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Microliths-small, retouched, often-backed stone tools-are often interpreted to be the product of composite tools, including projectile weapons, and efficient hunting strategies by modern humans. In Europe and Africa these lithic toolkits are linked to hunting of medium- and large-sized game found in grassland or woodland settings, or as adaptations to risky environments during periods of climatic change. Here, we report on a recently excavated lithic assemblage from the Late Pleistocene cave site of Fa-Hien Lena in the tropical evergreen rainforest of Sri Lanka. Our analyses demonstrate that Fa-Hien Lena represents the earliest microlith assemblage in South Asia (c. 48,000-45,000 cal. years BP) in firm association with evidence for the procurement of small to medium size arboreal prey and rainforest plants. Moreover, our data highlight that the lithic technology of Fa-Hien Lena changed little over the long span of human occupation (c. 48,000-45,000 cal. years BP to c. 4,000 cal. years BP) indicating a successful, stable technological adaptation to the tropics. We argue that microlith assemblages were an important part of the environmental plasticity that enabled Homo sapiens to colonise and specialise in a diversity of ecological settings during its expansion within and beyond Africa. The proliferation of diverse microlithic technologies across Eurasia c. 48-45 ka was part of a flexible human 'toolkit' that assisted our species' spread into all of the world's environments, and the development of specialised technological and cultural approaches to novel ecological situations., Competing Interests: The authors have declared that no competing interests exist.

Defining the distinctive capacities of Homo sapiens relative to other hominins is a major focus for human evolutionary studies. It has been argued that the procurement of small, difficult-to-catch, agile prey is a hallmark of complex behavior unique to our species; however, most research in this regard has been limited to the last 20,000 years in Europe and the Levant. Here, we present detailed faunal assemblage and taphonomic data from Fa-Hien Lena Cave in Sri Lanka that demonstrates specialized, sophisticated hunting of semi-arboreal and arboreal monkey and squirrel populations from ca. 45,000 years ago, in a tropical rainforest environment. Facilitated by complex osseous and microlithic technologies, we argue these data highlight that the early capture of small, elusive mammals was part of the plastic behavior of Homo sapiens that allowed it to rapidly colonize a series of extreme environments that were apparently untouched by its hominin relatives.

Batadomba-lena, a rockshelter in the rainforest of southwestern Sri Lanka, has yielded some of the earliest evidence of Homo sapiens in South Asia. H. sapiens foragers were present at Batadomba-lena from ca. 36,000 cal BP to the terminal Pleistocene and Holocene. Human occupation was sporadic before the global Last Glacial Maximum (LGM). Batadomba-lena's Late Pleistocene inhabitants foraged for a broad spectrum of plant and mainly arboreal animal resources (monkeys, squirrels and abundant rainforest snails), derived from a landscape that retained equatorial rainforest cover through periods of pronounced regional aridity during the LGM. Juxtaposed hearths, palaeofloors with habitation debris, postholes, excavated pits, and animal and plant remains, including abundant Canarium nutshells, reflect intensive habitation of the rockshelter in times of monsoon intensification and biome reorganisation after ca. 16,000 cal BP. This period corresponds with further broadening of the economic spectrum, evidenced though increased contribution of squirrels, freshwater snails and Canarium nuts in the diet of the rockshelter occupants. Microliths are more abundant and morphologically diverse in the earliest, pre-LGM layer and decline markedly during intensified rockshelter use on the wane of the LGM. We propose that changing toolkits and subsistence base reflect changing foraging practices, from shorter-lived visits of highly mobile foraging bands in the period before the LGM, to intensified use of Batadomba-lena and intense foraging for diverse resources around the site during and, especially, following the LGM. Traces of ochre, marine shell beads and other objects from an 80 km-distant shore, and, possibly burials reflect symbolic practices from the outset of human presence at the rockshelter. Evidence for differentiated use of space (individual hearths, possible habitation structures) is present in LGM and terminal Pleistocene layers. The record of Batadomba-lena demonstrates that Late Pleistocene pathways to (aspects of) behavioural 'modernity' (composite tools, practice of symbolism and ritual, broad spectrum economy) were diverse and ecologically contingent., (Copyright © 2011 Elsevier Ltd. All rights reserved.)