INTRODUCTION
When Charles Darwin visited the Falkland Islands in 1833, he noted the unusual occurrence of a single terrestrial mammal species:
Dusicyon australis, a fox-like canid known as the Falkland Islands wolf or “warrah” (
1). In his journals, Darwin observed the warrah’s lack of fear and its inquisitive nature, hypothesizing that “within a very few years … this fox will be classed with the dodo, as an animal which has perished from the face of the earth” (
2). By 1876, the warrah became the first canid to go extinct in the global historic period, as a result of overhunting (
3). While the warrah’s extinction is well documented, the nature and timing of its initial dispersal to the Falkland Islands has been the subject of speculation for centuries. Genetic evidence indicates that
D. australis diverged from
Dusicyon avus, a South American mainland species of fox, between 31 and 8 thousand years (ka) BP (before present;
4). When the first Europeans arrived in 1690, they found no evidence of people nor have any archaeological sites been discovered that support a pre-European human presence in the islands to date. It has thus been hypothesized that
D. australis evolved from populations of
D. avus that crossed the South Atlantic during the Last Glacial Maximum, when sea levels were ~125 m lower and the exposed continental shelf reduced the distance of a transoceanic crossing (
4). However, this hypothesis is undermined by a paucity of evidence for a sea-ice bridge, as well as the lack of any other native terrestrial mammals in the Falkland Islands (
4). Alternately, if
D. australis diverged from
D. avus in mainland South America, perhaps via domestication, then it could have arrived in the Falkland Islands during the Holocene via transport by Yaghan (Yámana) or other Indigenous peoples before European exploration of the South Atlantic (
1).
The position of the Falkland Islands relative to the nutrient-rich Falkland Current (
Fig. 1) could have facilitated the islands’ accessibility to capable seafaring peoples, who may have taken advantage of the current’s rich biodiversity as a food resource (
5). By 6400 BP, most of the Indigenous communities of the Beagle Channel and the Strait of Magellan subsisted primarily on marine resources (
6). Both vertebrate remains preserved in shell middens and historic ethnographic accounts indicate that Indigenous Fuegians had developed the capacity to hunt pinnipeds in open water from canoes (
6). Several lines of evidence indicate the possibility of a pre-European human presence in the Falkland Islands, including (i) anecdotal evidence of canoes during Darwin’s visit (
2), (ii) a previously reported increase in fossil charcoal centered at 4800 BP near the present-day town of Stanley (
7), (iii) reports of lithics and a bone harpoon (
8,
9), and (iv) the presence of the warrah, the Falkland Islands’ lone terrestrial mammal. In 1979, a local landowner found a stone projectile point on the north side of New Island (
Fig. 2). This artifact is made of locally sourced quartzite abundant throughout the Falkland Islands (
10) and is consistent with lithic technology used in Tierra del Fuego over the past 1000 years (
Fig. 2) (
8). However, in the absence of archaeological data, the origin of the stone point (and the canoes observed by Darwin in 1833) remained an enigma.
Establishing the timing of human arrival on islands can be challenging, as small or transient populations may not leave behind cultural material (
11). However, the paleoecological record can provide evidence of anthropogenic impacts on sensitive island ecosystems, including increases in fire activity, species introductions, or extinctions (
11–
13). Of these, fire has proven to be a particularly sensitive indicator of initial human activity in island ecosystems (
11). Sedimentary charcoal accumulation rates (CHAR) have been found to increase by orders of magnitude immediately following human arrival, ultimately declining to an elevated baseline relative to the preanthropogenic background (
11,
14,
15). By providing indirect proxy evidence of human impacts integrated across whole landscapes, the paleoecological record may be more sensitive than the archaeological record at detecting initial or ephemeral human activities on island ecosystems. To leverage this potential, we combined methods from paleoecology and archaeology to investigate a potential prehistoric human presence in the Falkland Islands. Our interdisciplinary investigation re-opens the question of a potential human prehistory of the Falkland Islands, which are an important biodiversity hotspot and the site of the first canid extinction in the global historic record.
RESULTS
In 2018, we conducted a ground surface survey on New Island in the area where a stone point made of locally sourced quartzite had been found on the surface by a local landowner in 1979. During the survey, we identified seven distinct deposits of disarticulated seabird and marine mammal bones just upslope of the location where the stone point was found (
Fig. 2). We excavated two of these deposits, which were primarily composed of
Otaria flavescens [South American sea lion; New Island bone pile 6 (NEWIBP6): minimum number of individuals (MNI) = 61; NEWIBP7: MNI = 73; table S3] and
Eudyptes chrysocome (southern rockhopper penguin; NEWIBP6: MNI = 9; NEWIBP7: MNI = 77; table S3). We dated 10 unique elements from these piles, representing 10
O. flavescens individuals (table S1). The New Island bone piles yielded dates between 675 and 530 BP (1275 to 1420 CE) [all dates reported as calibrated calendar years before present (1950), except when indicated as CE or otherwise noted].
We analyzed charcoal records from three sites for evidence of prehistoric anthropogenic burning: a peat column from New Island, a peat core from a bog on Mount Usborne (705 m; East Falkland), and a coastal peat column from Bleaker Island (
Fig. 1). Mount Usborne is the highest location in the Falkland Islands, located in the interior of East Falkland; its isolation, exposure, and distance to coastal resources make it an unlikely candidate for past human land use, and the site is not visited frequently today. We thus consider the Mount Usborne charcoal record to be a control sample representing the natural, climate-driven fire regime for the Falkland Islands. In contrast, Bleaker Island is a small (20.7 km
2) outlying island to the south of East Falkland. We included this location because Darwin noted several dugout canoes on a beach at Bleaker in 1833. He speculated that the Falklands Current was the most likely explanation for the presence of the canoes but did not rule out Indigenous visitors (
2).
The Mount Usborne fire history was relatively uniform (
Fig. 3), with low CHAR (<1 piece cm
−2 year
−1;
Fig. 3) throughout the 13,000-year record. The 15,000-year record at Bleaker Island is similar, with only a slight increase in CHAR and fire frequency at ~7000 BP. In contrast, the New Island charcoal record shows a marked increase in CHAR at ~1800 BP that is not seen at the other two sites (
Fig. 3). From 550 to 400 BP, CHAR abruptly increased by two orders of magnitude, contemporaneous with the radiocarbon distributions of the dated
O. flavescens bones (
Fig. 4). A similar abrupt increases in CHAR occurred from 180 to 120 BP during the initial arrival of Europeans in New Island, after which CHAR values were reduced but fire activity remained constant until the present day (
Fig. 4). The increases in CHAR at 550 and 180 BP contain the highest values of any of the records that we analyzed throughout the Falkland Islands and were several orders of magnitude greater than the maximum CHAR values from Mount Usborne or Bleaker Island (
Fig. 3), as well as from a previously published charcoal record from Surf Bay on East Falkland (
16).
To assess the possibility of a human introduction of the warrah, we analyzed six
D. australis elements from a shallow, seasonal pond at Spring Point Farm, West Falkland. Spring Point is the only location where fossil
D. australis bones have been found in the Falkland Islands despite large populations of the canids reported in historic accounts. The Falkland Islands are largely composed of acidic peat, which is poor for bone preservation and may account for the apparent scarcity of warrah fossils. Of the six samples sent for accelerator mass spectrometry (AMS) dating, only three yielded sufficient collagen for dating (table S1). Of these, a lower M1 tooth yielded the oldest known age for
D. australis, 3396–3752 BP (3790 ± 25
14C BP), which is currently the minimum arrival date of the warrah in the Falkland Islands. The other two
D. australis dates were 236–500 BP (895 ± 15
14C BP) and 264–511 BP (925 ± 15
14C BP) (table S1). Carbon and nitrogen isotopes suggest that the three
D. australis individuals that we analyzed had a marine-based diet consisting primarily of apex marine predators; δ
13C values were −9.6, −12.3, and −12.2 per mil (‰) (
Fig. 5 and table S1), and δ
15N values were 22, 17.3, and17.7 ‰, respectively. These δ
15N values place the warrah on par with sea lions, fur seals, and cetaceans, which range from 11 to 22‰ (
17–
19); typically, a 3 to 5‰ enrichment in δ
15N is observed with each increase in trophic level (
20). While these results may be consistent with selective coastal scavenging, they could also indicate a commensal relationship with humans, subsisting on littoral resources such as
O. flavescens and
E. chrysocome (
21).
DISCUSSION
Multiple independent lines of evidence from New Island indicate that people were in the Falkland Islands centuries before European colonization: an orders-of-magnitude increase in charcoal that overlaps in age with multiple mixed vertebrate bone deposits and a locally sourced stone point consistent with technology used by Indigenous Fuegian cultures. The earliest possible European sightings of the Falkland Islands date to the 16th century (
22), and the first confirmed landing was in 1690 CE (Supplementary Text) (
23). The Falkland Islands were not permanently settled until 1764 CE (
7); there are no recorded landings at New Island until 1774 CE (
24). Critically, while the earliest dates for European sighting of the Falkland Islands are debated, the first landing was not made until more than 250 years after the charcoal peaks and bone piles reported in this study (1275 to 1420 CE).
The fire history at New Island is consistent with that seen in other island systems following human arrival (
11), indicated by an orders-of-magnitude increase in charcoal (
Fig. 3). While fire can have natural or human origins, we interpret the increase in fire activity ~550 BP at New Island to be anthropogenic for several reasons. First, there are only two potential ignition sources in the Falkland Islands: lightning strikes and people. Fires caused by lightning strikes throughout the Falkland Islands are rare today (
7). As in Tierra del Fuego, sustained Westerlies counteract the strong convective currents that produce thunderstorms (although the strength and position of the Westerlies have shifted through time) (
25). An average of four thunderstorms per year (
7), coupled with low average temperatures (29° to −9°C) and low precipitation (642 mm/year) (
26), creates conditions in which lightning-caused fires are rare. Lightning is thus unlikely to produce the prolonged periods of high-frequency fire events seen in the New Island charcoal record, and it is unlikely that lightning strikes would be more common on New Island than on Mount Usborne or Bleaker Island. Furthermore, the observed increase in fire at New Island in the last millennium cannot be explained by changes in grassland productivity, temperature (
27), or precipitation (
28), none of which increased over the last millennium (fig. S1), so we rule these out as drivers.
The precontact increase in CHAR at 550 BP is similar in magnitude to the later increase at 1790 CE, when European and American sailors regularly used fire to improve visibility on New Island, as well as to flush seals and penguins for hunting and to render whale, seal, and penguin oil (
24,
29). This historic increase in CHAR demonstrates the sensitivity of our peat record to anthropogenic burning and provides a useful basis of comparison for the pre-European increase in CHAR (
Fig. 3). The gap in fire events between 550 and 180 BP is also consistent with the documented absence of human habitants in the Falkland Islands at the time of early European exploration of the region. While the earliest fire peaks at 1000 BP appear small compared to subsequent intervals, they represent an order-of-magnitude increase in CHAR over earlier values in the New Island record, when fires were rare and small by comparison, and may represent an earlier prehistoric human presence (
Figs. 3 and
4).
We also interpret the mixed New Island bone assemblages as anthropogenic for several reasons: First,
E. chrysocome is a prey species for
O. flavescens, and it is unlikely that these two species would be found together in abundance in natural death assemblages (table S3). Second, the conspicuous absence of
O. flavescens stomach contents, including fish bones or otoliths, cephalopod beaks, and bone fragments in the surrounding matrix, suggests that these bone piles were not the original death sites (
30). Furthermore, the piles are located on a 9° slope undergoing active erosion, with no depressions that would promote accumulation over time (
Fig. 2C). These features appear more consistent with a secondary processing or midden site rather than natural death assemblages. No cut marks or other evidence of butchering were found during our excavations. However, the absence of cut marks does not necessarily preclude processing by people (
30,
31). The presence of processing marks depend on several factors, including the skill of the butchers, carcass size, tool materials, the elements in an assemblage, and the taphonomy of the site (
31,
32). Preferential use of bone and shell tools by several Indigenous Fuegian cultures (
33) may explain the lack of butchery marks, as bone and mussel shell tools likely would have been too soft to leave easily observable cut marks on bone. It may also explain the absence of lithic debris across the site.
Of the several groups inhabiting Tierra del Fuego during the Holocene, the evidence at New Island is most consistent with culture of the Yaghan (Yámana) people, highly capable seafarers subsisting on a marine diet composed largely of seals, penguins, and shellfish (
33). They hunted sea lions and seals with bone harpoons hafted to wooden handles, and meat and skins were processed using mussel shells hafted to wooden handles (
33). While stone points were used by the Yaghan, they were used infrequently and typically only for arrows, which they sometimes used to hunt penguins (
33). Before European colonization, the Yaghan were highly mobile people without permanent settlements, and families (including canids) traveled together by dugout canoe as far as the Diego Ramírez Islands, located 105 km southwest of Cape Horn in the Drake Passage (
33). Fire was an integral part of Yaghan culture, whose fires may even have given Tierra del Fuego (“Land of Fire”) its name (
33). According to contemporary ethnographic accounts, the Yaghan wore little clothing despite the harsh climate and kept constant fires centrally located for warmth and cooking (
33). They brought fire with them when they traveled by sea, where a small hearth was stoked by children who crouched in the middle of their canoes (
33). The Yaghan also used signal fires to alert neighboring bands of whale carcasses or other important resources or when someone had died (
33). While there are no ethnographic accounts of the Yaghan using fire to flush game, as later Europeans did in the Falkland Islands (dense tussac grass is less common in Tierra del Fuego), it is possible that Indigenous visitors used fire as a hunting tool in the Falkland Islands. It is also possible that accidental burns resulted from smaller, intentional fires, as dry tussac grass and peat are highly flammable.
Based on ethnographic accounts of Yaghan people, it follows that evidence of their presence in the Falkland Islands would include (i) increased fire activity, (ii) utilization of coastal resources such as sea lions and penguins, and (iii) the presence of canids. The findings reported here are consistent with these expectations, although we are unable to determine whether these were intentional visits. We think it unlikely that prehistoric peoples had a sustained presence in the Falkland Islands. Rather, the evidence supports one or more temporary visitations or accidental landings that left behind little cultural material but resulted in a discernable anthropogenic fingerprint in the fossil record. While the evidence from New Island suggests a local arrival of people between 1000 and 550 BP, we cannot rule out earlier visits elsewhere across the Falkland Islands. Both the presence of the warrah on West Falkland by at least 3400 BP and an increase in charcoal abundance recorded in a previous peat record near Stanley from 5400 BP (
7) to ~3500 BP (
16) could indicate an earlier arrival.
The possibility of a prehistoric human presence in the Falkland Islands reopens the debate about the origin of the warrah and could explain the long-debated anomaly of a sole terrestrial mammal on an isolated archipelago. Southern South America has a long history of close human-canid interactions (
34–
36), including the extinct Fuegian dog [a domesticated form of the culpeo fox,
Lycalopex culpaeus (
36)], and foxes played an important role in Yaghan culture. According to ethnographic accounts, each family kept several canids, who routinely traveled in canoes and were never intentionally left behind when traveling (
33). The Fuegian dog bore a close morphological similarity to the culpeo and played an active role in the hunt for otters, birds, and foxes, as well as a protective role when unfamiliar humans were present. Even in times of resource scarcity, these canids were never eaten and their skins were never used, as their role in the family was more akin to hunting partner or companion (
33). Ethnographic accounts and recent genetic studies both suggest that
L. culpaeus underwent self-domestication through the construction of a new ecological niche shared with humans (
33,
36). While
D. australis was not closely related to
L. culpaeus, its closest relative,
D. avus, is the most commonly identified canid in Fuego-Patagonian archaeological sites until its extinction by ~400 BP (
3,
4,
34). At the Loma de los Muertos mortuary site, a
D. avus individual (2792 ± 50
14C BP) was found to be the primary subject of an interment, which suggests an intimate and mutualistic interspecies relationship (
35). It has been suggested that the warrah may have been a semi-domesticated form of
D. avus (
1), which could explain the observed docile and curious nature of the warrah described by Darwin and others, as well as some of the warrah’s physical traits (
2,
37).
When combined with the long history of human marine adaptation, semi-domestication of the warrah in mainland South America could help explain its presence in the Falkland Islands. The δ
15N values reported here reflect a marine diet at higher trophic levels, including seals, which the small canid was unlikely to have hunted. While the warrah may have been a selective coastal scavenger, their enriched δ
15N values could also have resulted from a mutualism with humans who were primarily subsisting on sea lions, penguins, and other marine predators (
21). Both pre- and post-contact hunter-gatherers from Tierra del Fuego had isotopic values (δ
13C = −12.8, δ
15N = 17.2;
Fig. 5), reflecting their specialization on marine resources from high tropic levels (
38,
39). In contrast, hunter-gatherers in the more interior Pampas region of northern Argentina exhibited much lower values (δ
13C = −17.4, δ
15N = 11.9;
Fig. 5) associated with more terrestrial diets (
40). While the dietary isotopic signatures of the warrah samples do not unequivocally prove human introduction, neither do they rule out a mutualism with people. Future studies of warrah remains collected during the European period, when abundant terrestrial food sources (e.g., sheep, cattle, horses, rabbits, pigs, rats, and mice) had been introduced to the islands, may provide further insight into the warrah’s dietary preferences and potential interaction with humans.
Ecological changes follow closely on the heels of human arrival, particularly in sensitive island ecosystems (although it is important to note that these impacts are not necessarily negative) (
41). Although our study did not directly quantify human impacts on native flora and fauna, our findings raise questions about the influence of fire, hunting, or the arrival of the warrah on the Falkland Islands’ flora and fauna. Oceanic islands are often hotspots of biodiversity, and many suffered extensive extinctions throughout the Holocene, which began during initial human settlement and accelerated during European colonization (
42). Ground-nesting birds were hit especially hard by Holocene extinctions, particularly because of the introduction of predators (
42,
43), so resolving the timing and origin of the warrah is especially relevant. Today, the Falkland Islands are globally important breeding grounds for many species of ground-nesting seabirds that are threatened by multiple global changes, including sea level rise, habitat loss, and climate change (
44). Further analysis of vertebrate fossil deposits, sedimentary archives (including ancient DNA), and historic museum specimens will be critical for establishing historic biodiversity baselines and assessing the long-term resilience of Falkland Islands biodiversity to global changes (
16).
A pre-European human presence in the Falkland Islands broadens our understanding of the human history of not only the Falkland Islands but also the South Atlantic more generally and highlights the skill and adaptability of southern South America’s adept seafaring peoples. New Island has many features that would have been attractive to seafarers (a sheltered sandy cove, fresh water, seabird, and pinniped colonies;
Fig. 1); these are shared with many other islands in the Falkland Islands and elsewhere. The highly mobile lifestyles and material cultures of people such as the Yaghan may mask their presence in the archaeological record, limiting our ability to fully recognize the complex histories of human migration. Our work supports interdisciplinary investigations where conspicuous evidence of human activity remains absent and highlights the utility of paleoecological proxies in identifying the indirect signatures of initial, ephemeral, or nomadic peopling. Future research may yield new potential sites in the Falkland Islands and beyond. Ideally, this work should be done in collaboration with regional Indigenous communities, whose oral histories and traditional ecological knowledge represent millennia of expertise in the prehistory of Tierra del Fuego and the South Atlantic.
Acknowledgments
We are grateful to the Falkland Islanders who lent expertise and graciously allowed access to field sites and samples: the Evans, Strange, Rendell, Smith, and Rowland families and S. Massam. We thank the Denver Museum of Nature and Science for providing historic O. flavescens samples. The South Atlantic Environmental Research Institute, New Island Conservation Trust, Falkland Islands Museum and National Trust, and Tony Smith of Discovery Falklands for providing invaluable logistical support. Columns were scanned at LacCore (National Lacustrine Core Facility), Department of Earth Sciences, University of Minnesota-Twin Cities. A. Cassels created the three-dimensional model of the stone point. We are grateful to three anonymous reviewers whose comments greatly improved this manuscript.
Funding: This material is based on work supported by the NSF Graduate Research Fellowship under grant no.1840992 to K.M.H., an Explorer’s Club Student Research Grant, a Rolex/Explorer’s Club grant, the Dan and Betty Churchill Exploration Fund, and >200 crowdfunders on
experiment.com. J.L.G. was supported by NSF CAREER grant EAR-1753186.
Author contributions: K.M.H., J.L.G., and D.H.S. conceptualized this study. K.M.H., J.L.G., D.V.G., K.E.K., J.R.S., T.V.L., and B.L.H. conducted field and laboratory analyses. K.M.H., J.L.G., D.V.G., K.E.K., B.L.H., D.H.S., J.R.S., P.B., and T.V.L. contributed to the interpretation and wrote the manuscript.
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 (LOI, charcoal, radiocarbon dates, and δ
15N and δ
13C isotopes) are present in the paper, the Supplementary Materials, and the Neotoma Paleoecology Database (
www.neotomadb.org/).
Sci. Adv.: Not Falklands: Islas Malvinas
The real name is Islas Malvinas