INTRODUCTION
South Asia, and Sri Lanka more specifically, has emerged as a particularly important region for understanding how our species managed to successfully colonize a wide variety of environments among a backdrop of changing climates and interhominin contacts (
1–
4). As early as the 1980s, it was proposed that microlith technologies alongside bone technologies and ochre use appeared in Sri Lanka earlier than they did in Europe (
5). Despite these suggestions, and the recognized importance of the region for human evolutionary studies, detailed studies of the material culture recovered from its most ancient sites have thus far been lacking, particularly with regard to postulated personal ornaments (
6–
7), potential projectile technologies (
4,
8), and other forms of material culture that provide insight into how human societies negotiated the South Asian tropics during the Late Pleistocene. It is only recently that multidisciplinary excavation of long, well-dated cave and rockshelter sequences is enabling these questions to be investigated and the findings to be compared to records from more customarily discussed regions (
9).
As the site of the earliest fossil appearance of
Homo sapiens in South Asia (
5), Fa-Hien Lena cave in southwestern Sri Lanka is a crucial locale for understanding the adaptive capacities and cultural flexibility that humans required as they first moved throughout the diverse environments of Asia. Ongoing analysis of the site has already found that it holds the earliest microlith assemblage in the region (
9) and attests to targeted hunting of prime-aged semi-arboreal and arboreal monkeys and squirrels (
4). Here, we present evidence for the earliest use of bow-and-arrow technology outside of Africa—a unique tradition using innovative osseous-based arrow heads. Also described is a diverse toolkit of bone and tooth tools indicating the manufacture of plant- and/or hide-based items that may represent some of the earliest clothing or nets in a tropical setting, alongside a complex array of symbolic artifacts—this record stretching from c. 48 ka (thousand years) through to c. 4 ka BP (before present) (
Fig. 1).
DISCUSSION
The proliferation of parietal and portable art along with the dynamic change in technological complexity during the European Upper Paleolithic (c. 45 to 11 ka BP) has been held up as a “gold standard” in human cultural development (
37–
40). In Western and Central Europe, tailored clothing, osseous technologies, and figurative art have been argued to enable resilience in the face of cold climates and growing populations (
41). More recently, finds in Africa placed the earliest evidence for symbolism and extensive social signaling alongside sophisticated production of blades for projectile technologies in the coastal contexts of southern and northern Africa c. 100 to 80 ka (
42–
44). Long traditions of archaeological excavation and prioritization in these regions have meant that they eclipse other parts of Africa, Asia, Australasia, and the Americas in quantity of archaeological residues reported and, consequently, discussions of the origins and adaptive context of material culture characteristic of our species. This trend has prevailed despite the fact we now know that by the end of the Late Pleistocene,
H. sapiens populations were occupying arid environments in Southern Africa and Australia (
45,
46), paleoarctic settings in Siberia (
47), high-altitude environments on the Tibetan Plateau (
48), and, most significantly here, tropical forests in Africa (
49), Southeast Asia (
50), Melanesia (
51), and Sri Lanka.
By providing the first in-depth traceological insights into the use of organic technologies in the Wet Zone of Pleistocene Sri Lanka, we have identified different types of pointed bone technologies in the record of Fa-Hien Lena—including notched unipoints and bipoints that likely tipped arrows, as well as geometric points that were potentially used in freshwater fishing. While the use of fiber at the site may also have supported the application of trapping in hunting arboreal and freshwater resources, it is now definitively evident that the majority of the osseous technologies were used in the high-velocity projectile hunting of small semi-arboreal and arboreal game. This finding is notable in a global context as it represents the earliest definitive evidence for high-powered projectile hunting in a tropical rainforest environment at 48 ka BP—and the earliest evidence for bow-and-arrow technology outside of Africa (
52). While the use of high-velocity projectile weaponry has been tentatively suggested at 32 ka BP at Lobang Hangus (Niah Caves, Sarawak), more definitive evidence in Southeast Asia is limited to the Terminal Pleistocene and Holocene but concentrated around the Pleistocene-Holocene boundary (
53,
54).
The Fa-Hien Lena osseous technologies share a number of commonalities with that recovered from the similarly aged Niah Caves contexts some 3000 km away in Sarawak, for which a tropical rainforest environment has also been reconstructed during the Late Pleistocene (
55,
56). In particular, the use of fine grinding to shape the artifacts and the use of pointed long bones leaving the diaphysis intact are recorded at both sites, as is the dominance of projectile points over other osseous tool forms in the assemblages. Teeth were used as tools at both locations (suids at Niah and Cercopithecid and
Muntiacus at Fa-Hien Lena). Differences are also apparent, however, the most important being that while the tools at Fa-Hien Lena were almost exclusively made on Cercopithecid bone (with the exceptions of the few made on large mammal bone from the Terminal Pleistocene), at Niah, the tools were manufactured mostly from large or intermediate-sized mammals (
55,
56).
We also present the first high-resolution analysis of symbolic material culture from a Late Pleistocene site in tropical Sri Lanka. Microscopic analysis has demonstrated that
H. sapiens populations in this part of the world—as those recorded in Africa and Eurasia at this same period—continued to value mineral colorants and marine shell beads for social signaling practices. Unique to this site, however, is the heavy usage of silver (mica) pigments alongside both bright red and bright yellow. While the collection and use of silver pigments are noted for other early
H. sapiens sites in Asia and Australasia, such as Madjedbebe (
57), as far as we are aware, we report the first instance of beads made from ochre nodules. The use of both ochre and marine shells for ornamentation purposes is demonstrated for
H. sapiens in the Middle and Later Stone Ages of Africa from c. 100 ka BP (
58), from c. 45 ka BP in Eurasia (
36), and from 45 ka BP in Island Southeast Asia (
59). It is noteworthy that, yet again, we find light-colored shells and colorants in bright hues forming the backbone of the symbolling system. Given the stable isotope evidence that highlights the reliance of humans living at the Late Pleistocene Wet Zone rainforest sites on forest resources (
2,
3), the attainment of marine shells, which were then elaborated, also highlights the significance of exchange networks between populations on different parts of the island. Alongside later, Terminal Pleistocene evidence for shark teeth and marine shell beads at Fa-Hien Lena (
6) and Balangoda Kuragala (
2), this initial finding implies the development of long-distance trading networks between populations exploiting the interior rainforests and groups located on the coast.
It is becoming increasingly apparent that if we are to understand the technology, symbolic material culture developments, social contexts, and adaptive strategies that accompanied our species during its expansion within and beyond Africa, it is important to study records beyond the traditional heartlands of paleoanthropological and Paleolithic archaeological research (
59–
63). Notably, we have demonstrated that elements such as clothing and fiber traps, projectile technologies, and symbolic networks, once thought to be characteristically early in Europe and Africa (
39,
40), were present among early human populations at the tip of South Asia inhabiting a tropical rainforest environment. Recent research has highlighted the nonlinear and divergent way in which traits of supposed “behavioral modernity” among our species appeared (
63–
65). Yet, if we are to move away from essentialist ideas of “modernity,” toward understandings of “variability” (
63), it is also essential to determine the diverse environmental contexts in which personal ornamentation, projectile technologies, and long-distance social networks emerged in the Pleistocene, not just in Africa but also beyond. Only then can we fully understand the adaptive contexts behind these behaviors and the nature of their application in the unique diversity of environments that our species was to colonize by the start of the Holocene.
Acknowledgments
We thank C. Thero, A. Thero, and the other monks at the Fa-Hien Lena Cave temple. We also thank S. Disanayaka, P. B. Mandawala, S. A. T. G. Priyantha, L. V. A. De. Mel, and other members at the excavation Branch of the Department of Archaeology, Sri Lanka. Funding: This research was supported by the Research Council of the University of Sri Jayewardenepura (RC-USJ) and the Max Planck Society. Author contributions: M.C.L., N.A., O.W., M.D.P., N.B., and P.R. designed the research; O.W., N.A., M.C.L., S.D., N.B., and M.M.P. collected the data; O.W., N.A., and M.C.L. analyzed the data; and M.C.L., N.A., O.W., S.D., M.M.P., N.P., N.B., M.D.P., and P.R. wrote the paper. Competing interests: The authors declare that they have no competing interests. Data and materials availability: All data needed to evaluate the conclusions in the paper are present in the paper and/or the Supplementary Materials. Additional data related to this paper may be requested from the authors. The artifacts from the Fa-Hien Lena excavations are curated at the Department of Archaeology, Government of Sri Lanka, under the site code BYP and the suffixes 10, 11, and 12 (denoting the year of excavation).