Published December 31, 2023
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Twenty-five well-justified fossil calibrations for primate divergences
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Vries, Dorien de, Beck, Robin M.D. (2023): Twenty-five well-justified fossil calibrations for primate divergences. Palaeontologia Electronica (a8) 26 (1): 1-52, DOI: 10.26879/1249, URL: http://dx.doi.org/10.26879/1249
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- Figure: 10.5281/zenodo.12655322 (DOI)
References
- Calibrating taxon. Sivapithecus indicus Specimen. (GSP) Y 16075, maxilla (Raza et al., 1983; Kappelman et al., 1991) with the connection between the maxilla and premaxilla partially preserved (Begun, 2015), from locality Y494 from the Chinji Formation, Pakistan (Pilgrim, 1910).
- Phylogenetic justification. Sivapithecus has been consistently recovered as a pongine in recent phylogenetic analyses (e.g., Begun et al., 2012; Nengo et al., 2017; Gilbert et al., 2020; Pugh, 2022). Y 16075 preserves the derived subnasal anatomy characteristic of modern orangutans (Pongo spp.; Kappelman et al., 1991). Isolated teeth from slightly older sites in the Chinji Formation have been referred to Sivapithecus, but they lack diagnostic features to support this referral (Kappelman et al., 1991), and so we do not use these for calibration purposes. We note that the slightly older Kenyapithecus (see "Crown Hominoidea" above) has been recovered as a pongine in some recent phylogenetic analyses (Nengo et al., 2017; Gilbert et al., 2020), but others place it as a stem hominid (e.g., Young and MacLatchy, 2004; Worthington, 2012; Begun et al., 2012; Pugh, 2022), and so it is not suitable for calibrating this node.
- Suggested prior distribution. Offset exponential Age justifications. We base our minimum age on the reported age of 12.3 Ma for another site in the mid-Chinji Formation, Y647 (which also preserves Sivapithecus indicus specimens), which is stated to be at the same stratigraphic level as Y494 (Morgan et al., 2015); this age is stated to be based on magnetostratigraphy, but Morgan et al. (2015) do not provide further details, and so it should be treated as tentative.
- We base the maximum bound on the maximum age of the oldest stem hominoid Rukwapithecus (see "Crown Catarrhini", "Crown Cercopithecidae", and "Crown Hominoidea" above). A potential alternative maximum bound would be to use the maximum age of the oldest hominoid Kenyapithecus, which is 14.0 Ma (see Crown Hominoidea above). However, given that a few published analyses have placed Kenyapithecus as a crown hominid (Nengo et al., 2017; Gilbert et al., 2020), it may be unduly restrictive to use this taxon to inform our maximum bound. We therefore choose a more conservative approach based on the age of Rukwapithecus, as this taxon has been consistently found to be a stem hominoid in recent phylogenetic analyses (see "Crown Catarrhini" and "Crown Cercopithecidae" above). We consider the offset exponential distribution to be most appropriate for this calibration, based on the same arguments given for the crown Hominoidea node (see "Crown Hominoidea" above). Assuming a 5% probability of exceeding the soft maximum bound, this would give a mean and median prior on this divergence of 16.6 and 15.3 Ma, respectively.
- Additional CladeAge calibration. We consider Sivapithecus indicus to be the oldest definitive pongine. Pugh (2022) presented a series of phylogenetic analyses of Miocene hominoids based on a large (41 taxa, 274 characters) morphological character matrix, using different character coding schemes and analytical methods. Most of these analyses found Nakalipithecus (described by Kunimatsu et al., 2007) to be the oldest (stem) hominine among the taxa included (see Pugh, 2022: figure 5); total evidence tip-dating analysis of the Pugh (2022) matrix in combination with DNA and protein sequence data also strongly supports Nakalipithecus as a stem hominine (Beck, in prep.). All known specimens of Nakalipithecus are from Upper Member of the Nakali Formation of Kenya, within Chron C5n.1n (Kunimatsu et al., 2007), which is 9.937-9.786 Ma (Raffi et al., 2020), and provides our additional CladeAge calibration here.
- Comments. Roos et al. (2019) used Kenyapithecus wickeri (with a maximum age of 14.9 Ma) as their maximum bound on this node, on the assumption that it is a stem hominid. The stem hominid position of Kenyapithecus has been supported by most recent phylogenetic analyses (e.g., Pugh, 2022), but it was placed as a crown hominid (pongine) by Nengo et al. (2017) and Gilbert et al. (2022), hence our decision to use the maximum age of the oldest stem hominoid Rukwapithecus fleaglei (see "Crown Hominoidea" above) for setting the soft maximum bound at 25.235 Ma. Benton et al. (2015), meanwhile, used a maximum of 33.9 Ma based on the age of the oldest known crown anthropoids from the L-41 Quarry of the Fayum Depression, Egypt (see "Crown Anthropoidea" above), which seems excessively conservative given the diversity of stem hominoids but absence of crown forms in the early Miocene African record (see "Crown Hominoidea" above).