A revised systematic scheme for the Eurasian Miocene fossil Hominidae

doi:10.1006/jhev.1997.0145

Journal of Human Evolution

Volume 33, Issue 4, October 1997, Pages 449-477

https://doi.org/10.1006/jhev.1997.0145 Get rights and content

Abstract

The current systematics of the European hominids has recently been re-examined in light of new and important fossil discoveries. It has been argued by Begun (1992a) that the European dryopithecines (includingGraecopithecusfrom Greece) represent one of the earliest members of the Hominidae. Andrews (1992) and Dean & Delson (1992) agree thatGraecopithecusshould probably be placed within the Homininae, but thatDryopithecusshould be excluded from this subfamily and retained within the Dryopithecinae. De Bonis & Koufos (1993, 1994) suggest thatGraecopithecusshares a sister-group relationship with hominins. Moyà-Solà & Köhler (1993, 1995), however, argue thatDryopithecusandGraecopithecusboth represent primitive members of the Ponginae. A phylogenetic and taxonomic revision of these taxa presented here supports the allocation ofGraecopithecusto Homininae. It is also shown that the apparent disagreement concerningDryopithecusis a result of the presence of two distinct genera within the fossil samples examined. It is concluded that the Hungarian hominid should be retained within the genusDryopithecusand the subfamily Dryopithecinae; the Spanish hominids are allocated to the resurrected genusHispanopithecusVillalta & Crusafont-Pairó, 1944 which is likely to be a primitive member of the Ponginae.

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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.
Systematics of Miocene apes: State of the art of a neverending controversy
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Hominoids diverged from cercopithecoids during the Oligocene in Afro-Arabia, initially radiating in that continent and subsequently dispersing into Eurasia. From the Late Miocene onward, the geographic range of hominoids progressively shrank, except for hominins, which dispersed out of Africa during the Pleistocene. Although the overall picture of hominoid evolution is clear based on available fossil evidence, many uncertainties persist regarding the phylogeny and paleobiogeography of Miocene apes (nonhominin hominoids), owing to their sparse record, pervasive homoplasy, and the decimated current diversity of this group. We review Miocene ape systematics and evolution by focusing on the most parsimonious cladograms published during the last decade. First, we provide a historical account of the progress made in Miocene ape phylogeny and paleobiogeography, report an updated classification of Miocene apes, and provide a list of Miocene ape species-locality occurrences together with an analysis of their paleobiodiversity dynamics. Second, we discuss various critical issues of Miocene ape phylogeny and paleobiogeography (hylobatid and crown hominid origins, plus the relationships of Oreopithecus) in the light of the highly divergent results obtained from cladistic analyses of craniodental and postcranial characters separately. We conclude that cladistic efforts to disentangle Miocene ape phylogeny are potentially biased by a long-branch attraction problem caused by the numerous postcranial similarities shared between hylobatids and hominids—despite the increasingly held view that they are likely homoplastic to a large extent, as illustrated by Sivapithecus and Pierolapithecus—and further aggravated by abundant missing data owing to incomplete preservation. Finally, we argue that—besides the recovery of additional fossils, the retrieval of paleoproteomic data, and a better integration between cladistics and geometric morphometrics—Miocene ape phylogenetics should take advantage of total-evidence (tip-dating) Bayesian methods of phylogenetic inference combining morphologic, molecular, and chronostratigraphic data. This would hopefully help ascertain whether hylobatid divergence was more basal than currently supported.
Phylogenetic analysis of Middle-Late Miocene apes
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This position accords well with previous morphological observations of Ou. macedoniensis, which is found at three Late Miocene sites in northern Greece (Andrews, 1992; Begun, 1992; Dean and Delson, 1992; de Bonis and Koufos, 1994; Benefit and McCrossin, 1995; Begun et al., 1997; Cameron, 1997; Harrison, 2010a). Favorable comparisons between Ou.
Despite intensive study, many aspects of the evolutionary history of great apes and humans (Hominidae) are not well understood. In particular, the phylogenetic relationships of many fossil taxa remain poorly resolved. This study aims to provide an updated hypothesis of phylogenetic relationships for Middle-Late Miocene fossil apes, focusing on those taxa typically considered to be great apes. The character matrix compiled here samples 274 characters from the skull, dentition, and postcranium. Multiple iterations were performed to examine the effects of ingroup taxon selection, outgroup constraints, treatment of continuous data, character partitions (craniodental, postcranial), and missing data. Parsimony and Bayesian methods were used to infer phylogenetic relationships. Most European hominoids (Hispanopithecus, Rudapithecus, Dryopithecus, Pierolapithecus) are recovered as stem hominids, not more closely related to orangutans or to African apes and humans (Homininae), whereas Ouranopithecus, Graecopithecus, and Nakalipithecus are inferred to be members of the hominine clade. Asian fossil hominoids, with the exception of Lufengpithecus hudienensis, are recovered as part of the orangutan clade (Ponginae). Results suggest that Kenyapithecus and Griphopithecus are possible stem hominids, whereas Equatorius and Nacholapithecus are consistently recovered as stem hominoids. Oreopithecus and Samburupithecus are not recovered as hominids. Results of Bayesian analyses differ from those of parsimony analyses. Craniodental and postcranial character partitions are incongruent in the placement of hylobatids, which is interpreted as evidence that hylobatids and hominids independently evolved adaptations to suspensory positional behaviors. An understanding of phylogenetic relationships is necessary to address many of the questions asked in paleoanthropology. Thus, the updated hypothesis of phylogenetic relationships presented here can be used to gain a better understanding of important morphological transitions that took place during hominid evolution, ancestral morphotypes at key nodes, and the biogeography of the clade.
Phylogenetic signal in molar dental shape of extant and fossil catarrhine primates
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Despite the distinct anatomical traits in both groups, the classification of extinct catarrhines has been extensively debated (Fleagle, 1999; Begun, 2002, 2010; Harrison, 2002; Rasmussen, 2002; Casanovas-Vilar et al., 2011). In this context, phylogenetic relationships for the Eurasian Miocene hominoids have been a long standing topic of debate (Moyà-Solà and Köhler, 1995; Begun et al., 1997, 2012; Cameron, 1997; Stewart and Disotell, 1998; Andrews and Bernor, 1999; Begun and Nargolwalla, 2004; Moyà-Solà et al., 2004, 2009a,b; Begun, 2007, 2010; Alba et al., 2010b; Senut, 2010; Casanovas-Vilar et al., 2011). This is especially true for the controversial phylogenetic position of the dryopithecinae clade that some researchers closely relate to the African great apes (Homininae, hominines; Begun et al., 1997, 2012; Begun, 2009, 2010), while others relate to the Asian pongines (Ponginae; Moyà-Solà and Köhler, 1995; Alba et al., 2010b; Casanovas-Vilar et al., 2011).
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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 dP³ 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 I² and a female C¹ 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 P³, M¹ and M² proportions as well as I¹ 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)
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Citation Excerpt :
Nevertheless, controversies and uncertainties persist regarding the phylogenetic relationships of Hispanopithecus with extant and other fossil great apes. Thus, several authors have considered it to be a stem pongine (Moyà-Solà and Köhler, 1993; Köhler et al., 2001; Grehan and Schwartz, 2009), in agreement with the extended concept of the Ponginae favored by some authors (Schwartz, 1990, 1997; Cameron, 1997). This is further reinforced by the orang-like locomotor adaptations displayed by this taxon (Moyà-Solà and Köhler, 1996a; Almécija et al., 2007; Alba et al., 2010a).
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.

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^☆: Tobias, P. V.Coppens, Y.

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Regular ArticleA revised systematic scheme for the Eurasian Miocene fossil Hominidae☆

Abstract

J. hum. Evol.

J. hum. Evol.

J. hum. Evol.

J. hum. Evol.

Geobios

Molec. Phylogene. Evol.

Evolution and environment in the Hominidea

Nature

Ramapithecus

Les plus anciens hominides: Colloque VI, IX Cong

A Revision of the Turkish Miocene Hominid Sivapithecus meteai

Palaeontology

Hominid dietary evolution

Phil. Trans. R. Soc. Lond. B.

The nature of the evidence

Nature

Miocene fossil hominids and the chimp–human clade

Science

Dryopithecus crusafonti

Am. J. phys. Anthrop.

Phyletic Diversity and Locomotion in Primitive European Hominids

Am. J. phys. Anthrop.

Relations Among the Great Apes and Humans: New Interpretations Based on the Fossil Great Ape Dryopithecus

Yearb. Phys. Anthropol.

Observations on the cranial anatomy ofOuranopithecus

Am. J. phys. Anthrop.

Restoration and reinterpretation of the facial specimen attributed toSivapithecus meteai

Am. J. phys. Anthrop.

Phyletic Relationships of Miocene Hominids and Higher Primate Classification

Première découverte d'un Primate hominidé dans le Miocène supérieur de Macédoine (Grèce)

C. R. Acad. Sci. Paris

Nouveaux restes de Primates hominidés dans le Vallésien de Macédoine (Grèce)

C. R. Acad. Sci. Paris

Un nouveau genre de primate hominı̈de dans le Vallésien (Miocène supérieur) de Macédoine

C. R. Acad. Sci. Paris

Variations métriques de la denture chez les Hominidés primitifs: comparaison entreAustralopithecus afarensisOuranopithecus macedonensis

C. R. Acad. Sci. Paris, t. 292 série II

La position phylétique d'Ouranopithecus

Cour. Forsch.-Inst. Senkenberg.

La place du genreOuranopithecus

C. R. Acad. Sci. Paris, t.300 Serie II

Découverte d'un nouveau gisement de Vertébrés dans le Miocène supérieur de Macédoine (Grèce)

C. R. Acad. Sci. Paris, t.303 Série II

Intérét de l'anatomie naso-maxillaire pour la phylogénie des Hominidés

C. R. Acad. Sci. Paris, t.304 Serie II

New hominid skull material from the late Miocene of Macedonia in Northern Greece

Nature

The face and the mandible ofOuranopithecus macedonensis

J. hum. Evol.

Our Ancestor:Ouranopithecus

Evol. Anthrop.

Regular Article
A revised systematic scheme for the Eurasian Miocene fossil Hominidae☆