Late-surviving megafauna in Tasmania, Australia, implicate human involvement in their extinction

Establishing the cause of past extinctions is critical if we are to understand better what might trigger future occurrences and how to prevent them. The mechanisms of continental late Pleistocene megafaunal extinction, however, are still fiercely contested. Potential factors contributing to their demise include climatic change, human impact, or some combination. On the Australian mainland, 90% of the megafauna became extinct by ≈46 thousand years (ka) ago, soon after the first archaeological evidence for human colonization of the continent. Yet, on the neighboring island of Tasmania (which was connected to the mainland when sea levels were lower), megafaunal extinction appears to have taken place before the initial human arrival between 43 and 40 ka, which would seem to exonerate people as a contributing factor in the extirpation of the island megafauna. Age estimates for the last megafauna, however, are poorly constrained. Here, we show, by direct dating of fossil remains and their associated sediments, that some Tasmanian megafauna survived until at least 41 ka (i.e., after their extinction on the Australian mainland) and thus overlapped with humans. Furthermore, a vegetation record for Tasmania spanning the last 130 ka shows that no significant regional climatic or environmental change occurred between 43 and 37 ka, when a land bridge existed between Tasmania and the mainland. Our results are consistent with a model of human-induced extinction for the Tasmanian megafauna, most probably driven by hunting, and they reaffirm the value of islands adjacent to continental landmasses as tests of competing hypotheses for late Quaternary megafaunal extinctions.     

Synchronous extinction of North America's Pleistocene mammals

The late Pleistocene witnessed the extinction of 35 genera of North American mammals. The last appearance dates of 16 of these genera securely fall between 12,000 and 10,000 radiocarbon years ago (≈13,800–11,400 calendar years B.P.), although whether the absence of fossil occurrences for the remaining 19 genera from this time interval is the result of sampling error or temporally staggered extinctions is unclear. Analysis of the chronology of extinctions suggests that sampling error can explain the absence of terminal Pleistocene last appearance dates for the remaining 19 genera. The extinction chronology of North American Pleistocene mammals therefore can be characterized as a synchronous event that took place 12,000–10,000 radiocarbon years B.P. Results favor an extinction mechanism that is capable of wiping out up to 35 genera across a continent in a geologic instant.     

Megafaunal meiolaniid horned turtles survived until early human settlement in Vanuatu,Southwest Pacific

Meiolaniid or horned turtles are members of the extinct Pleistocene megafauna of Australia and the southwest Pacific. The timing and causes of their extinction have remained elusive. Here we report the remains of meiolaniid turtles from cemetery and midden layers dating 3,100/3,000 calibrated years before present to approximately 2,900/2,800 calibrated years before present in the Teouma Lapita archaeological site on Efate in Vanuatu. The remains are mainly leg bones; shell fragments are scant and there are no cranial or caudal elements, attesting to off-site butchering of the turtles. The new taxon differs markedly from other named insular terrestrial horned turtles. It is the only member of the family demonstrated to have survived into the Holocene and the first known to have become extinct after encountering humans.     

The shifting baseline of northern fur seal ecology in the northeast Pacific Ocean

Historical data provide a baseline against which to judge the significance of recent ecological shifts and guide conservation strategies, especially for species decimated by pre-20th century harvesting. Northern fur seals (NFS; Callorhinus ursinus) are a common pinniped species in archaeological sites from southern California to the Aleutian Islands, yet today they breed almost exclusively on offshore islands at high latitudes. Harvest profiles from archaeological sites contain many unweaned pups, confirming the presence of temperate-latitude breeding colonies in California, the Pacific Northwest, and the eastern Aleutian Islands. Isotopic results suggest that prehistoric NFS fed offshore across their entire range, that California populations were distinct from populations to the north, and that populations breeding at temperate latitudes in the past used a different reproductive strategy than modern populations. The extinction of temperate-latitude breeding populations was asynchronous geographically. In southern California, the Pacific Northwest, and the eastern Aleutians, NFS remained abundant in the archaeological record up to the historical period approximately 200 years B.P.; thus their regional collapse is plausibly attributed to historical hunting or some other anthropogenic ecosystem disturbance. In contrast, NFS populations in central and northern California collapsed at approximately 800 years B.P., long before European contact. The relative roles of human hunting versus climatic factors in explaining this ecological shift are unclear, as more paleoclimate information is needed from the coastal zone.     

Prehistoric birds from New Ireland, Papua New Guinea: extinctions on a large Melanesian island

At least 50 species of birds are represented in 241 bird bones from five late Pleistocene and Holocene archaeological sites on New Ireland (Bismarck Archipelago, Papua New Guinea). The bones include only two of seabirds and none of migrant shorebirds or introduced species. Of the 50 species, at least 12 (petrel, hawk, megapode, quail, four rails, cockatoo, two owls, and crow) are not part of the current avifauna and have not been recorded previously from New Ireland. Larger samples of bones undoubtedly would indicate more extirpated species and refine the chronology of extinction. Humans have lived on New Ireland for ca. 35,000 years, whereas most of the identified bones are 15,000 to 6,000 years old. It is suspected that most or all of New Ireland's avian extinction was anthropogenic, but this suspicion remains undetermined. Our data show that significant prehistoric losses of birds, which are well documented on Pacific islands more remote than New Ireland, occurred also on large, high, mostly forested islands close to New Guinea.     

Younger Dryas "black mats" and the Rancholabrean termination in North America

Of the 97 geoarchaeological sites of this study that bridge the Pleistocene-Holocene transition (last deglaciation), approximately two thirds have a black organic-rich layer or “black mat” in the form of mollic paleosols, aquolls, diatomites, or algal mats with radiocarbon ages suggesting they are stratigraphic manifestations of the Younger Dryas cooling episode 10,900 B.P. to 9,800 B.P. (radiocarbon years). This layer or mat covers the Clovis-age landscape or surface on which the last remnants of the terminal Pleistocene megafauna are recorded. Stratigraphically and chronologically the extinction appears to have been catastrophic, seemingly too sudden and extensive for either human predation or climate change to have been the primary cause. This sudden Rancholabrean termination at 10,900 ± 50 B.P. appears to have coincided with the sudden climatic switch from Allerød warming to Younger Dryas cooling. Recent evidence for extraterrestrial impact, although not yet compelling, needs further testing because a remarkable major perturbation occurred at 10,900 B.P. that needs to be explained.     

Prolonged coexistence of humans and megafauna in Pleistocene Australia

Recent claims for continent wide disappearance of megafauna at 46.5 thousand calendar years ago (ka) in Australia have been used to support a "blitzkrieg" model, which explains extinctions as the result of rapid overkill by human colonizers. A number of key sites with megafauna remains that significantly postdate 46.5 ka have been excluded from consideration because of questions regarding their stratigraphic integrity. Of these sites, Cuddie Springs is the only locality in Australia where megafauna and cultural remains are found together in sequential stratigraphic horizons, dated from 36-30 ka. Verifying the stratigraphic associations found here would effectively refute the rapid-overkill model and necessitate reconsideration of the regional impacts of global climatic change on megafauna and humans in the lead up to the last glacial maximum. Here, we present geochemical evidence that demonstrates the coexistence of humans and now-extinct megafaunal species on the Australian continent for a minimum of 15 ka.     

Colloquium paper: Megafauna biomass tradeoff as a driver of Quaternary and future extinctions

Earth's most recent major extinction episode, the Quaternary Megafauna Extinction, claimed two-thirds of mammal genera and one-half of species that weighed >44 kg between approximately 50,000 and 3,000 years ago. Estimates of megafauna biomass (including humans as a megafauna species) for before, during, and after the extinction episode suggest that growth of human biomass largely matched the loss of non-human megafauna biomass until approximately 12,000 years ago. Then, total megafauna biomass crashed, because many non-human megafauna species suddenly disappeared, whereas human biomass continued to rise. After the crash, the global ecosystem gradually recovered into a new state where megafauna biomass was concentrated around one species, humans, instead of being distributed across many species. Precrash biomass levels were finally reached just before the Industrial Revolution began, then skyrocketed above the precrash baseline as humans augmented the energy available to the global ecosystem by mining fossil fuels. Implications include (i) an increase in human biomass (with attendant hunting and other impacts) intersected with climate change to cause the Quaternary Megafauna Extinction and an ecological threshold event, after which humans became dominant in the global ecosystem; (ii) with continued growth of human biomass and today's unprecedented global warming, only extraordinary and stepped-up conservation efforts will prevent a new round of extinctions in most body-size and taxonomic spectra; and (iii) a near-future biomass crash that will unfavorably impact humans and their domesticates and other species is unavoidable unless alternative energy sources are developed to replace dwindling supplies of fossil fuels.     

Explaining the Pleistocene megafaunal extinctions: models,chronologies, and assumptions

Understanding of the Pleistocene megafaunal extinctions has been advanced recently by the application of simulation models and new developments in geochronological dating. Together these have been used to posit a rapid demise of megafauna due to over-hunting by invading humans. However, we demonstrate that the results of these extinction models are highly sensitive to implicit assumptions concerning the degree of prey naivety to human hunters. In addition, we show that in Greater Australia, where the extinctions occurred well before the end of the last Ice Age (unlike the North American situation), estimates of the duration of coexistence between humans and megafauna remain imprecise. Contrary to recent claims, the existing data do not prove the "blitzkrieg" model of overkill.     

Directly dated starch residues document early formative maize (Zea mays L.) in tropical Ecuador

The study of maize (Zea mays L.) domestication has advanced from questions of its origins to the study-and debate-of its dietary role and the timing of its dispersal from Mexico. Because the investigation of maize's spread is hampered by poor preservation of macrobotanical remains in the Neotropics, research has focused on microbotanical remains whose contexts are often dated by association, leading some to question the dates assigned. Furthermore, some scholars have argued that maize was not introduced to southwestern Ecuador until approximately 4150-3850 calendar years before the present (cal B.P.), that it was used first and foremost as a fermented beverage in ceremonial contexts, and that it was not important in everyday subsistence, challenging previous studies based on maize starch and phytoliths. To further investigate these questions, we analyzed every-day cooking vessels, food-processing implements, and sediments for starch and phytoliths from an archaeological site in southwestern Ecuador constituting a small Early Formative village. Employing a new technique to recover starch granules from charred cooking-pot residues we show that maize was present, cultivated, and consumed here in domestic contexts by at least 5300-4950 cal B.P. Directly dating the residues by accelerator mass spectrometry (AMS) radiocarbon measurement, our results represent the earliest direct dates for maize in Early Formative Ecuadorian sites and provide further support that, once domesticated approximately 9000 calendar years ago, maize spread rapidly from southwestern Mexico to northwestern South America.     

Rapid prehistoric extinction of iguanas and birds in Polynesia

The Tongoleleka archaeological site on Lifuka Island, Kingdom of Tonga, is a rich accumulation of pottery, marine mollusks, and nonhuman bones that represents first human contact on a small island in Remote Oceania approximately 2,850 years ago. The lower strata contain decorated Lapita-style pottery and bones of an extinct iguana (Brachylophus undescribed sp.) and numerous species of extinct birds. The upper strata instead feature Polynesian Plainware pottery and bones of extant species of vertebrates. A stratigraphic series of 20 accelerator-mass spectrometer radiocarbon dates on individual bones of the iguana, an extinct megapode (Megapodius alimentum), and the non-native chicken (Gallus gallus) suggests that anthropogenic loss of the first two species and introduction of the latter occurred on Lifuka within a time interval too short (a century or less) to be resolved by radiometric dating. The geologically instantaneous prehistoric collapse of Lifuka's vertebrate community contrasts with the much longer periods of faunal depletion on some other islands, thus showing that the elapse time between human arrival and major extinction events was highly variable on oceanic islands as well as on continents.     

Global archaeological evidence for proboscidean overkill

One million years ago, proboscideans occupied most of Africa, Europe, Asia, and the Americas. Today, wild elephants are only found in portions of sub-Saharan Africa and South Asia. Although the causes of global Pleistocene extinctions in the order Proboscidea remain unresolved, the most common explanations involve climatic change and/or human hunting. In this report, we test the overkill and climate-change hypotheses by using global archaeological spatiotemporal patterning in proboscidean kill/scavenge sites. Spanning approximately 1.8 million years, the archaeological record of human subsistence exploitation of proboscideans is preferentially located on the edges of the human geographic range. This finding is commensurate with global overkill, suggesting that prehistoric human range expansion resulted in localized extinction events. In the present and the past, proboscideans have survived in refugia that are largely inaccessible to human populations.     

Paleoindian demography and the extraterrestrial impact hypothesis

Recently it has been suggested that one or more large extraterrestrial (ET) objects struck northern North America 12,900 ± 100 calendar years before present (calBP) [Firestone RB, et al. (2007) Proc Natl Acad Sci USA 104: 16016–16021]. This impact is claimed to have triggered the Younger Dryas major cooling event and resulted in the extinction of the North American megafauna. The impact is also claimed to have caused major cultural changes and population decline among the Paleoindians. Here, we report a study in which ≈1,500 radiocarbon dates from archaeological sites in Canada and the United States were used to test the hypothesis that the ET resulted in population decline among the Paleoindians. Following recent studies [e.g., Gamble C, Davies W, Pettitt P, Hazelwood L, Richards M (2005) Camb Archaeol J 15:193–223), the summed probability distribution of the calibrated dates was used to identify probable changes in human population size between 15,000 and 9,000 calBP. Subsequently, potential biases were evaluated by modeling and spatial analysis of the dated occupations. The results of the analyses were not consistent with the predictions of extraterrestrial impact hypothesis. No evidence of a population decline among the Paleoindians at 12,900 ± 100 calBP was found. Thus, minimally, the study suggests the extraterrestrial impact hypothesis should be amended.     

Paleoclimate and Amerindians: evidence from stable isotopes and atmospheric circulation

Two Amerindian demographic shifts are attributed to climate change in the northwest plains of North America: at approximately 11,000 calendar years before present (yr BP), Amerindian culture apparently split into foothills-mountains vs. plains biomes; and from 8,000-5,000 yr BP, scarce archaeological sites on the open plains suggest emigration during xeric "Altithermal" conditions. We reconstructed paleoclimates from stable isotopes in prehistoric bison bone and relations between weather and fractions of C(4) plants in forage. Further, we developed a climate-change model that synthesized stable isotope, existing qualitative evidence (e.g., palynological, erosional), and global climate mechanisms affecting this midlatitude region. Our isotope data indicate significant warming from approximately 12,400 to 11,900 yr BP, supporting climate-driven cultural separation. However, isotope evidence of apparently wet, warm conditions at 7,300 yr BP refutes emigration to avoid xeric conditions. Scarcity of archaeological sites is best explained by rapid climate fluctuations after catastrophic draining of the Laurentide Lakes, which disrupted North Atlantic Deep Water production and subsequently altered monsoonal inputs to the open plains.     

Dating the late prehistoric dispersal of Polynesians to New Zealand using the commensal Pacific rat

The pristine island ecosystems of East Polynesia were among the last places on Earth settled by prehistoric people, and their colonization triggered a devastating transformation. Overhunting contributed to widespread faunal extinctions and the decline of marine megafauna, fires destroyed lowland forests, and the introduction of the omnivorous Pacific rat (Rattus exulans) led to a new wave of predation on the biota. East Polynesian islands preserve exceptionally detailed records of the initial prehistoric impacts on highly vulnerable ecosystems, but nearly all such studies are clouded by persistent controversies over the timing of initial human colonization, which has resulted in proposed settlement chronologies varying from approximately 200 B.C. to 1000 A.D. or younger. Such differences underpin radically divergent interpretations of human dispersal from West Polynesia and of ecological and social transformation in East Polynesia and ultimately obfuscate the timing and patterns of this process. Using New Zealand as an example, we provide a reliable approach for accurately dating initial human colonization on Pacific islands by radiocarbon dating the arrival of the Pacific rat. Radiocarbon dates on distinctive rat-gnawed seeds and rat bones show that the Pacific rat was introduced to both main islands of New Zealand approximately 1280 A.D., a millennium later than previously assumed. This matches with the earliest-dated archaeological sites, human-induced faunal extinctions, and deforestation, implying there was no long period of invisibility in either the archaeological or palaeoecological records.     

Life and extinction of megafauna in the ice-age Arctic

Understanding the population dynamics of megafauna that inhabited the mammoth steppe provides insights into the causes of extinctions during both the terminal Pleistocene and today. Our study area is Alaska''s North Slope, a place where humans were rare when these extinctions occurred. After developing a statistical approach to remove the age artifacts caused by radiocarbon calibration from a large series of dated megafaunal bones, we compare the temporal patterns of bone abundance with climate records. Megafaunal abundance tracked ice age climate, peaking during transitions from cold to warm periods. These results suggest that a defining characteristic of the mammoth steppe was its temporal instability and imply that regional extinctions followed by population reestablishment from distant refugia were characteristic features of ice-age biogeography at high latitudes. It follows that long-distance dispersal was crucial for the long-term persistence of megafaunal species living in the Arctic. Such dispersal was only possible when their rapidly shifting range lands were geographically interconnected. The end of the last ice age was fatally unique because the geographic ranges of arctic megafauna became permanently fragmented after stable, interglacial climate engendered the spread of peatlands at the same time that rising sea level severed former dispersal routes.One of the most intriguing examples of mass extinction and the most accessible in terms of its geological record occurred around the end of the Wisconsin ice age ca. 10–45 calendar ka B.P. (10,000–45,000 calendar y ago) when some 65% of terrestrial megafauna genera (animals weighing >45 kg) became globally extinct (1). Based on what we know about recent species extinctions, the causes of extinction are usually synergistic, often species-specific, and therefore, complex, which implies that there is no universal explanation for end-Pleistocene extinctions (2, 3). Globally and specifically in the Arctic (3–10), megafaunal extinctions have been variously blamed on overhunting, rapid climate change, habitat loss, and introduced diseases (3–10). Further complicating a clear understanding of the causes of ice-age extinctions is that the magnitude and tempo of environmental change during the last 100,000 y of the Pleistocene were fundamentally different than during the Holocene (11), and these differences had far-reaching implications for community structure, evolution, and extinction causes (12).A recent survey comparing the extinction dates of circumboreal megafauna with ice-age climate suggests that extinctions and genetic turnover were most frequent during warm, interstadial events (13). However, the mechanisms for these extinctions remain unclear, partly because this previous study considered multiple taxa living in many different ecosystems. Here, we focus on five megafaunal species that coinhabited a region of the Arctic with an ecological setting that is relatively well-understood. To avoid the methodological problems involved in pinpointing extinction dates (13), we infer population dynamics from changes in the relative abundance of megafauna over time. Using a uniquely large dataset of dated megafaunal bones from one particular area, we test a specific paleoecological hypothesis relating rapid climate change to population dynamics—namely, that transitions from cold to warm intervals were briefly optimal for grazing megafauna.The study area is Alaska’s North Slope, the tundra region bordered to the south by the Brooks Range and to the north by the Arctic Ocean (Fig. 1). The North Slope is a particularly interesting place to study end-Pleistocene extinctions for several reasons. First, its ice-age megafauna included iconic species like woolly mammoth (Mammuthus primigenius), steppe bison (Bison priscus), and cave lion (Panthera spelaea) (14). Second, the local extinctions of megafauna on Alaska’s North Slope occurred at a time when archaeological remains are rare, suggesting that people seldom ventured there (15, 16). Third, bone preservation in arctic environments tends to be excellent because of the presence of permafrost (perennially frozen ground), which makes it possible to ¹⁴C date large numbers of bones from many different species (SI Appendix, Table S1). Our record of dated bones provides key insights into the temporal dynamics and biogeographical characteristics of the mammoth steppe, a biome that was unique to the ice ages and the exact nature of which has been long debated (17).Open in a separate windowFig. 1.The North Slope is the tundra region between the Brooks Range and the Arctic Ocean. The light blue area shows the extent of the Bering Land Bridge during the last glacial maximum (LGM) ca. 19,000 calendar y B.P. Glacier extent (gray) during the LGM is based on the works by Dyke (64) and Brigham-Grette et al. (71). The timing of the opening of the ice-free corridor is still uncertain.     

Timing and dynamics of Late Pleistocene mammal extinctions in southwestern Australia

Explaining the Late Pleistocene demise of many of the world's larger terrestrial vertebrates is arguably the most enduring and debated topic in Quaternary science. Australia lost >90% of its larger species by around 40 thousand years (ka) ago, but the relative importance of human impacts and increased aridity remains unclear. Resolving the debate has been hampered by a lack of sites spanning the last glacial cycle. Here we report on an exceptional faunal succession from Tight Entrance Cave, southwestern Australia, which shows persistence of a diverse mammal community for at least 100 ka leading up to the earliest regional evidence of humans at 49 ka. Within 10 millennia, all larger mammals except the gray kangaroo and thylacine are lost from the regional record. Stable-isotope, charcoal, and small-mammal records reveal evidence of environmental change from 70 ka, but the extinctions occurred well in advance of the most extreme climatic phase. We conclude that the arrival of humans was probably decisive in the southwestern Australian extinctions, but that changes in climate and fire activity may have played facilitating roles. One-factor explanations for the Pleistocene extinctions in Australia are likely oversimplistic.     

A northern glacial refugium for bank voles (Clethrionomys glareolus)

There is controversy and uncertainty on how far north there were glacial refugia for temperate species during the Pleistocene glaciations and in the extent of the contribution of such refugia to present-day populations. We examined these issues using phylogeographic analysis of a European woodland mammal, the bank vole (Clethrionomys glareolus). A Bayesian coalescence analysis indicates that a bank vole population survived the height of the last glaciation (approximately 25,000-10,000 years B.P.) in the vicinity of the Carpathians, a major central European mountain chain well north of the Mediterranean areas typically regarded as glacial refugia for temperate species. Parameter estimates from the fitted isolation with migration model show that the divergence of the Carpathian population started at least 22,000 years ago, and it was likely followed by only negligible immigration from adjacent regions, suggesting the persistence of bank voles in the Carpathians through the height of the last glaciation. On the contrary, there is clear evidence for gene flow out of the Carpathians, demonstrating the contribution of the Carpathian population to the colonization of Europe after the Pleistocene. These findings are consistent with data from animal and plant fossils recovered in the Carpathians and provide the clearest phylogeographic evidence to date of a northern glacial refugium for temperate species in Europe.     

Probable extirpation of a breeding colony of Short-tailed Albatross (Phoebastria albatrus) on Bermuda by Pleistocene sea-level rise

Albatrosses (Diomedeidae) do not occur in the North Atlantic Ocean today except as vagrants, although five species were present in the early Pliocene. No fossil breeding sites of albatrosses were known previously. The timing of extinction of albatrosses in the North Atlantic was likewise unknown. Deposits that formed near present-day sea level along the southeastern shore of Bermuda contain remains of a former breeding colony and include intact eggshells and bones of embryos, juveniles, and adults of Short-tailed Albatross (Phoebastria albatrus), a critically endangered species now confined to a few islets in the northwestern Pacific Ocean. These deposits are correlated with the middle Pleistocene Lower Town Hill Formation, which at other sites have a radiometric age of 405,000 years ago. This equates with the marine isotope stage 11 interglacial, which culminated in a rise in sea-level to >+20 m. Bones of a juvenile Short-tailed Albatross were also found in beach deposits at +21.3 m from this same interglacial. We interpret the extirpation of albatrosses on Bermuda as probably resulting from lack of nesting sites protected from storm surges over the little emergent land that remained at the height of the marine isotope stage 11 sea level rise.     

Late Pleistocene and Holocene environmental history of the Iguala Valley, Central Balsas Watershed of Mexico

The origin of agriculture was a signal development in human affairs and as such has occupied the attention of scholars from the natural and social sciences for well over a century. Historical studies of climate and vegetation are closely associated with crop plant evolution because they can reveal the ecological contexts of plant domestication together with the antiquity and effects of agricultural practices on the environment. In this article, we present paleoecological evidence from three lakes and a swamp located in the Central Balsas watershed of tropical southwestern Mexico that date from 14,000 B.P. to the modern era. [Dates expressed in B.P. years are radiocarbon ages. Calibrated (calendar) ages, expressed as cal B.P., are provided for dates in the text.] Previous molecular studies suggest that maize (Zea mays L.) and other important crops such as squashes (Cucurbita spp.) were domesticated in the region. Our combined pollen, phytolith, charcoal, and sedimentary studies indicate that during the late glacial period (14,000-10,000 B.P.), lake beds were dry, the climate was cooler and drier, and open vegetational communities were more widespread than after the Pleistocene ended. Zea was a continuous part of the vegetation since at least the terminal Pleistocene. During the Holocene, lakes became important foci of human activity, and cultural interference with a species-diverse tropical forest is indicated. Maize and squash were grown at lake edges starting between 10,000 and 5,000 B.P., most likely sometime during the first half of that period. Significant episodes of climatic drying evidenced between 1,800 B.P. and 900 B.P. appear to be coeval with those documented in the Classic Maya region and elsewhere, showing widespread instability in the late Holocene climate.