Quaternary Science Reviews 140 (2016) 39e48 Contents lists available at ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev Size variation in Tachyoryctes splendens (East African mole-rat) and its implications for late Quaternary temperature change in equatorial East Africa * J. Tyler Faith a, , David B. Patterson b, Nick Blegen c, Chris J. O'Neill a, Curtis W. Marean d, e, Daniel J. Peppe f, Christian A. Tryon c a Archaeology Program, School of Social Science, University of Queensland, Brisbane, QLD 4072, Australia b Center for the Advanced Study of Human Paleobiology, Department of Anthropology, The George Washington University, Washington, DC 20052, USA c Department of Anthropology, Harvard University, Cambridge, MA 02138, USA d Institute of Human Origins, School of Human Evolution and Social Change, Arizona State University, Tempe, AZ 85287, USA e Centre for Coastal Palaeoscience, Nelson Mandela Metropolitan University, Port Elizabeth, Eastern Cape 6031, South Africa f Terrestrial Paleoclimatology Research Group, Department of Geosciences, Baylor University, Waco, TX 76798, USA article info abstract Article history: This study develops a new proxy for Quaternary temperature change in tropical Africa through analysis Received 4 September 2015 of size variation in East African mole-rat (Tachyoryctes splendens). In modern mole-rats, mandibular Received in revised form alveolar length is unrelated to annual precipitation, precipitation seasonality, temperature seasonality, or 1 March 2016 primary productivity. However, it is inversely correlated with mean annual temperature, in agreement Accepted 14 March 2016 with Bergmann's rule. This relationship is observed at temperatures below ~17.3 C, but not at higher Available online 28 March 2016 temperatures. We apply these observations to late Quaternary mole-rats from Wakondo (~100 ka) and Kisaaka (~50 ka) in the Lake Victoria region and Enkapune ya Muto (EYM; ~7.2e3.2 ka) in Kenya's central Keywords: Bergmann's rule rift. The Lake Victoria mole-rats are larger than expected for populations from warm climates typical of Enkapune ya Muto the area today, implying cooler temperatures in the past. The magnitude of temperature decline needed Karungu to drive the size shift is substantial (~4e6 C), similar in magnitude to the degree of change between the Lake Naivasha Last Glacial Maximum and Holocene, but is consistent with regional temperature records and with Lake Victoria scenarios linking equatorial African temperature to northern hemisphere summer insolation. Size Paleoclimate changes through time at EYM indicate that rising temperatures during the middle Holocene accompa- Paleoenvironment nied and potentially contributed to a decline in Lake Naivasha and expansion of grassland vegetation. Rusinga Island © 2016 Elsevier Ltd. All rights reserved. 1. Introduction regional climate histories and limits our understanding of the drivers of climate change in tropical Africa. Developing new climate East African records of paleoclimate and paleoenvironment are proxies provides one opportunity to rectify this situation. Such central to understanding the mechanisms underlying Quaternary work is not only important to understanding Quaternary climate climate change in tropical Africa (e.g., deMenocal, 1995; Trauth dynamics, but is also relevant to human origins research (e.g., et al., 2003; Verschuren et al., 2009). It is clear that orbital forcing deMenocal, 2004; Trauth et al., 2010; Blome et al., 2012). Paleo- mechanisms translate to different responses at high latitudes anthropologists have proposed links between changes in climate versus the tropics (Clement et al., 2004), but a limited number of and human biology, behavior, and biogeography (Potts, 1998; Vrba records from equatorial Africa prohibits the construction of et al., 1995; Blome et al., 2012; Potts and Faith, 2015), yet the details of these relationships are limited by few climate proxies that can be associated with archaeological and paleontological records (e.g., Blome et al., 2012) and by lack of a theoretically-grounded under- * Corresponding author. E-mail addresses: [email protected] (J.T. Faith), [email protected] standing of the relationships between climate, environment, and (D.B. Patterson), [email protected] (N. Blegen), [email protected] human populations (Behrensmeyer, 2006; Marean et al., 2015). (C.J. O'Neill), [email protected] (C.W. Marean), [email protected] Our aim here is to develop a new proxy of Quaternary climate (D.J. Peppe), [email protected] (C.A. Tryon). http://dx.doi.org/10.1016/j.quascirev.2016.03.017 0277-3791/© 2016 Elsevier Ltd. All rights reserved. 40 J.T. Faith et al. / Quaternary Science Reviews 140 (2016) 39e48 change through analysis of body size variation of modern and fossil temperature annual range, annual precipitation, and precipitation specimens of East African mole-rat (Tachyoryctes splendens). Among seasonality (coefficient of variation of monthly totals), were contemporary vertebrates, size clines across environmental gradi- extracted from the WorldClim global climate database (Hijmans ents are well-documented (e.g., Millien et al., 2006), potentially et al., 2005). We use the intermediate resolution 5 arc-minute tracking changes in temperature, seasonality, and primary pro- resolution climate layers, which average the climate signal from a ductivity, among other variables (e.g., Rosenzweig, 1968; James, broader area, to account for the lack of precise locality coordinates 1970; Boyce, 1978; Ashton et al., 2000; Gür, 2010). The best for some samples. In addition, we calculated annual net primary known of these relationships is Bergmann's Rule, which (broadly productivity (NPP), averaged from 2010 to 2014, from the MODIS defined) proposes that populations living in cooler climates are MOD17 data (Zhao et al., 2005) over a 5 km radius around each larger than their conspecifics in warmer climates (Meiri and Dayan, locality. 2003). Such relationships have long been studied by paleoecolo- Global climate change over the last ~100 years could contribute gists to reconstruct paleoclimate change in southern Africa from to a slight mismatch between mole-rat specimens, many collected body size shifts of fossil carnivores and various small mammals in the early 1900s, and their associated climate data. However, the (Avery, 1982, 2004; Klein, 1986, 1991; Klein and Cruz-Uribe, 1996). magnitude of such error is minor compared to magnitude of cli- Parallel research in East Africa is lacking, although Marean et al. matic variation between localities. For example, the IPCC (2015) (1994) document potentially climate-mediated size shifts in the reports an increase in average global temperature of ~0.85 C since mandibular alveolar lengths of Holocene Tachyoryctes splendens 1880, two orders of magnitude lower than the range of mean from Enkapune ya Muto in Kenya's Central Rift Valley. At the time, annual temperatures across modern localities (13.6 C). While dif- the lack of modern data relating alveolar length to climate pre- ferences in climate between today and when the specimens were cluded a definitive interpretation of these patterns. collected may contribute to a minor amount of analytical noise, this Tachyoryctes splendens is a solitary animal that is discontinu- should be swamped out by the variation between localities. ously distributed across portions of East Africa and Central Africa The modern localities encompass substantial variation in annual that receive >500 mm annual rainfall. It inhabits a wide range of precipitation (805e1943 mm/yr) and mean annual temperature environments, including tropical forests, open woodlands, and (10.0e23.6 C). Temperature variation is related to elevation gra- grasslands and occurs at altitudes up to ~4000 m, preferring well- dients (Table 1), as confirmed by a tight inverse correlation be- drained soils suitable for digging extensive burrows (Jarvis and tween mean annual temperature and elevation (r ¼0.982, Sale, 1971; Schlitter et al., 2008). Tachyoryctes is well represented p < 0.001). Other studies of size clines in mammals typically in the East African fossil record (Winkler et al., 2010) with the explore variation across broad latitudinal gradients (e.g., earliest records of the genus dating to the late Miocene in Ethiopia Rosenzweig, 1968; Klein, 1986; Koch, 1986), over which tempera- (Haile-Selassie et al., 2004; Wesselman et al., 2009) and extant ture, seasonality, and primary productivity often co-vary, making it T. splendens known from Late Pleistocene and Holocene sites in difficult to disentangle the factors driving size variation (see also Kenya and Tanzania (Mehlman, 1989; Marean, 1992b; Marean et al., Ashton et al., 2000; Gür, 2010). Our sample is restricted to the 1994; Gifford-Gonzalez, 1998; Faith et al., 2015). Together, the tropics and is characterized by a limited latitudinal range occurrence of T. splendens in diverse habitats with varied climate (5.55Se11.25N), with 16 of the 23 sample localities falling within regimes, its abundance in the fossil record, and previously docu- 1.5 of the equator. There is a significant, but weak, correlation mented size shifts make it an ideal candidate to explore body size- between the two variables related to seasonality (temperature climate relationships. We examine these relationships in modern annual range and precipitation seasonality: r ¼ 0.431, p ¼ 0.040), T. splendens and apply our results to address paleoclimate