodied catarrhines from Africa (dendropithecids), considered to be stem hominoids by some authors, are viewed as more primitive than the larger-bodied stem ape Proconsul. This has led to the assumption that hylobatids are a dwarfed lineage that evolved from a larger-bodied and more great ape–like common ancestor with hominids (great apes and humans). RATIONALE Here we describe a new genus of small-bodied (4 to 5 kg) ape from the Miocene (11.6 Ma), discovered in the Abocador de Can Mata stratigraphic series (Vallès-Penedès Basin, northeast Iberian Peninsula), that challenges current views on the last common ancestor of extant hominoids. This genus is based on a partial skeleton that enables a reliable reconstruction of cranial morphology and a detailed assessment of elbow and wrist anatomy. It exhibits a mosaic of primitive (stem catarrhine–like) and derived (extant hominoid–like) features that forces us to reevaluate the role played by small-bodied catarrhines in ape evolution. RESULTS The new genus retains some features that are suggestive of generalized above-branch quadrupedalism, but it possesses more extensive hominoid-like postcranial features (mostly related to enhanced forearm rotation and ulnar deviation capabilities) than those convergently displayed by atelids. Its overall body plan is more compatible with an emphasis on cautious and eclectic climbing, combined with some degree of below-branch forelimb-dominated suspension (although less acrobatic than in extant gibbons). Its relative brain size implies a monkey-like degree of encephalization (similar to that of hylobatids but below that of great apes), and dental microwear indicates a frugivorous diet. From a phylogenetic viewpoint, the new genus combines craniodental and postcranial primitive features (similar to those of dendropithecids) with multiple derived cranial and postcranial features shared with extant hominoids. Some cranial similarities with gibbons would support a closer phylogenetic link between the new genus and hylobatids. However, this possibility is not supported by the total evidence. A cladistic analysis based on more than 300 craniodental and postcranial features reveals that the new genus is a stem hominoid (preceding the divergence between hylobatids and hominids), although more derived than previously known small catarrhines and Proconsul. CONCLUSION As the first known Miocene small-bodied catarrhine to share abundant derived features with extant hominoids, the new genus indicates a greater morphological diversity than previously recognized among this heterogeneous group, and it provides key insight into the last common ancestor of hylobatids and hominids. Our cladistic results, coupled with the chronology and location of the new genus, suggest that it represents a late-surviving offshoot of a small African stem hominoid that is more closely related to crown hominoids than Proconsul is. These results suggest that, at least in size and cranial morphology, the last common ancestor of extant hominoids might have been more gibbon-like (less great ape–like) than generally assumed. Cranial reconstruction and life appearance. Artist’s representation of the cranial reconstruction (in frontal view) and of the life appearance (in lateral oblique view) of the new genus of small-bodied ape from the Iberian Miocene. [Artwork by M. Palmero] Miocene small-bodied anthropoid primates from Africa and Eurasia are generally considered to precede the divergence between the two groups of extant catarrhines—hominoids (apes and humans) and Old World monkeys—and 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).

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.

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 craniodental remains of the barbourofelid Albanosmilus jourdani (Filhol, 1883) from the Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula) and the phylogeny of the Barbourofelini

Available remains of the barbourofelin Albanosmilus jourdani from the Middle to Late Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula) are described. In addition to the dentognathic remains described by previous authors, the new material includes a complete cranium, a calvarium and several mandibles from Abocador de Can Mata, Creu Conill 20 and Hostalets Superior. It is concluded that Albanosmilus, previously considered a subjective junior synonym of Sansanosmilus, must be resurrected as a polytypic genus including A. jourdani (= A. vallesiensis). The most plesiomorphic North American barbourofelin, previously included in Barbourofelis, is also transferred into Albanosmilus as A. whitfordi. An emended diagnosis of Albanosmilus is provided. The results of a cladistic analysis support the monophyly of the family Barbourofelidae and the tribe Barbourofelini, further indicating that amongst the latter, Sansanosmilus occupies the basalmost position. The two Albanosmilus species are more derived, although the analysis fails to resolve conclusively whether A. whitfordi is more closely related to A. jourdani or Barbourofelis s.s. From a palaeobiogeographical viewpoint, our results suggest that: (1) barbourofelins originated in Eurasia during the early Middle Miocene; (2) Barbourofelis originated in North America during the late Middle Miocene, following the dispersal of Eurasian Albanosmilus into that continent; and (3) the presence of Barbourofelis in Turkey during the Late Miocene may represent a later independent dispersal event from North America back into Eurasia.