Abstract
Torso-orthograde (TO)-positional behavior and an associated suite of postcranial morphological specializations , both of which are often linked to the evolution of large body size , are considered hallmarks of hominoid evolution . These features are thought to have played an important role in the Old World monkey-ape divergence and the development of upright bipedal locomotion in our own lineage. Compared to early theorists, who considered hylobatids an appropriate model for investigating the initial stages of hominoid evolution, more recent reconstructions of the last common ancestor of hominoids have advocated a larger-bodied, more generalized ape very different from hylobatids as the most likely morphotype. While field observations from the large-bodied non-human hominids (i.e., Asian and African apes) confirm that they do in fact utilize an expansive TO-positional repertoire, no detailed data from hylobatids has been available until now to fully evaluate this hypothesis in light of all hominoid taxa. Using recently published positional behavior data from a population of adult white-handed gibbons ( Hylobates lar ) from Khao Yai National Park , Thailand , and integrating these data with a diverse anthropoid primate comparative sample, we reevaluate differences in positional behavior of extant cercopithecoids and hominoids. The comparative dataset agrees with recent suggestions that all living apes share a diverse TO-positional repertoire, relative to that of cercopithecoid monkeys. The question, whether or not TO-anatomical specializations and behavior among the Hominoidea represent homologous or homoplastic traits or both, could not yet be answered conclusively, despite the fact that homologous evolution is the more parsimonious explanation, due to a lack of support from the fossil record. Nevertheless, an upright (TO) and enhanced exploitation of the arboreal tree canopy by our ape ancestors were likely key evolutionary novelties that helped to shape the origin of hominins. Only an expansive understanding of the diversity of both extant and extinct apes will help to comprehensively unravel our enigmatic origins.
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Notes
- 1.
Torso-orthograde locomotion includes bipedalism, brachiation/forelimb swing, clamber/transfer, and vertical climb, as outlined by Hunt et al. 1996.
- 2.
Torso-orthograde posture includes bipedalism and forelimb suspension, as outlined by Hunt et al. 1996.
- 3.
Extant apes share a number of postcranial anatomical features, including a relatively short stable lumbar region, a mediolaterally broad and dorsoventrally shallow thorax, relatively long forelimbs, a more dorsal positioning of the scapula, a mobile shoulder joint, an elbow that is fully extendable, a forearm that is stable during a wide range of pronation and supination, a mobile wrist, long curved phalanges, the lack of a tail, in addition to myological specializations associated with arm extension/flexion, forearm pronation/supination, and hand flexion (Keith 1923; Schultz 1930; Harrison 1987; 2016; Hunt 1991b, 2016; Rose 1993, 1994, 1997; Gebo 1996; Ward 1997a, 2015; Larson 1998; Stern and Larson 2001; Gibbs et al. 2002; Young 2003; Kagaya et al. 2008, 2009, 2010; Diogo and Wood 2011; Chan 2014; Williams and Russo 2015).
- 4.
We follow Harrison (2010a, 2013) in placing Proconsul within the subfamily Proconsulinae; Afropithecus, Equatorius, Heliopithecus, Morotopithecus (Afropithecus?), and Nacholapithecus within the subfamily Afropithecinae; and Mabokopithecus, Nyanzapithecus, Rangwapithecus, Turkanopithecus, and Xenopithecus in the subfamily Nyanzapithecinae. Rukwapithecus is also tentatively placed in the subfamily Nyanzapithecinae, following Stevens et al. (2013). We recognize that McNulty et al. (2015) separate the fossil material commonly associated with the genus Proconsul, into two genera, Ekembo and Proconsul; however, given that the taxonomic changes are relatively recent and that the available postcranial material has been described as relatively homogeneous (Ward 1998), we tentatively place Ekembo as a separate genus within the subfamily Proconsulinae and utilize Ekembo/Proconsul to collectively designate the group, except in instances where a specific species is highlighted. Also similar to Harrison (2010a, 2013) we group the aforementioned subfamilies in the family Proconsulidae and superfamily Proconsuloidea (which may be paraphyletic); however, we place the Proconsuloidea as stem hominoids (Begun et al. 1997; Rae 1999; Fleagle 2013). Some recent studies prefer to place the Proconsulidae into the superfamily Hominoidea (Zalmout et al. 2010; Stevens et al. 2013). The placement of Otavipithecus is unclear, and we therefore place this taxon incertae sedis among the Proconsuloidea. Following Harrison et al. (2008), Yuanmoupithecus currently remains the only fossil within the family Hylobatidae (Harrison 2016). Within the family Hominidae, we tentatively place the genera Anoiapithecus, Dryopithecus, Graecopithecus, Hispanopithecus, Oreopithecus, Pierolapithecus, Rudapithecus, and Ouranopithecus, within the subfamily Dryopithecinae (i.e., stem hominids) (Harrison 2010b; Fleagle 2013). Furthermore, the species Ankarapithecus, Gigantopithecus, Khoratpithecus, and Sivapithecus are tentatively grouped into the hominid subfamily Ponginae (i.e., stem pongines) (Harrison 2010b; Fleagle 2013). As incertae sedis within the family Hominidae, we place the species Chororapithecus, Lufengpithecus, Nakalipithecus, and Samburupithecus (Harrison 2010a; Ji et al. 2013). The taxonomic placement of Griphopithecus and Kenyapithecus is highly enigmatic (i.e., they may represent stem hominoids, early crown hominoids, or stem hominids); however, we tentatively group them incertae sedis in the family Hominidae (Harrison 2010a; Fleagle 2013). Finally, we highlight the recent work of Alba et al. (2015), who named a new fossil genus Pliobates, which they placed within the superfamily Hominoidea, in a newly named family Pliobatidae (i.e., stem hominoids). However, Benefit and McCrossin (2015), in response to the aforementioned publication have made a good case that Pliobates may actually be better placed within the early catarrhine superfamily Pliopithecoidea. Until subsequent studies confirm the placement of Pliobates within the Hominoidea, we follow Benefit and McCrossin’s (2015) suggestion and view Pliobates as a pliopithecoid.
- 5.
From this point on, we highlight in bold the specific modes detailed in Hunt et al. (1996) to maintain consistency and to differentiate these modes from expansive and/or vague locomotor and postural categories used in previous studies of positional behavior or in discussions of fossil taxa.
- 6.
The study by Sarringhaus et al. (2014) was not included in the comparative sample presented here, as their study combined data from both arboreal and terrestrial contexts, whereas in this study, the focus was strictly arboreal locomotion and posture.
- 7.
The fossil locations for Dryopithecus, Hispanopithecus, Oreopithecus, Pierolapithecus, and Rudapithecus collectively span from ~13 to 7 Ma (Casanovas-Vilar et al. 2011).
- 8.
The taxonomic placement of Oreopithecus is highly enigmatic. Currently, many authors consider Oreopithecus to be a stem hominid (Harrison and Rook 1997; Begun 2007, 2015; Harrison 2010b; Begun et al. 2012). Some analyses suggest, however, that Oreopithecus shares derived dental traits with one or more of the nyanzapithecines (McCrossin 1992; Harrison 2002, 2010a). If this later scenario is proven to be correct, the postcranial specializations that are shared between Oreopithecus and extant hominoids are likely homoplastic.
- 9.
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Acknowledgements
This chapter greatly benefited from field studies on Sumatran Hylobates agilis and Symphalangus syndactylus, which would not have been possible without the assistance of an NSF Doctoral Dissertation Improvement Grant (BCS 1061477 awarded to MGN).
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Nowak, M.G., Reichard, U.H. (2016). Locomotion and Posture in Ancestral Hominoids Prior to the Split of Hylobatids. In: Reichard, U., Hirai, H., Barelli, C. (eds) Evolution of Gibbons and Siamang. Developments in Primatology: Progress and Prospects. Springer, New York, NY. https://doi.org/10.1007/978-1-4939-5614-2_3
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