Southwest China, the last refuge of continental primates in East Asia

This study is based on two databases collected from a broad literature review: 1) Primate fossil locations (please see Appendix) discovered in the Late Miocene (11.6 – 5.3 mya), Pliocene (5.3 – 2.58 mya), Early Pleistocene (2.58 – 0.77 may), Middle Pleistocene (0.77– 0.129 mya) and Late Pleistocene (0.129 – 0.0117 mya). However, as the fossil sites of the catarrhines in the Pliocene are very scarce, especially regarding the Pongidae and Hylobatidae (Harrison 2016), the reconstruction of the dispersal scenarios focused on the Pleistocene. 2) Pongidae was extinct in the Late Pleistocene, so later information in the Holocene comes from historical geographic distributions of the existing Colobinae (from 1304), Cercopithecinae (from 1175), and Hylobatidae (from 1182): a remarkable 800-year period. Such information is from the records of Chinese local governmental historical archives, annals, journals, and books; more details have been given in our previous publications: (Li et al., 2020; Li et al., 2002). Public records categorize catarrhines into four different groups: 1) gibbons, called “Yuan” in Chinese, are characterized by long upper limbs, slim body structure, unique calls, and acrobatic behavior (Zhang, 2015; Zhou and Zhang, 2013). Their records appeared since the Zhou Dynasty (1027-221 BC) in art, mainly in paintings (Geissmann, 2008), and other social services, such as funeral services, being buried with the dead (Vogel, 2018); 2) snub-nosed monkeys (Rhinopithecus) with up-turned nostrils and sturdy body structure; 3) leaf-eating monkeys (Presbytis, Trachypithecus, Pygathrix, and Semnopithecus), showing slender body structure with typical arboreal lifestyle and special folivorous diets; and 4) macaques that are primarily terrestrial and morphologically quite different from the other groups (Li et al., 2020). Among the four, 1) belongs to the Hylobatidae (gibbons), containing Nomascus (4 species), Hylobates (1 species), and Hoolock (1 species); 2) and 3) are classified as the Colobinae (colobines), including snub-nosed monkeys (Rhinopithecus, five species) and langurs (Trachypithecus, six species; Pygathrix, one species; and Semnopithecus, one species); and 4) is regarded as the Cercopithecinae (Macaca), consisting eight species (Li et al., 2018).

本研究基于两项通过广泛文献调研采集的数据库：1）晚中新世（11.6–5.3百万年前）、上新世（5.3–2.58百万年前）、早更新世（2.58–0.77百万年前）、中更新世（0.77–0.129百万年前）及晚更新世（0.129–0.0117百万年前）发现的灵长类化石点（详见附录）。但由于上新世狭鼻类（catarrhines）的化石遗址极为匮乏，尤其是人猿科（Pongidae）和长臂猿科（Hylobatidae）（Harrison 2016），因此本研究的扩散场景重建聚焦于更新世时期。2）人猿科已于晚更新世灭绝，因此全新世的后续分布数据来自现存疣猴亚科（Colobinae，记录始于公元1304年）、猕猴亚科（Cercopithecinae，记录始于公元1175年）以及长臂猿科（Hylobatidae，记录始于公元1182年）的历史地理分布——这一时段跨度长达800年。此类信息源自中国地方官府历史档案、地方志、期刊与典籍的记载；更多细节已在我们此前的发表成果中详述：(Li et al., 2020; Li et al., 2002)。公共记录将狭鼻类划分为四大类群：1）长臂猿，中文俗称“猿”，以上肢修长、躯体纤细、鸣声独特且兼具杂技般的行为特征为典型（Zhang, 2015; Zhou and Zhang, 2013）。其相关记载最早可追溯至西周时期（公元前1027年-公元前221年），多见于绘画等艺术形式中（Geissmann, 2008），亦见于丧葬等社会习俗中，如作为随葬品（Vogel, 2018）；2）仰鼻猴属（Rhinopithecus），以鼻孔上翻、躯体强健为特征；3）叶猴类（包括Presbytis、Trachypithecus、Pygathrix及Semnopithecus），躯体纤细，营典型树栖生活，且以特殊的叶食性为食性特征；4）猕猴属（Macaca），主要营陆地生活，形态与其他类群差异显著（Li et al., 2020）。在这四大类群中，1）隶属于长臂猿科（Hylobatidae），包含Nomascus（4种）、Hylobates（1种）及Hoolock（1种）；2）与3）归为疣猴亚科（Colobinae），涵盖仰鼻猴属（Rhinopithecus，共5种）及叶猴类（Trachypithecus，6种；Pygathrix，1种；Semnopithecus，1种）；4）则属于猕猴亚科（Cercopithecinae），包含8个物种（Li et al., 2018）。